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The fields of ecology and conservation have evolved rapidly over the past century. Synthesizing larger trends in these disciplines remains a challenge yet is critical to bridging subdisciplines, guiding research, and informing educational frameworks. Here, we provide what we believe is the largest full‐text culturomic analysis of ecology and conservation journals, covering 80 years, 52 journals, and half a billion words. Our analysis illuminates the boom‐and‐bust of ecological hypotheses and theories; the adoption of statistical, genetic, and social‐science approaches; and the domination of terms that have emerged in recent decades (eg climate change, invasive species, ecosystem services, meta‐analysis, and supplementary material, which largely replaced unpublished data). We track the evolution of ecology from a largely descriptive field focused on natural history and observational studies to a more data‐driven, multidisciplinary field focused on applied environmental issues. Overall, our analysis highlights the increasing breadth of the field, illustrating that there is room for more diversity of ecologists and conservationists today than ever before.

Human activities alter ecosystems everywhere, causing rapid biodiversity loss and biotic homogenization. These losses necessitate coordinated conservation actions guided by biodiversity and species distribution spatial data that cover large areas yet have fine-enough resolution to be management-relevant (i.e., ≤5 km). However, most biodiversity products are too coarse for management or are only available for small areas. Furthermore, many maps generated for biodiversity assessment and conservation do not explicitly quantify the inherent tradeoff between resolution and accuracy when predicting biodiversity patterns. Our goals were to generate predictive models of overall breeding bird species richness and species richness of different guilds based on nine functional or life-history-based traits across the conterminous United States at three resolutions (0.5, 2.5, and 5 km) and quantify the tradeoff between resolution and accuracy and, hence, relevance for management of the resulting biodiversity maps. We summarized 18 years of North American Breeding Bird Survey data (1992-2019) and modeled species richness using random forests, including 66 predictor variables (describing climate, vegetation, geomorphology, and anthropogenic conditions), 20 of which we newly derived. Among the three spatial resolutions, the percentage variance explained ranged from 27% to 60% (median = 54%; mean = 57%) for overall species richness and 12% to 87% (median = 61%; mean = 58%) for our different guilds. Overall species richness and guild-specific species richness were best explained at 5-km resolution using ~24 predictor variables based on percentage variance explained, symmetric mean absolute percentage error, and root mean square error values. However, our 2.5-km-resolution maps were almost as accurate and provided more spatially detailed information, which is why we recommend them for most management applications. Our results represent the first consistent, occurrence-based, and nationwide maps of breeding bird richness with a thorough accuracy assessment that are also spatially detailed enough to inform local management decisions. More broadly, our findings highlight the importance of explicitly considering tradeoffs between resolution and accuracy to create management-relevant biodiversity products for large areas.

Previous estimates of tropical forest carbon loss in the twenty-first century using satellite data typically focus on its magnitude, whereas regional loss trajectories and associated drivers are rarely reported. Here we used different high-resolution satellite datasets to show a doubling of gross tropical forest carbon loss worldwide from 0.97 ± 0.16 PgC yr−1 in 2001–2005 to 1.99 ± 0.13 PgC yr−1 in 2015–2019. This increase in carbon loss from forest conversion is higher than in bookkeeping models forced by land-use statistical data, which show no trend or a slight decline in land-use emissions in the early twenty-first century. Most (82%) of the forest carbon loss is at some stages associated with large-scale commodity or small-scale agriculture activities, particularly in Africa and Southeast Asia. We find that ~70% of former forest lands converted to agriculture in 2001–2019 remained so in 2020, confirming a dominant role of agriculture in long-term pan-tropical carbon reductions on formerly forested landscapes. The acceleration and high rate of forest carbon loss in the twenty-first century suggest that existing strategies to reduce forest loss are not successful; and this failure underscores the importance of monitoring deforestation trends following the new pledges made in Glasgow.

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

Many global environmental agendas, including halting biodiversity loss, reversing land degradation, and limiting climate change, depend upon retaining forests with high ecological integrity, yet the scale and degree of forest modification remain poorly quantified and mapped. By integrating data on observed and inferred human pressures and an index of lost connectivity, we generate a globally consistent, continuous index of forest condition as determined by the degree of anthropogenic modification. Globally, only 17.4 million km

Many global environmental agendas, including halting biodiversity loss, reversing land degradation, and limiting climate change, depend upon retaining forests with high ecological integrity, yet the scale and degree of forest modification remain poorly quantified and mapped. By integrating data on observed and inferred human pressures and an index of lost connectivity, we generate a globally consistent, continuous index of forest condition as determined by the degree of anthropogenic modification. Globally, only 17.4 million km2 of forest (40.5%) has high landscape-level integrity (mostly found in Canada, Russia, the Amazon, Central Africa, and New Guinea) and only 27% of this area is found in nationally designated protected areas. Of the forest inside protected areas, only 56% has high landscape-level integrity. Ambitious policies that prioritize the retention of forest integrity, especially in the most intact areas, are now urgently needed alongside current efforts aimed at halting deforestation and restoring the integrity of forests globally., Mapping and quantifying degree of forest modification is critical to conserve and manage forests. Here the authors propose a new quantitative metric for landscape integrity and apply it to a global forest map, showing that less than half of the world’s forest cover has high integrity, most of which is outside nationally designed protected areas.

Species loss is occurring globally at unprecedented rates, and effective conservation planning requires an understanding of landscape characteristics that determine biodiversity patterns. Habitat heterogeneity is an important determinant of species diversity, but is difficult to measure across large areas using field-based methods that are costly and logistically challenging. Satellite image texture analysis offers a cost-effective alternative for quantifying habitat heterogeneity across broad spatial scales. We tested the ability of texture measures derived from 30-m resolution Enhanced Vegetation Index (EVI) data to capture habitat heterogeneity and predict bird species richness across the conterminous United States. We used Landsat 8 satellite imagery from 2013-2017 to derive a suite of texture measures characterizing vegetation heterogeneity. Individual texture measures explained up to 21% of the variance in bird richness patterns in North American Breeding Bird Survey (BBS) data during the same time period. Texture measures were positively related to total breeding bird richness, but this relationship varied among forest, grassland, and shrubland habitat specialists. Multiple texture measures combined with mean EVI explained up to 41% of the variance in total bird richness, and models including EVI-based texture measures explained up to 10% more variance than those that included only EVI. Models that also incorporated topographic and land cover metrics further improved predictive performance, explaining up to 51% of the variance in total bird richness. A texture measure contributed predictive power and characterized landscape features that EVI and forest cover alone could not, even though the latter two were overall more important variables. Our results highlight the potential of texture measures for mapping habitat heterogeneity and species richness patterns across broad spatial extents, especially when used in conjunction with vegetation indices or land cover data. By generating 30-m resolution texture maps and modeling bird richness at a near-continental scale, we expand on previous applications of image texture measures for modeling biodiversity that were either limited in spatial extent or based on coarse-resolution imagery. Incorporating texture measures into broad-scale biodiversity models may advance our understanding of mechanisms underlying species richness patterns and improve predictions of species responses to rapid global change.

Many global environmental agendas, including halting biodiversity loss, reversing land degradation, and limiting climate change, depend upon retaining forests with high ecological integrity, yet the scale and degree of forest modification remains poorly quantified and mapped. By integrating data on observed and inferred human pressures and an index of lost connectivity, we generate the first globally-consistent, continuous index of forest condition as determined by degree of anthropogenic modification. Globally, only 17.4 million km2 of forest (40.5%) have high landscape level integrity (mostly found in Canada, Russia, the Amazon, Central Africa and New Guinea) and only 27% of this area is found in nationally-designated protected areas. Of the forest in protected areas, only 56% has high landscape level integrity. Ambitious policies that prioritize the retention of forest integrity, especially in the most intact areas, are now urgently needed alongside current efforts aimed at halting deforestation and restoring the integrity of forests globally.

Climate change is leading to widespread elevational shifts thought to increase species extinction risk in mountains. We integrate digital elevation models with a metric of human pressure to examine changes in the amount of intact land area available for species undergoing elevational range shifts in all major mountain ranges globally (n = 1010). Nearly 60% of mountainous area is under intense human pressure, predominantly at low elevations and mountain bases. Consequently, upslope range shifts generally resulted in modeled species at lower elevations expanding into areas of lower human pressure and, due to complex topography, encountering more intact land area relative to their starting position. Such gains were often attenuated at high elevations as land-use constraints diminished and topographic constraints increased. Integrating patterns of topography and human pressure is essential for accurate species vulnerability assessments under climate change, as priorities for protecting, connecting, and restoring mountain landscapes may otherwise be misguided., It is often assumed that many species will move upslope in mountainous regions as the climate warms. However, the authors show here that as many species move to higher elevations they will enter areas of lower human footprint but potentially more constraining topography.

Protected areas (PAs) that span elevational gradients enhance protection for taxonomic and phylogenetic diversity and facilitate species range shifts under climate change. We quantified the global protection of elevational gradients by analyzing the elevational distributions of 44,155 PAs in 1,010 mountain ranges using the highest resolution digital elevation models available. We show that, on average, mountain ranges in Africa and Asia have the lowest elevational protection, ranges in Europe and South America have intermediate elevational protection, and ranges in North America and Oceania have the highest elevational protection. We use the Convention on Biological Diversity’s Aichi Target 11 to assess the proportion of elevational gradients meeting the 17% suggested minimum target and examine how different protection categories contribute to elevational protection. When considering only strict PAs [International Union for Conservation of Nature (IUCN) categories I–IV, n = 24,706], nearly 40% of ranges do not contain any PAs, roughly half fail to meet the 17% target at any elevation, and ∼75% fail to meet the target throughout ≥50% of the elevational gradient. Observed elevational protection is well below optimal, and frequently below a null model of elevational protection. Including less stringent PAs (IUCN categories V–VI and nondesignated PAs, n = 19,449) significantly enhances elevational protection for most continents, but several highly biodiverse ranges require new or expanded PAs to increase elevational protection. Ensuring conservation outcomes for PAs with lower IUCN designations as well as strategically placing PAs to better represent and connect elevational gradients will enhance ecological representation and facilitate species range shifts under climate change.

To understand ecological phenomena, it is necessary to observe their behaviour across multiple spatial and temporal scales. Since this need was first highlighted in the 1980s, technology has opened previously inaccessible scales to observation. To help to determine whether there have been corresponding changes in the scales observed by modern ecologists, we analysed the resolution, extent, interval and duration of observations (excluding experiments) in 348 studies that have been published between 2004 and 2014. We found that observational scales were generally narrow, because ecologists still primarily use conventional field techniques. In the spatial domain, most observations had resolutions ≤1 m2 and extents ≤10,000 ha. In the temporal domain, most observations were either unreplicated or infrequently repeated (>1 month interval) and ≤1 year in duration. Compared with studies conducted before 2004, observational durations and resolutions appear largely unchanged, but intervals have become finer and extents larger. We also found a large gulf between the scales at which phenomena are actually observed and the scales those observations ostensibly represent, raising concerns about observational comprehensiveness. Furthermore, most studies did not clearly report scale, suggesting that it remains a minor concern. Ecologists can better understand the scales represented by observations by incorporating autocorrelation measures, while journals can promote attentiveness to scale by implementing scale-reporting standards.

The many Gondwanic vegetation types found across the extensive latitudes and elevation gradients of South America's southern cone contribute to Chile's global biodiversity hotspot ranking. Species loss in global biodiversity hotspots is an ongoing climate change concern and land managers need spatially explicit climate risk maps to adapt conservation strategies to climate change in these areas. We modeled future climate risk for Chile's terrestrial vegetation using a high-resolution vegetation map and tested the relationship to climate risk for each type's latitudinal and elevation range. We found that 43.6% of all vegetation has high climate risk in Global Circulation Models (GCMs) under a high emissions scenario (RCP8.5). All forest types in the country, including Southern Beech (Nothofagus sp.), Alerce (Fitzroya cupressoides), Araucaria (Araucaria araucana), and Sclerophyllous, as well as the Valdivian rainforest, Altiplanic Steppes, and Salares, face high levels of climate risk. Tests for trends in risk across elevation and latitude showed that exposure for all types increased with elevation based on the MIROC5 GCM, and decreased with latitude based on the Had2GEM-ES GCM. Our results suggest that vegetation types with smaller latitudinal ranges typically have higher levels of climate risk, but a type's elevation range is not significantly correlated with risk of exposure. We identified climatically stable areas which could act as vegetation refugia in Patagonia, the central Andes mountains between latitudes 27.5°S and 32.5°S, and some coastal areas. Conservation strategies in Chile should include the protection of climatically stable areas to safeguard current Gondwanic biodiversity and active habitat restoration in climatically exposed areas to facilitate vegetation shifts.

Over the course of a year, vegetation and temperature have strong phenological and seasonal patterns, respectively, and many species have adapted to these patterns. High inter-annual variability in the phenology of vegetation and in the seasonality of temperature pose a threat for biodiversity. However, areas with high spatial variability likely have higher ecological resilience where inter-annual variability is high, because spatial variability indicates presence of a range of resources, microclimatic refugia, and habitat conditions. The integration of inter-annual and spatial variability is thus important for biodiversity conservation. Areas where spatial variability is low and inter-annual variability is high are likely to limit resilience to disturbance. In contrast, areas of high spatial variability may be high priority candidates for protection. Our goal was to develop spatio-temporal remotely sensed indices to identify hotspots of biodiversity conservation concern. We generated indices that capture the inter-annual and spatial variability of vegetation greenness and land surface temperature and integrated them to identify areas of high, medium, and low biodiversity conservation concern. We applied our method in Argentina (2.8 million km2), a country with a wide range of climates and biomes. To generate the inter-annual variability indices, we analyzed MODIS Enhanced Vegetation Index (EVI) and Land Surface Temperature (LST) time series from 2001 to 2018, fitted curves to obtain annual phenological and seasonal metrics, and calculated their inter-annual variability. To generate the spatial variability indices, we calculated standard deviation image texture of Landsat 8 EVI and LST. When we integrated our inter-annual and spatial variability indices, areas in the northeast and parts of southern Argentina were the hotspots of highest conservation concern. High inter-annual variability poses a threat in these areas, because spatial variability is low. These are areas where management efforts could be valuable. In contrast, areas in the northwest and central-west are where protection should be strongly considered because the high spatial variability may confer resilience to disturbance, due to the variety of conditions and resources within close proximity. We developed remotely sensed indices to identify hotspots of high and low conservation concern at scales relevant to biodiversity conservation, which can be used to target management actions in order to minimize biodiversity loss.

There is clear evidence that species' ranges along environmental gradients are constrained by both biotic and abiotic factors, yet their relative importance in structuring realized distributions remains uncertain. We surveyed breeding bird communities while collecting in situ temperature and vegetation data along five elevational transects in the Himalayas differing in temperature variability, habitat zonation, and bird richness in order to disentangle temperature, habitat, and congeneric competition as mechanisms structuring elevational ranges. Our results from species' abundance models representing these three mechanisms differed markedly from previous, foundational research in the tropics. Contrary to general expectations, we found little evidence for competition as a major determinant of range boundaries, with congeneric species limiting only 12% of ranges. Instead, temperature and habitat were found to structure the majority of species' distributions, limiting 48 and 40% of ranges, respectively. Our results suggest that different mechanisms may structure species ranges in the temperate Himalayas compared to tropical systems. Despite recent evidence suggesting temperate species have broader thermal tolerances than tropical species, our findings reinforce the notion that the abiotic environment has significant control over the distributions of temperate species.

Climatic representation in protected areas—and not just the amount—is critical to safeguarding biodiversity under climate change., Protected areas (PAs) are essential to biodiversity conservation, but their static boundaries may undermine their potential for protecting species under climate change. We assessed how the climatic conditions within global terrestrial PAs may change over time. By 2070, protection is expected to decline in cold and warm climates and increase in cool and hot climates over a wide range of precipitation. Most countries are expected to fail to protect >90% of their available climate at current levels. The evenness of climatic representation under protection—not the amount of area protected—positively influenced the retention of climatic conditions under protection. On average, protection retention would increase by ~118% if countries doubled their climatic representativeness under protection or by ~102% if countries collectively reduced emissions in accordance with global targets. Therefore, alongside adoption of mitigation policies, adaptation policies that improve the complementarity of climatic conditions within PAs will help countries safeguard biodiversity.

The thermal environment limits species ranges through its influence on physiology and resource distributions and thus affects species richness patterns over broad spatial scales. Understanding how temperature drives species richness patterns is particularly important in the context of global change and for effective conservation planning. Landsat 8's Thermal Infrared Sensor (TIRS) allows direct mapping of temperature at moderate spatial resolutions (100 m, downscaled by the USGS to 30 m), overcoming limitations inherent in coarse interpolated weather station data that poorly capture fine-scale temperature patterns over broad areas. TIRS data thus offer the unique opportunity to understand how the thermal environment influences species richness patterns. Our aim was to develop and assess the ability of TIRS-based temperature metrics to predict patterns of winter bird richness across the conterminous United States during winter, a period of marked temperature stress for birds. We used TIRS data from 2013-2018 to derive metrics of relative temperature and intra-seasonal thermal heterogeneity. To quantify winter bird richness across the conterminous US, we tabulated the richness only for resident bird species, i.e., those species that do not move between the winter and breeding seasons, from the North American Breeding Bird Survey, the most extensive survey of birds in the US. We expected that relative temperature and thermal heterogeneity would have strong positive associations with winter bird richness because colder temperatures heighten temperature stress for birds, and thermal heterogeneity is a proxy for thermal niches and potential thermal refugia that can support more species. We further expected that both the strength of the effects and the relative importance of these variables would be greater for species with greater climate sensitivity, such as small-bodied species and climate-threatened species (i.e., those with large discrepancies between their current and future distributions following projected climate change). Consistent with our predictions, relative temperature and thermal heterogeneity strongly positively influenced winter bird richness patterns, with statistical models explaining 37.3% of the variance in resident bird richness. Thermal heterogeneity was the strongest predictor of small-bodied and climate-threatened species in our models, whereas relative temperature was the strongest predictor of large-bodied and climate-stable species. Our results demonstrate the important role that the thermal environment plays in governing winter bird richness patterns and highlight the previously underappreciated role that intra-seasonal thermal heterogeneity may have in supporting high winter bird species richness. Our findings thus illustrate the exciting potential for TIRS data to guide conservation planning in an era of global change.

In their Letter, You et al. (1) raise concerns about the use of the World Database on Protected Areas (WDPA) (https://www.protectedplanet.net/) in conservation assessments and planning. Their concern arises from potential differences in protected area (PA) delineations and designations between the WDPA and alternate, national PA datasets, citing China’s National Nature Reserves (CNNR) as an example with apparent deviations. You et al. (1) highlight that using the CNNR in calculations of elevational protection yields results that contrast with those published in our original paper (2) using the WDPA. Several issues with You et al.’s (1) analysis prevent a straightforward comparison with … [↵][1]1To whom correspondence should be addressed. Email: pelsen{at}berkeley.edu. [1]: #xref-corresp-1-1

The Himalayas are a global biodiversity hotspot threatened by widespread agriculture and pasture expansion. To determine the impact of these threats on biodiversity and to formulate appropriate conservation strategies, we surveyed birds along elevational gradients in primary forest and in human-dominated lands spanning a gradient of habitat alteration, including forest-agriculture mosaics, mixed agriculture mosaics, and pasture. We surveyed birds during the breeding season and in winter to account for pronounced seasonal migrations. Bird abundance and richness in forest-agriculture and mixed agriculture mosaics were equal to or greater than in primary forest and greater than in pasture at local and landscape scales during both seasons. Pasture had greater abundance and richness of birds in winter than primary forest, but richness was greater in primary forest at the landscape scale during the breeding season. All 4 land-use types held unique species, suggesting that all must be retained in the landscape to conserve the entire avifauna. Our results suggest forest-agriculture and mixed agriculture mosaics are particularly important for sustaining Himalayan bird communities during winter and primary forests are vital for sustaining Himalayan bird communities during the breeding season. Further conversion of forest-agriculture and mixed agriculture mosaics to pasture would likely result in significant biodiversity losses that would disproportionately affect breeding species. To ensure comprehensive conservation, strategies in the western Himalayas must balance the protection of intact primary forest with the minimization of pasture expansion.

Surface area does not decrease monotonically with elevation for two-thirds of mountain ranges. Consequently many mountain species might not experience reduced habitat area as they move upslope under climate change.

The impacts of land-use change on biodiversity in the Himalayas are poorly known, notwithstanding widespread deforestation and agricultural intensification in this highly biodiverse region. Although intact primary forests harbor many Himalayan birds during breeding, a large number of bird species use agricultural lands during winter. We assessed how Himalayan bird species richness, abundance, and composition during winter are affected by forest loss stemming from agriculture and grazing. Bird surveys along 12 elevational transects within primary forest, low-intensity agriculture, mixed subsistence agriculture, and intensively grazed pastures in winter revealed that bird species richness and abundance were greatest in low-intensity and mixed agriculture, intermediate in grazed pastures, and lowest in primary forest at both local and landscape scales; over twice as many species and individuals were recorded in low-intensity agriculture than in primary forest. Bird communities in primary forests were distinct from those in all other land-use classes, but only 4 species were unique to primary forests. Low-, medium-, and high-intensity agriculture harbored 32 unique species. Of the species observed in primary forest, 80% had equal or greater abundance in low-intensity agricultural lands, underscoring the value of these lands in retaining diverse community assemblages at high densities in winter. Among disturbed landscapes, bird species richness and abundance declined as land-use intensity increased, especially in high-intensity pastures. Our results suggest that agricultural landscapes are important for most Himalayan bird species in winter. But agricultural intensification-especially increased grazing-will likely result in biodiversity losses. Given that forest reserves alone may inadequately conserve Himalayan birds in winter, comprehensive conservation strategies in the region must go beyond protecting intact primary forests and ensure that low-intensity agricultural lands are not extensively converted to high-intensity pastures.

Longstanding theory predicts that competitive interactions set species' range limits in relatively aseasonal, species-rich regions, while temperature limits distributions in more seasonal, species-poor areas. More recent theory holds that species evolve narrow physiological tolerances in aseasonal regions, with temperature being an important determining factor in such zones. We tested how abiotic (temperature) and biotic (competition) factors set range limits and structure bird communities along strong, opposing, temperature-seasonality and species-richness gradients in the Himalayas, in two regions separated by 1500 km. By examining the degree to which seasonal elevational migration conserves year-round thermal niches across species, we show that species in the relatively aseasonal and speciose east are more constrained by temperature compared with species in the highly seasonal west. We further show that seasonality has a profound effect on the strength of competition between congeneric species. Competition appears to be stronger in winter, a period of resource scarcity in the Himalayas, in both the east and the west, with similarly sized eastern species more likely to segregate in thermal niche space in winter. Our results indicate that rather than acting in isolation, abiotic and biotic factors mediate each other to structure ecological communities.

Incentivizing carbon storage can be a win-win pathway to conserving biodiversity and mitigating climate change. In savannas, however, the situation is more complex. Promoting carbon storage through woody encroachment may reduce plant diversity of savanna endemics, even as the diversity of encroaching forest species increases. This trade-off has important implications for the management of biodiversity and carbon in savanna habitats, but has rarely been evaluated empirically. We quantified the nature of carbon-diversity relationships in the Brazilian Cerrado by analyzing how woody plant species richness changed with carbon storage in 206 sites across the 2.2 million km(2) region at two spatial scales. We show that total woody plant species diversity increases with carbon storage, as expected, but that the richness of endemic savanna woody plant species declines with carbon storage both at the local scale, as woody biomass accumulates within plots, and at the landscape scale, as forest replaces savanna. The sharpest trade-offs between carbon storage and savanna diversity occurred at the early stages of carbon accumulation at the local scale but the final stages of forest encroachment at the landscape scale. Furthermore, the loss of savanna species quickens in the final stages of forest encroachment, and beyond a point, savanna species losses outpace forest species gains with increasing carbon accumulation. Our results suggest that although woody encroachment in savanna ecosystems may provide substantial carbon benefits, it comes at the rapidly accruing cost of woody plant species adapted to the open savanna environment. Moreover, the dependence of carbon-diversity trade-offs on the amount of savanna area remaining requires land managers to carefully consider local conditions. Widespread woody encroachment in both Australian and African savannas and grasslands may present similar threats to biodiversity.

Protected areas (PAs) have long been criticized as creations of and for an elite few, where associated costs, but few benefits, are borne by marginalized rural communities. Contrary to predictions of this argument, we found that average human population growth rates on the borders of 306 PAs in 45 countries in Africa and Latin America were nearly double average rural growth, suggesting that PAs attract, rather than repel, human settlement. Higher population growth on PA edges is evident across ecoregions, countries, and continents and is correlated positively with international donor investment in national conservation programs and an index of park-related funding. These findings provide insight on the value of PAs for local people, but also highlight a looming threat to PA effectiveness and biodiversity conservation.