Climate Change, Elevational Range Shifts, and Bird Extinctions

Abstract:  Limitations imposed on species ranges by the climatic, ecological, and physiological effects of elevation are important determinants of extinction risk. We modeled the effects of elevational limits on the extinction risk of landbirds, 87% of all bird species. Elevational limitation of range size explained 97% of the variation in the probability of being in a World Conservation Union category of extinction risk. Our model that combined elevational ranges, four Millennium Assessment habitat-loss scenarios, and an intermediate estimate of surface warming of 2.8° C, projected a best guess of 400–550 landbird extinctions, and that approximately 2150 additional species would be at risk of extinction by 2100. For Western Hemisphere landbirds, intermediate extinction estimates based on climate-induced changes in actual distributions ranged from 1.3% (1.1° C warming) to 30.0% (6.4° C warming) of these species. Worldwide, every degree of warming projected a nonlinear increase in bird extinctions of about 100–500 species. Only 21% of the species predicted to become extinct in our scenarios are currently considered threatened with extinction. Different habitat-loss and surface-warming scenarios predicted substantially different futures for landbird species. To improve the precision of climate-induced extinction estimates, there is an urgent need for high-resolution measurements of shifts in the elevational ranges of species. Given the accelerating influence of climate change on species distributions and conservation, using elevational limits in a tested, standardized, and robust manner can improve conservation assessments of terrestrial species and will help identify species that are most vulnerable to global climate change. Our climate-induced extinction estimates are broadly similar to those of bird species at risk from other factors, but these estimates largely involve different sets of species.     

Scenarios of change in the realized climatic niche of mountain carnivores and ungulates

Mountains are among the natural systems most affected by climate change, and mountain mammals are considered particularly imperiled, given their high degree of specialization to narrow tolerance bands of environmental conditions. Climate change mitigation policies, such as the Paris Agreement, are essential to stem climate change impacts on natural systems. But how significant is the Paris Agreement to the survival of mountain mammals? We investigated how alternative emission scenarios may determine change in the realized climatic niche of mountain carnivores and ungulates in 2050. We based our predictions of future change in species niches based on how species have responded to past environmental changes, focusing on the probabilities of niche shrink and niche stability. We found that achieving the Paris Agreement's commitments would substantially reduce climate instability for mountain species. Specifically, limiting global warming to below 1.5°C would reduce the probability of niche shrinkage by 4% compared with a high-emission scenario. Globally, carnivores showed greater niche shrinkage than ungulates, whereas ungulates were more likely to shift their niches (i.e., face a level of climate change that allows adaptation). Twenty-three species threatened by climate change according to the IUCN Red List had greater niche contraction than other species we analyzed (3% higher on average). We therefore argue that climate mitigation policies must be coupled with rapid species-specific conservation intervention and sustainable land-use policies to avoid high risk of loss of already vulnerable species.     

Scenarios of large mammal loss in Europe for the 21st century

Distributions and populations of large mammals are declining globally, leading to an increase in their extinction risk. We forecasted the distribution of extant European large mammals (17 carnivores and 10 ungulates) based on 2 Rio+20 scenarios of socioeconomic development: business as usual and reduced impact through changes in human consumption of natural resources. These scenarios are linked to scenarios of land‐use change and climate change through the spatial allocation of land conversion up to 2050. We used a hierarchical framework to forecast the extent and distribution of mammal habitat based on species’ habitat preferences (as described in the International Union for Conservation of Nature Red List database) within a suitable climatic space fitted to the species’ current geographic range. We analyzed the geographic and taxonomic variation of habitat loss for large mammals and the potential effect of the reduced impact policy on loss mitigation. Averaging across scenarios, European large mammals were predicted to lose 10% of their habitat by 2050 (25% in the worst‐case scenario). Predicted loss was much higher for species in northwestern Europe, where habitat is expected to be lost due to climate and land‐use change. Change in human consumption patterns was predicted to substantially improve the conservation of habitat for European large mammals, but not enough to reduce extinction risk if species cannot adapt locally to climate change or disperse.     

Effects of climate and land-cover change on the conservation status of gibbons

Species shift their distribution in response to climate and land-cover change, which may result in a spatial mismatch between currently protected areas (PAs) and priority conservation areas (PCAs). We examined the effects of climate and land-cover change on potential range of gibbons and sought to identify PCAs that would conserve them effectively. We collected global gibbon occurrence points and modeled (ecological niche model) their current and potential 2050s ranges under climate-change and different land-cover-change scenarios. We examined change in range and PA coverage between the current and future ranges of each gibbon species. We applied spatial conservation prioritization to identify the top 30% PCAs for each species. We then determined how much of the PCAs are conserved in each country within the global range of gibbons. On average, 31% (SD 22) of each species’ current range was covered in PAs. PA coverage of the current range of 9 species was <30%. Nine species lost on average 46% (SD 29) of their potential range due to climate change. Under climate-change with an optimistic land-cover-change scenario (B1), 12 species lost 39% (SD 28) of their range. In a pessimistic land-cover-change scenario (A2), 15 species lost 36% (SD 28) of their range. Five species lost significantly more range under the A2 scenario than the B1 scenario (p = 0.01, SD 0.01), suggesting that gibbons will benefit from effective management of land cover. PA coverage of future range was <30% for 11 species. On average, 32% (SD 25) of PCAs were covered by PAs. Indonesia contained more species and PCAs and thus has the greatest responsibility for gibbon conservation. Indonesia, India, and Myanmar need to expand their PAs to fulfill their responsibility to gibbon conservation. Our results provide a baseline for global gibbon conservation, particularly for countries lacking gibbon research capacity.     

Ecological Consequences of Recent Climate Change

Abstract: Global climate change is frequently considered a major conservation threat. The Earth's climate has already warmed by 0.5° C over the past century, and recent studies show that it is possible to detect the effects of a changing climate on ecological systems. This suggests that global change may be a current and future conservation threat. Changes in recent decades are apparent at all levels of ecological organization: population and life-history changes, shifts in geographic range, changes in species composition of communities, and changes in the structure and functioning of ecosystems. These ecological effects can be linked to recent population declines and to both local and global extinctions of species. Although it is impossible to prove that climate change is the cause of these ecological effects, these findings have important implications for conservation biology. It is no longer safe to assume that all of a species' historic range remains suitable. In drawing attention to the importance of climate change as a current threat to species, these studies emphasize the need for current conservation efforts to consider climate change in both in situ conservation and reintroduction efforts. Additional threats will emerge as climate continues to change, especially as climate interacts with other stressors such as habitat fragmentation. These studies can contribute to preparations for future challenges by providing valuable input to models and direct examples of how species respond to climate change.     

Economic costs of achieving current conservation goals in the future as climate changes

Conservation of biologically diverse regions has thus far been accomplished largely through the establishment and maintenance of protected areas. Climate change is expected to shift climate space of many species outside existing reserve boundaries. We used climate-envelope models to examine shifts in climate space of 11 species that are representative of the Mount Hamilton Project area (MHPA) (California, U.S.A.), which includes areas within Alameda, Santa Clara, San Joaquin, Stanislaus, Merced, and San Benito counties and is in the state's Central Coast ecoregion. We used Marxan site-selection software to determine the minimum area required as climate changes to achieve a baseline conservation goal equal to 80% of existing climate space for all species in the MHPA through 2050 and 2100. Additionally, we assessed the costs associated with use of existing conservation strategies (land acquisition and management actions such as species translocation, monitoring, and captive breeding) necessary to meet current species-conservation goals as climate changes. Meeting conservation goals as climate changes through 2050 required an additional 256,000 ha (332%) of protected area, primarily to the south and west of the MHPA. Through 2050 the total cost of land acquisition and management was estimated at US$1.67-1.79 billion, or 139-149% of the cost of achieving the same conservation goals with no climate change. To maintain 80% of climate space through 2100 required nearly 380,000 additional hectares that would cost $2.46-2.62 billion, or 209-219% of the cost of achieving the same conservation goals with no climate change. Furthermore, maintaining 80% of existing climate space within California for 27% of the focal species was not possible by 2100 because climate space for these species did not exist in the state. The high costs of conserving species as the climate changes-that we found in an assessment of one conservation project-highlights the need for tools that will aid in iterative goal setting given the uncertainty of the effects of climate change and adaptive management that includes new conservation strategies and consideration of the long-term economic costs of conservation.     

Multispecies and Multiscale Conservation Planning: Setting Quantitative Targets for Red‐Listed Lichens on Ancient Oaks

Abstract: Species occurrence in a habitat patch depends on local habitat and the amount of that habitat in the wider landscape. We used predictions from empirical landscape studies to set quantitative conservation criteria and targets in a multispecies and multiscale conservation planning effort. We used regression analyses to compare species richness and occurrence of five red‐listed lichens on 50 ancient oaks (Quercus robur; 120–140 cm in diameter) with the density of ancient oaks in circles of varying radius from each individual oak. Species richness and the occurrence of three of the five species were best explained by increasing density of oaks within 0.5 km; one species was best explained by the density of oaks within 2 km, and another was best predicted by the density of oaks within 5 km. The minimum numbers of ancient oaks required for “successful conservation” was defined as the number of oaks required to obtain a predicted local occurrence of 50% for all species included or a predicted local occurrence of 80% for all species included. These numbers of oaks were calculated for two relevant landscape scales (1 km² and 13 km²) that corresponded to various species responses, in such a way that calculations also accounted for local number of oaks. Ten and seven of the 50 ancient oaks surveyed were situated in landscapes that already fulfilled criteria for successful conservation when the 50% and 80% criteria, respectively, were used to define the level of successful conservation. For cost‐efficient conservation, oak stands in the landscapes most suitable for successful conservation should be prioritized for conservation and management (e.g., grazing and planting of new oaks) at the expense of oak stands situated elsewhere.     

Using Montane Mammals to Model Extinctions Due to Global Change

We use data on the species-area relationship and the nested subset structure of the boreal mammal faunas inhabiting isolated mountaintops in the Great Basin to develop a simple quantitative model that predicts the number and identity of species that would go extinct under an assumed scenario of changing climate and vegetation. Global warming of 3°C is predicted to cause the loss of 9–62% of the species inhabiting each mountain range and the extinction of three of fourteen species throughout the region. These results suggest (1) that it is possible to make highly plausible predictions about the susceptibility of species to extinction without detailed information about their population biology, and (2) that global and regional environmental changes seriously threaten the survival of species that are restricted in distribution to both natural "habitat islands" and biological reserves.     

Seeding phenology influences wood mouse seed choices: the overlooked role of timing in the foraging decisions by seed-dispersing rodents

Scatter-hoarding rodents influence the population dynamics of plants by acting as seed predators and dispersers. Therefore, rodent foraging preferences for certain seed traits (species, size, condition) have been extensively studied. However, to what extent these preferences are fixed or they track the temporal changes on seed characteristics due to phenological differences has been seldom explored. We studied the temporal variability in seed preferences by wood mouse (Apodemus sylvaticus), according to phenological changes in seed characteristics of two co-occurring oaks (Quercus ilex and Quercus pubescens). The phenology of acorn abundance and the acorn predation/dispersal patterns by rodents were monitored over an entire seeding season. Results revealed temporal changes in rodent preferences for acorns of the two oaks, matching their different seeding phenology (earlier in Q. pubescens and later in Q. ilex). On the other hand, whatever the species considered, rodents preferred larger and sound acorns along the entire season, although the dispersal of infested ones increased slightly during the peaks of acorn drop. The observed influence of seeding phenology on seed choices by rodents warns about inferring definite conclusions regarding their foraging behavior when arising from short-term experiments. Indeed, this study reveals that foraging preferences may be highly dynamic and context-dependent for some seed traits (e.g., species and condition), rather than fixed behavioral patterns. Plasticity in rodent foraging choices may allow them to successfully exploit different oaks with uncoupled seeding phenologies, while potentially favoring their coexistence.     

Climate heterogeneity modulates impact of warming on tropical insects

Evolutionary history and physiology mediate species responses to climate change. Tropical species that do not naturally experience high temperature variability have a narrow thermal tolerance compared to similar taxa at temperate latitudes and could therefore be most vulnerable to warming. However, the thermal adaptation of a species may also be influenced by spatial temperature variations over its geographical range. Spatial climate gradients, especially from topography, may also broaden thermal tolerance and therefore act to buffer warming impacts. Here we show that for low-seasonality environments, high spatial heterogeneity in temperature correlates significantly with greater warming tolerance in insects globally. Based on this relationship, we find that climate change projections of direct physiological impacts on insect fitness highlight the vulnerability of tropical lowland areas to future warming. Thus, in addition to seasonality, spatial heterogeneity may play a critical role in thermal adaptation and climate change impacts particularly in the tropics.     

Antagonistic effect of natural habitat conversion on community adjustment to climate warming in nonbreeding waterbirds

Although the impacts of climate and land-use changes on biodiversity have been widely documented, their joint effects remain poorly understood. We evaluated how nonbreeding waterbird communities adjust to climate warming along a gradient of land-use change. Using midwinter waterbird counts (132 species) at 164 major nonbreeding sites in 22 Mediterranean countries, we assessed the changes in species composition from 1991 to 2010, relative to thermal niche position and breadth, in response to regional and local winter temperature anomalies and conversion of natural habitats. We observed a low-level, nonsignificant community adjustment to the temperature increase where natural habitat conversion occurred. At the sites affected by natural habitat conversion, the relative increase of warm-dwelling species in response to climate warming was 6 times lower and the relative species decline was 3 times higher than in the sites without natural habitat conversion. We found no evidence of community adjustment to climate warming when natural habitat conversion was >5% over 15 years. This strong negative effect suggests an antagonistic interaction between climate warming and habitat change. These results underline the importance of habitat conservation in community adjustment to climate warming.     

Causes and consequences of unequal seedling production in forest trees: a case study in red oaks

Inequality in reproductive success has important implications for ecological and evolutionary dynamics, but lifetime reproductive success is challenging to measure in long-lived species such as forest trees. While seed production is often used as a proxy for overall reproductive success, high mortality of seeds and the potential for trade-offs between seed number and quality draw this assumption into question. Parentage analyses of established seedlings can bring us one step closer to understanding the causes and consequences of variation in reproductive success. In this paper we demonstrate a new method for estimating individual seedling production and average percentage germination, using data from two mixed-species populations of red oaks (Quercus rubra, Q. velutina, Q. falcata, and Q. coccinea). We use these estimates to examine the distribution of female reproductive success and to test the relationship between seedling number and individual seed production, age, and growth rate. We show that both seed and seedling production are highly skewed, roughly conforming to zero-inflated lognormal distributions, rather than to the Poisson or negative-binomial distributions often assumed by population genetics analyses. While the number of established offspring is positively associated with mean annual seed production, a lower proportion of seeds from highly fecund individuals become seedlings. Our red oak populations also show evidence of trade-offs between growth rate and reproductive success. The high degree of inequality in seedling production shown here for red oaks, and by previous studies in other species, suggests that many trees may be more vulnerable to genetic drift than previously thought, if immigration in limited by fragmentation or other environmental changes.     

Phenological tracking enables positive species responses to climate change

Earlier spring phenology observed in many plant species in recent decades provides compelling evidence that species are already responding to the rising global temperatures associated with anthropogenic climate change. There is great variability among species, however, in their phenological sensitivity to temperature. Species that do not phenologically "track" climate change may be at a disadvantage if their growth becomes limited by missed interactions with mutualists, or a shorter growing season relative to earlier-active competitors. Here, we set out to test the hypothesis that phenological sensitivity could be used to predict species performance in a warming climate, by synthesizing results across terrestrial warming experiments. We assembled data for 57 species across 24 studies where flowering or vegetative phenology was matched with a measure of species performance. Performance metrics included biomass, percent cover, number of flowers, or individual growth. We found that species that advanced their phenology with warming also increased their performance, whereas those that did not advance tended to decline in performance with warming. This indicates that species that cannot phenologically "track" climate may be at increased risk with future climate change, and it suggests that phenological monitoring may provide an important tool for setting future conservation priorities.     

Using environmental heterogeneity to plan for sea‐level rise

Environmental heterogeneity is increasingly being used to select conservation areas that will provide for future biodiversity under a variety of climate scenarios. This approach, termed conserving nature's stage (CNS), assumes environmental features respond to climate change more slowly than biological communities, but will CNS be effective if the stage were to change as rapidly as the climate? We tested the effectiveness of using CNS to select sites in salt marshes for conservation in coastal Georgia (U.S.A.), where environmental features will change rapidly as sea level rises. We calculated species diversity based on distributions of 7 bird species with a variety of niches in Georgia salt marshes. Environmental heterogeneity was assessed across six landscape gradients (e.g., elevation, salinity, and patch area). We used 2 approaches to select sites with high environmental heterogeneity: site complementarity (environmental diversity [ED]) and local environmental heterogeneity (environmental richness [ER]). Sites selected based on ER predicted present‐day species diversity better than randomly selected sites (up to an 8.1% improvement), were resilient to areal loss from SLR (1.0% average areal loss by 2050 compared with 0.9% loss of randomly selected sites), and provided habitat to a threatened species (0.63 average occupancy compared with 0.6 average occupancy of randomly selected sites). Sites selected based on ED predicted species diversity no better or worse than random and were not resilient to SLR (2.9% average areal loss by 2050). Despite the discrepancy between the 2 approaches, CNS is a viable strategy for conservation site selection in salt marshes because the ER approach was successful. It has potential for application in other coastal areas where SLR will affect environmental features, but its performance may depend on the magnitude of geological changes caused by SLR. Our results indicate that conservation planners that had heretofore excluded low‐lying coasts from CNS planning could include coastal ecosystems in regional conservation strategies.     

Predicting extinctions as a result of climate change

Widespread extinction is a predicted ecological consequence of global warming. Extinction risk under climate change scenarios is a function of distribution breadth. Focusing on trees and birds of the eastern United States, we used joint climate and environment models to examine fit and climate change vulnerability as a function of distribution breadth. We found that extinction vulnerability increases with decreasing distribution size. We also found that model fit decreases with decreasing distribution size, resulting in high prediction uncertainty among narrowly distributed species. High prediction uncertainty creates a conservation dilemma in that excluding these species under-predicts extinction risk and favors mistaken inaction on global warming. By contrast, including narrow endemics results in over-predicting extinction risk and promotes mistaken inaction on behalf of individual species prematurely considered doomed to extinction.     

Divergent effects of climate change on the egg-laying opportunity of species in cold and warm regions

Climate warming can substantially impact embryonic development and juvenile growth in oviparous species. Estimating the overall impacts of climate warming on oviparous reproduction is difficult because egg-laying events happen throughout the reproductive season. Successful egg laying requires the completion of embryonic development as well as hatching timing conducive to offspring survival and energy accumulation. We propose a new metric—egg-laying opportunity (EO)—to estimate the annual hours during which a clutch of freshly laid eggs yields surviving offspring that store sufficient energy for overwintering. We estimated the EO within the distribution of a model species, Sceloporus undulatus, under recent climate condition and a climate-warming scenario by combining microclimate data, developmental functions, and biophysical models. We predicted that EO will decline as the climate warms at 74.8% of 11,407 sites. Decreasing hatching success and offspring energy accounted for more lost EO hours (72.6% and 72.9%) than the occurrence of offspring heat stress (59.9%). Nesting deeper (at a depth of 12 cm) may be a more effective behavioral adjustment for retaining EO than using shadier (50% shade) nests because the former fully mitigated the decline of EO under the considered warming scenario at more sites (66.1%) than the latter (28.3%). We advocate for the use of EO in predicting the impacts of climate warming on oviparous animals because it encapsulates the integrative impacts of climate warming on all stages of reproductive life history.     

Global Warming and the Species Richness of Bats in Texas

General circulation models provide predictions for global climate under scenarios of increased atmospheric CO₂. Climate change is expected to lead directly to changes in distributions of vegetation associations. Distribution of animals will also change to the extent that animals rely on vegetation for food or shelter. Bat species in Texas appear to be restricted, in part, by the availability of roosts. We used geographic information systems and the Holdridge vegetation-climate association scheme to model the effect of climate change on bat distributions and species richness in Texas. Habitat characteristics for each species were identified from the literature and included vegetation, topography, and availability of caves. We assumed caves and topography to be fixed relative to climate. Vegetation changes were predicted from the Holdridge vegetation-climate association scheme. The redistribution of bats following climate change was predicted based on the new locations of suitable habitat characteristics. Under conditions of global warming tropical forests were predicted to expand into Texas; tree-roosting bats were sensitive to this change in vegetation. Cavity-roosting bats were less affected by changes in vegetation, but, where response was predicted, ranges decline.     

Benefits of protected areas for nonbreeding waterbirds adjusting their distributions under climate warming

Climate warming is driving changes in species distributions and community composition. Many species have a so-called climatic debt, that is, shifts in range lag behind shifts in temperature isoclines. Inside protected areas (PAs), community changes in response to climate warming can be facilitated by greater colonization rates by warm-dwelling species, but also mitigated by lowering extirpation rates of cold-dwelling species. An evaluation of the relative importance of colonization-extirpation processes is important to inform conservation strategies that aim for both climate debt reduction and species conservation. We assessed the colonization-extirpation dynamics involved in community changes in response to climate inside and outside PAs. To do so, we used 25 years of occurrence data of nonbreeding waterbirds in the western Palearctic (97 species, 7071 sites, 39 countries, 1993–2017). We used a community temperature index (CTI) framework based on species thermal affinities to investigate species turnover induced by temperature increase. We determined whether thermal community adjustment was associated with colonization by warm-dwelling species or extirpation of cold-dwelling species by modeling change in standard deviation of the CTI (CTI_SD). Using linear mixed-effects models, we investigated whether communities in PAs had lower climatic debt and different patterns of community change than communities outside PAs. For CTI and CTI_SD combined, communities inside PAs had more species, higher colonization, lower extirpation, and lower climatic debt (16%) than communities outside PAs. Thus, our results suggest that PAs facilitate 2 independent processes that shape community dynamics and maintain biodiversity. The community adjustment was, however, not sufficiently fast to keep pace with the large temperature increases in the central and northeastern western Palearctic. Our results underline the potential of combining CTI and CTI_SD metrics to improve understanding of the colonization-extirpation patterns driven by climate warming.     

Global Warming, Human Population Pressure, and Viability of the World's Smallest Butterfly

Abstract:  The effects of climate change and habitat destruction and their interaction are likely to be the greatest challenge to animal and plant conservation in the twenty-first century. We used the world's smallest butterfly, the Sinai baton blue ( Pseudophilotes sinaicus ), as an exemplar of how global warming and human population pressures may act together to cause species extinctions. We mapped the entire global range of this butterfly and obtained extensive data on the intensity of livestock grazing. As with an increasing number of species, it is confined to a network of small habitat patches and is threatened both by indirect human-induced factors (global warming) and by the direct activities of humans (in this case, livestock grazing and collection of medicinal plants). In the absence of global warming, grazing, and plant collection, our model suggested that the butterfly will persist for at least 200 years. Above a threshold intensity of global warming, the chance of extinction accelerated rapidly, implying that there may be an annual average temperature, specific to each endangered species, above which extinction becomes very much more likely. By contrast, there was no such threshold of grazing pressure—the chance of extinction increased steadily with increasing grazing. The impact of grazing, however, decreased with higher levels of year-to-year variation in habitat quality. The effect of global warming did not depend on the future level of grazing, suggesting that the impacts of global warming and grazing are additive. If the areas of habitat patches individually fall below certain prescribed levels, the butterfly is likely to go extinct. Two patches were very important for persistence: if either were lost the species would probably go extinct. Our results have implications for the conservation management of all species whose habitats are at risk because of the direct activities of humans and in the longer term because of climate change.