Species coexistence by wide constant size spacing.

We consider the distribution of fruit pigeons of the genera Ptilinopus and Ducula on the island of New Guinea. Of the 21 species, between six and eight coexist inside humid lowland forests. We conducted or analyzed 31 surveys at 16 different sites, resurveying some sites in different years. The species coexisting at any single site in a single year are a highly nonrandom selection of the species to which that site is geographically accessible. Their sizes are both much more widely spread and more uniformly spaced than in random sets of species drawn from the locally available species pool. We also present a detailed case study of a highly mobile species that has been recorded on every ornithologically explored island in the West Papuan island group west of New Guinea. That species' rareness on just three well-surveyed islands within the group cannot be due to an inability to reach them. Instead, its local status decreases from abundant resident to rare vagrant in parallel with increasing weight proximity of the other resident species.

Region-wide retreats from lower elevations of range-restricted birds across the Northern Andes.

Local studies show upslope shifts in the distribution of tropical birds in response to warming temperatures. Unanswered is whether these upward shifts occur regionally across many species. We considered a nearly 2000-km length of the Northern Andes, where deforestation, temperature, and extreme weather events have increased during the past decades. Range-restricted bird species are particularly vulnerable to such events and occur in exceptionally high numbers in this region. Using abundant crowd-sourced data from the Cornell Lab of Ornithology database, eBird, and the Global Biodiversity Information Facility, we documented distributions of nearly 200 such species. We examined whether species shifted their elevational ranges over time by comparing observed versus expected occurrences below a low elevational threshold and above a high elevational threshold for 2 periods: before and after 2005. We predicted fewer observations at lower elevations (those below the threshold) and more at upper elevations (those above the threshold) after 2005. We also tested for deforestation effects at lower elevations within each species' distribution ranges. We compared relative forest loss with the differences between observed and expected occurrences across the elevational range. Species' retreats from lower elevations were ubiquitous and involved a 23-40% decline in prevalence at the lowest elevations. Increases at higher elevations were not consistent. The retreats occurred across a broad spectrum of species, from predominantly lowland to predominantly highland. Because deforestation showed no relationship with species retreats, we contend that a warming climate is the most parsimonious explanation for such shifts.

Repliegues regionales desde elevaciones más bajas de aves de distribución restringida en los Andes septentrionales Resumen Los estudios locales muestran cambios en la distribución altitudinal de las aves tropicales como respuesta al aumento de la temperatura. No sabemos si estos cambios suceden en muchas especies a nivel regional. Consideramos casi 2000 km de los Andes septentrionales, en donde la deforestación y los eventos climáticos extremos han incrementado en las últimas décadas. Las aves con distribución restringida son particularmente vulnerables a dichos eventos y su presencia es numerosa en esta región. Usamos datos abundantes de origen colectivo tomados de la base de datos del Laboratorio de Ornitología de Cornell, eBird y el Sistema Global de Información sobre Biodiversidad para documentar la distribución de aproximadamente 200 de estas especies. Analizamos si las especies cambiaron su distribución altitudinal con el tiempo al comparar entre la presencia observada y la esperada bajo un umbral de elevación reducida y por encima de un umbral de elevación alta durante dos periodos: antes y después de 2005. Pronosticamos una cantidad menor de observaciones por debajo del umbral y una mayor cantidad por encima del umbral para después de 2005. También analizamos los efectos de la deforestación en elevaciones más bajas dentro de los rangos de distribución de las especies y comparamos la pérdida relativa del bosque con las diferencias entre la presencia observada y la esperada en todo el rango altitudinal. El repliegue de las especies a partir de las elevaciones más bajas fue ubicuo e involucró una declinación del 23-40% de la prevalencia en las elevaciones más bajas. Los incrementos en las elevaciones más altas no fueron uniformes. Los repliegues ocurrieron a lo largo de un espectro amplio de especies, desde las que predominan en las tierras bajas hasta las que predominan en las tierras altas. Ya que la deforestación no se relacionó con el repliegue, sostenemos que un clima más cálido es la explicación más parsimoniosa para estos cambios.

Using metapopulation theory for practical conservation of mangrove endemic birds.

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.

Uso de la Teoría de Metapoblaciones para la Conservación Práctica de las Aves Endémicas de Manglares Resumen A medida que un paisaje se fragmenta cada vez más debido a la pérdida de hábitat, los parches se vuelven más pequeños y aislados y, por lo tanto, menos propensos a sostener a una población local. La teoría de metapoblaciones es adecuada para analizar paisajes fragmentados porque combina características empíricas del paisaje con información de cada especie para producir información directa sobre los riesgos de extinción de la población. Este enfoque contrasta con las descripciones de los fragmentos de hábitat que solo proporcionan información directa sobre el riesgo. Mediante la combinación de un modelo metapoblacional espacialmente explícito con datos empíricos de los rangos de distribución de especies endémicas y mapas de la cobertura del hábitat, calculamos la capacidad de la metapoblación - una medida de la capacidad del paisaje para sostener una metapoblación. Los manglares proporcionan un paisaje modelo ideal porque son de interés para la conservación y los límites de los parches son delineados fácilmente. Calculamos la capacidad de la metapoblación global para el período 2000-2015 de 99 metapoblaciones de 32 especies de aves endémicas de manglares. El norte de Australia y el sudeste de Asia tuvieron la mayor riqueza de aves endémicas de manglares. El Caribe, la costa del Pacífico de Centroamérica, Madagascar, Borneo y parches aislados en el sudeste de Asia en Myanmar y Malasia tuvieron las mayores pérdidas de metapoblaciones. Las regiones con mayor pérdida hábitat fueron necesariamente aquellas con mayor pérdida de capacidad de la metapoblación. A menudo no era una cuestión de cuánto, sino cómo se perdió el hábitat. Nuestro método se puede utilizar por manejadores para evaluar y priorizar un paisaje para la persistencia de la metapoblación.

Bird extirpations and community dynamics in an Andean cloud forest over 100 years of land-use change.

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.

Targeted habitat restoration can reduce extinction rates in fragmented forests.

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.

US protected lands mismatch biodiversity priorities.

Because habitat loss is the main cause of extinction, where and how much society chooses to protect is vital for saving species. The United States is well positioned economically and politically to pursue habitat conservation should it be a societal goal. We assessed the US protected area portfolio with respect to biodiversity in the country. New synthesis maps for terrestrial vertebrates, freshwater fish, and trees permit comparison with protected areas to identify priorities for future conservation investment. Although the total area protected is substantial, its geographic configuration is nearly the opposite of patterns of endemism within the country. Most protected lands are in the West, whereas the vulnerable species are largely in the Southeast. Private land protections are significant, but they are not concentrated where the priorities are. To adequately protect the nation's unique biodiversity, we recommend specific areas deserving additional protection, some of them including public lands, but many others requiring private investment.

Navigating the complexity of ecological stability.

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.

China's endemic vertebrates sheltering under the protective umbrella of the giant panda.

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 <20,000 km(2) after accounting for elevational restriction and remaining habitats. These species concentrate mainly in Sichuan, Yunnan, Nan Mountains, and Hainan. There is a high concentration in the east Daxiang and Xiaoxiang Mountains of Sichuan, where pandas are absent and where there are no national nature reserves. The others concentrate in Yunnan, Nan Mountains, and Hainan. Here, 10 prefectures might establish new protected areas or upgrade local nature reserves to national status.

Global patterns of terrestrial vertebrate diversity and conservation.

Identifying priority areas for biodiversity is essential for directing conservation resources. Fundamentally, we must know where individual species live, which ones are vulnerable, where human actions threaten them, and their levels of protection. As conservation knowledge and threats change, we must reevaluate priorities. We mapped priority areas for vertebrates using newly updated data on >21,000 species of mammals, amphibians, and birds. For each taxon, we identified centers of richness for all species, small-ranged species, and threatened species listed with the International Union for the Conservation of Nature. Importantly, all analyses were at a spatial grain of 10 × 10 km, 100 times finer than previous assessments. This fine scale is a significant methodological improvement, because it brings mapping to scales comparable with regional decisions on where to place protected areas. We also mapped recent species discoveries, because they suggest where as-yet-unknown species might be living. To assess the protection of the priority areas, we calculated the percentage of priority areas within protected areas using the latest data from the World Database of Protected Areas, providing a snapshot of how well the planet's protected area system encompasses vertebrate biodiversity. Although the priority areas do have more protection than the global average, the level of protection still is insufficient given the importance of these areas for preventing vertebrate extinctions. We also found substantial differences between our identified vertebrate priorities and the leading map of global conservation priorities, the biodiversity hotspots. Our findings suggest a need to reassess the global allocation of conservation resources to reflect today's improved knowledge of biodiversity and conservation.

Estimating the normal background rate of species extinction.

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.

Setting practical conservation priorities for birds in the Western Andes of Colombia.

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.

Biodiversity hotspots house most undiscovered plant species.

For most organisms, the number of described species considerably underestimates how many exist. This is itself a problem and causes secondary complications given present high rates of species extinction. Known numbers of flowering plants form the basis of biodiversity "hotspots"--places where high levels of endemism and habitat loss coincide to produce high extinction rates. How different would conservation priorities be if the catalog were complete? Approximately 15% more species of flowering plant are likely still undiscovered. They are almost certainly rare, and depending on where they live, suffer high risks of extinction from habitat loss and global climate disruption. By using a model that incorporates taxonomic effort over time, regions predicted to contain large numbers of undiscovered species are already conservation priorities. Our results leave global conservation priorities more or less intact, but suggest considerably higher levels of species imperilment than previously acknowledged.

Tackling debt, biodiversity loss, and climate change.

Experience tells us how to maximize debt-for-nature effectiveness.

Conservation gaps and priorities of range-restricted birds in the Northern Andes.

The ongoing destruction of habitats in the tropics accelerates the current rate of species extinction. Range-restricted species are exceptionally vulnerable, yet we have insufficient knowledge about their protection. Species' current distributions, range sizes, and protection gaps are crucial to determining conservation priorities. Here, we identified priority range-restricted bird species and their conservation hotspots in the Northern Andes. We employed maps of the Area of Habitat (AOH), that better reflect their current distributions than existing maps. AOH provides unprecedented resolution and maps a species in the detail essential for practical conservation actions. We estimated protection within each species' AOH and for the cumulative distribution of all 335 forest-dependent range-restricted birds across the Northern Andes. For the latter, we also calculated protection across the elevational gradient. We estimated how much additional protection community lands (Indigenous and Afro-Latin American lands) would contribute if they were conservation-focused. AOHs ranged from 8 to 141,000 km2. We identified four conservation priorities based on cumulative species richness: the number of AOHs stacked per unit area. These priorities are high-resolution mapped representations of Endemic Bird Areas for the Tropical Andes that we consider critically important. Protected areas cover only 31% of the cumulative AOH, but community lands could add 19% more protection. Sixty-two per cent of the 335 species have ranges smaller than their published estimates, yet IUCN designates only 23% of these as Threatened. We identified 50 species as top conservation priorities. Most of these concentrate in areas of low protection near community lands and at middle elevations where, on average, only 34% of the land is protected. We highlight the importance of collaborative efforts among stakeholders: governments should support private and community-based conservation practices to protect the region with the most range-restricted birds worldwide.

The synergy between protected area effectiveness and economic growth.

Protected areas conserve biodiversity and ecosystem functions but might impede local economic growth. Understanding the global patterns and predictors of different relationships between protected area effectiveness and neighboring community economic growth can inform better implementation of the Kunming-Montreal Global Biodiversity Framework. We assessed 10,143 protected areas globally with matched samples to address the non-random location of protected areas. Our results show that protected areas resist human-induced land cover changes and do not limit nightlight increases in neighboring settlements. This result is robust, using different matching techniques, parameter settings, and selection of covariates. We identify four types of relationships between land cover changes and nightlight changes for each protected area: "synergy," "retreat," and two tradeoff relationships. About half of the protected areas (47.5%) retain their natural land cover and do so despite an increase of nightlights in the neighboring communities. This synergy relationship is the most common globally but varies between biomes and continents. Synergy is less frequent in the Amazon, Southeast Asia, and some developing areas, where most biodiversity resides and which suffer more from poverty. Smaller protected areas and those with better access to cities, moderate road density, and better baseline economic conditions have a higher probability of reaching synergy. Our results are promising, as the expansion of protected areas and increased species protection will rely more on conserving the human-modified landscape with smaller protected areas. Future interventions should address local development and biodiversity conservation together to achieve more co-benefits.

Protecting and connecting landscapes stabilizes populations of the Endangered savannah elephant.

The influence of protected areas on the growth of African savannah elephant populations is inadequately known. Across southern Africa, elephant numbers grew at 0.16% annually for the past quarter century. Locally, much depends on metapopulation dynamics-the size and connections of individual populations. Population numbers in large, connected, and strictly protected areas typically increased, were less variable from year to year, and suffered less from poaching. Conversely, populations in buffer areas that are less protected but still connected have more variation in growth from year to year. Buffer areas also differed more in their growth rates, likely due to more threats and dispersal opportunities in the face of such dangers. Isolated populations showed consistently high growth due to a lack of emigration. This suggests that "fortress" conservation generally maintains high growth, while anthropogenic-driven source-sink dynamics within connected conservation clusters drive stability in core areas and variability in buffers.

Estimating extinction risk with metapopulation models of large-scale fragmentation.

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.