Minimum viable metapopulation size, extinction debt, and the conservation of a declining species.

A key question facing conservation biologists is whether declines in species' distributions are keeping pace with landscape change, or whether current distributions overestimate probabilities of future persistence. We use metapopulations of the marsh fritillary butterfly Euphydryas aurinia in the United Kingdom as a model system to test for extinction debt in a declining species. We derive parameters for a metapopulation model (incidence function model, IFM) using information from a 625-km2 landscape where habitat patch occupancy, colonization, and extinction rates for E. aurinia depend on patch connectivity, area, and quality. We then show that habitat networks in six extant metapopulations in 16-km2 squares were larger, had longer modeled persistence times (using IFM), and higher metapopulation capacity (lambdaM) than six extinct metapopulations. However, there was a > 99% chance that one or more of the six extant metapopulations would go extinct in 100 years in the absence of further habitat loss. For 11 out of 12 networks, minimum areas of habitat needed for 95% persistence of metapopulation simulations after 100 years ranged from 80 to 142 ha (approximately 5-9% of land area), depending on the spatial location of habitat. The area of habitat exceeded the estimated minimum viable metapopulation size (MVM) in only two of the six extant metapopulations, and even then by only 20%. The remaining four extant networks were expected to suffer extinction in 15-126 years. MVM was consistently estimated as approximately 5% of land area based on a sensitivity analysis of IFM parameters and was reduced only marginally (to approximately 4%) by modeling the potential impact of long-distance colonization over wider landscapes. The results suggest a widespread extinction debt among extant metapopulations of a declining species, necessitating conservation management or reserve designation even in apparent strongholds. For threatened species, metapopulation modeling is a potential means to identify landscapes near to extinction thresholds, to which conservation measures can be targeted for the best chance of success.     

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.     

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.     

Ecological Correlates of Extinction Proneness in Tropical Butterflies

Abstract:  Widespread and rapid losses of natural habitats and biodiversity have made the identification of extinction-prone species a major challenge in conservation biology. We assessed the relative importance of biologically relevant species traits (e.g., body size, ecological specialization) obtained from published records to determine the extinction probability of butterflies in a highly disturbed tropical landscape (i.e., Singapore). We also developed a taxon-specific model to estimate the extinction proneness of butterflies in Southeast Asia. Logistic regression analyses showed that adult habitat specialization, larval host plant specificity, geographical distribution, sexual dichromatism, and congenor density were significant and independent determinants of butterfly extinctions in Singapore. Among these traits, specificity of larval host plant and adult habitat specialization were the best correlates of extinction risks. We used this phenomenological extinction-regression model to estimate the relative extinction proneness of 416 butterfly species in Southeast Asia. Our results illustrate the utility of available taxon-specific data for a localized area in estimating the extinction proneness of closely related species on a regional scale. When intensive field studies are not forthcoming, especially in regions suffering from rapid biodiversity losses (e.g., Southeast Asia), similar approaches could be used to estimate extinction threats for other taxonomic groups.     

Influence of habitat quality, population size, patch size, and connectivity on patch-occupancy dynamics of the middle spotted woodpecker

Despite extensive research on the effects of habitat fragmentation, the ecological mechanisms underlying colonization and extinction processes are poorly known, but knowledge of these mechanisms is essential to understanding the distribution and persistence of populations in fragmented habitats. We examined these mechanisms through multiseason occupancy models that elucidated patch-occupancy dynamics of Middle Spotted Woodpeckers (Dendrocopos medius) in northwestern Spain. The number of occupied patches was relatively stable from 2000 to 2010 (15-24% of 101 patches occupied every year) because extinction was balanced by recolonization. Larger and higher quality patches (i.e., higher density of oaks >37 cm dbh [diameter at breast height]) were more likely to be occupied. Habitat quality (i.e., density of large oaks) explained more variation in patch colonization and extinction than did patch size and connectivity, which were both weakly associated with probabilities of turnover. Patches of higher quality were more likely to be colonized than patches of lower quality. Populations in high-quality patches were less likely to become extinct. In addition, extinction in a patch was strongly associated with local population size but not with patch size, which means the latter may not be a good surrogate of population size in assessments of extinction probability. Our results suggest that habitat quality may be a primary driver of patch-occupancy dynamics and may increase the accuracy of models of population survival. We encourage comparisons of competing models that assess occupancy, colonization, and extinction probabilities in a single analytical framework (e.g., dynamic occupancy models) so as to shed light on the association of habitat quality and patch geometry with colonization and extinction processes in different settings and species.     

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.     

Effects of Climate Change on Population Persistence of Desert-Dwelling Mountain Sheep in California

Abstract:  Metapopulations may be very sensitive to global climate change, particularly if temperature and precipitation change rapidly. We present an analysis of the role of climate and other factors in determining metapopulation structure based on presence and absence data. We compared existing and historical population distributions of desert bighorn sheep ( Ovis canadensis ) to determine whether regional climate patterns were correlated with local extinction. To examine all mountain ranges known to hold or to have held desert bighorn populations in California and score for variables describing climate, metapopulation dynamics, human impacts, and other environmental factors, we used a geographic information system (GIS) and paper maps. We used logistic regression and hierarchical partitioning to assess the relationship among these variables and the current status of each population (extinct or extant). Parameters related to climate—elevation, precipitation, and presence of dependable springs—were strongly correlated with population persistence in the twentieth century. Populations inhabiting lower, drier mountain ranges were more likely to go extinct. The presence of domestic sheep grazing allotments was negatively correlated with population persistence. We used conditional extinction probabilities generated by the logistic-regression model to rank native, naturally recolonized, and reintroduced populations by vulnerability to extinction under several climate-change scenarios. Thus risk of extinction in metapopulations can be evaluated for global-climate-change scenarios even when few demographic data are available.     

Extinction Rates in Archipelagoes: Implications for Populations in Fragmented Habitats

To study the effect of habitat fragmentation on population viability, I used extinction rates on islands in archipelagoes and estimated the relative probability of extinction per species on single large islands and sets of smaller islands with the same total area. Data on lizards, birds, and mammals on oceanic islands and mammals on mountaintops and in nature reserves yield similar results. Species are likely to go extinct on all the small islands before they go extinct on the single, large island. In the short term, the analysis indicates that extinction probabilities may be lower on a set of small islands. This is perhaps an artifact due to underestimation of extinction rates on small islands and/or the necessity of pooling species in a focal taxon to obtain estimates of extinction rates (which may obscure area thresholds and underestimate the slope and curvature of extinction rates as a function of area). Ultimately, cumulative extinction probabilities are higher for a set of small islands than for single large islands. Mean and median times to extinction tend to be shorter in the fragmented systems, in some cases much shorter. Thus, to minimize extinction rates in isolated habitat remnants and nature reserve systems, the degree of fragmentation should be minimized     

The difference conservation makes to extinction risk of the world's ungulates

Previous studies show that conservation actions have prevented extinctions, recovered populations, and reduced declining trends in global biodiversity. However, all studies to date have substantially underestimated the difference conservation action makes because they failed to account fully for what would have happened in the absence thereof. We undertook a scenario‐based thought experiment to better quantify the effect conservation actions have had on the extinction risk of the world's 235 recognized ungulate species. We did so by comparing species’ observed conservation status in 2008 with their estimated status under counterfactual scenarios in which conservation efforts ceased in 1996. We estimated that without conservation at least 148 species would have deteriorated by one International Union for Conservation of Nature (IUCN) Red List category, including 6 species that now would be listed as extinct or extinct in the wild. The overall decline in the conservation status of ungulates would have been nearly 8 times worse than observed. This trend would have been greater still if not for conservation on private lands. While some species have benefited from highly targeted interventions, such as reintroduction, most benefited collaterally from conservation such as habitat protection. We found that the difference conservation action makes to the conservation status of the world's ungulate species is likely to be higher than previously estimated. Increased, and sustained, investment could help achieve further improvements.     

Extinction, Colonization, and Metapopulations: Environmental Tracking by Rare Species

Extinction and metapopulation theories emphasize that stochastic fluctuations in local populations cause extinction and that local extinctions generate empty habitat patches that are then available for recolonization. Metapopulation persistence depends on the balance of extinction and colonization in a static environment. For many rare and declining species, I argue (1) that extinction is usually the deterministic consequence of the local environment becoming unsuitable (through habitat loss or modification, introduction of a predator, etc.); (2) that the local environment usually remains unsuitable following local extinction, so extinctions only rarely generate empty patches of suitable habitat; and (3) that colonization usually follows improvement of the local environment for a particular species (or long-distance transfer by humans). Thus, persistence depends predominantly on whether organisms are able to track the shifting spatial mosaic of suitable environmental conditions or on maintainance of good conditions locally.     

Thresholds in Songbird Occurrence in Relation to Landscape Structure

Abstract:  Theory predicts the occurrence of threshold levels of habitat in landscapes, below which ecological processes change abruptly. Simulation models indicate that below critical thresholds, fragmentation of habitat influences patch occupancy by decreasing colonization rates and increasing rates of local extinction. Uncovering such putative relationships is important for understanding the demography of species and in developing sound conservation strategies. Using segmented logistic regression, we tested for thresholds in occurrence of 15 bird species as a function of the amount of suitable habitat at multiple scales (150–2000-m radii). Suitable habitat was defined quantitatively based on previously derived, spatially explicit distribution models for each species. The occurrence of 10 out of 15 species was influenced by the amount of habitat at a landscape scale (≥500-m radius). Of these species all but one were best predicted by threshold models. Six out of nine species exhibited asymptotic thresholds; the effects of habitat loss intensified at low amounts of habitat in a landscape. Landscape thresholds ranged from 8.6% habitat to 28.7% (     = 18.5 ± 2.6%[95% CI]). For two species landscape thresholds coincided with sensitivity to fragmentation; both species were more likely to occur in large patches, but only when the amount of habitat in a landscape was low. This supports the fragmentation threshold hypothesis. Nevertheless, the occurrence of most species appeared to be unaffected by fragmentation, regardless of the amount of habitat present at landscape extents. The thresholds we identified may be useful to managers in establishing conservation targets. Our results indicate that findings of landscape-scale studies conducted in regions with relatively high proportions of habitat and low fragmentation may not be applicable in regions with low habitat proportions and high fragmentation.     

Synergisms among Habitat Fragmentation, Livestock Grazing, and Biotic Invasions in Southwestern Australia

Abstract: The agricultural development of southern Australia over the past 200 years has resulted in extensively fragmented systems, often with only small, isolated remnants of native vegetation remaining. Grazing by sheep and cattle has affected both the remaining fragments and the surrounding matrix, and non-native plant and animal species have had dramatic effects on the native biota. Invasive plant species have the potential to significantly alter ecosystem composition and functioning, and invasive animals, particularly rabbits ( Oryctolagus cuniculatus ), foxes (    Vulpes vulpes ) and cats (    Felis catus ) effectively alter habitat and drive native fauna to local extinction. These different influences often interact. For instance, smaller fragments are often more prone to plant invasion and are more likely to have been grazed in the past. Invasion of plant species is often linked with livestock grazing or rabbit invasion, and other higher-order interactions are also apparent. Classical fragmentation studies that concentrate on parameters such as habitat area and isolation but ignore changes in habitat condition brought about by livestock and invasive species are unlikely to yield meaningful results. Similarly, management of fragmented ecosystems must account for not only the spatial characteristics of the remaining habitat but also the importance of other influences, particularly those that impinge on fragments from the surrounding matrix.     

Direct and indirect effects of climate and habitat factors on butterfly diversity

Many factors, including climate, resource availability, and habitat diversity, have been proposed as determinants of global diversity, but the links among them have rarely been studied. Using structural equation modeling (SEM), we investigated direct and indirect effects of climate variables, host-plant richness, and habitat diversity on butterfly species richness across Britain, at 20-km grid resolution. These factors were all important determinants of butterfly diversity, but their relative contributions differed between habitat generalists and specialists, and whether the effects were direct or indirect. Climate variables had strong effects on habitat generalists, whereas host-plant richness and habitat diversity contributed relatively more for habitat specialists. Considering total effects (direct and indirect together), climate variables had the strongest link to butterfly species richness for all groups of species. The results suggest that different mechanistic hypotheses to explain species richness may be more appropriate for habitat generalists and specialists, with generalists hypothesized to show direct physiological limitations and specialists additionally being constrained by trophic interactions (climate affecting host-plant richness).     

Differences in gypsum plant communities associated with habitat fragmentation and livestock grazing.

The negative consequences of habitat fragmentation for plant communities have been documented in many regions of the world. In some fragmented habitats, livestock grazing has been proposed to be a dispersal mechanism reducing isolation between fragments. In others, grazing acts together with fragmentation in a way that increases habitat degradation. Iberian gypsum plant communities have been grazed and fragmented by agricultural practices for centuries. Although their conservation is considered a priority by the European Community, the effects of fragmentation on gypsum plant communities and the possible role of livestock grazing remain unknown. In addition, a substantial proportion of plant species growing in gypsum environments are gypsum specialists. They could be particularly affected by fragmentation, as was found for other habitat specialists (i.e., serpentine and calcareous specialists). In this study (1) we investigated the effect of fragmentation and grazing on gypsum plant community composition (species and life-forms), and (2) we tested to see if gypsum specialists were differently affected by fragmentation and grazing than habitat generalists. A vegetation survey was conducted in the largest gypsum outcrop of Europe (Middle Ebro Valley, northeast Spain). Fragmented and continuous sites in grazed and ungrazed areas were compared. Measurements related to species and composition of life-forms were contrasted first for the whole gypsum plant community and then specifically for the gypsum specialists. In the whole community, our results showed lower plant species diversity in fragmented sites, mainly due to the larger dominance of species more tolerant to fragmented habitat conditions. With livestock grazing, the plant species richness and the similarity in plant species composition between remnants was larger, suggesting that animals were acting as dispersal agents between fragments. As expected, gypsum specialists were less abundant in fragmented areas, and grazing led to the disappearance of the rare gypsum specialist Campanula fastigiata. According to our results, conservation strategies for gypsum plant communities in human-dominated landscapes should consider that fragmentation and grazing modify plant community composition affecting gypsum specialists in particular.     

Significance of Sighting Rate in Inferring Extinction and Threat

Abstract:  We are now entering a time of immense environmental upheaval in which, increasingly, experts are required to provide conservation assessments. Quantitative assessment of trends in species' range and abundance is costly, requiring extensive field studies over a long period of time. Unfortunately, many species are only known through a few "chance" sightings or a handful of specimens, and therefore extinction may be even harder to ascertain. Several methods have been proposed for estimating the probability of extinction. However, comparison within and between species is difficult because of variations in sighting rates. We applied a probabilistic method that incorporates sighting rate to the sighting record of Vietnamese slipper orchids ( Paphiopedilum ). The method generates a probability that another sighting will occur given the previous sighting rate and the time since last observation. This allows greater comparability between species discovered at different times. Its predictions were more highly correlated with the World Conservation Union criteria than previous methods. Trends in data collection and the political climate of a country, which affects access to material, are important potential sources of variation that affect sighting rates. A lack of understanding of the process by which data are generated makes inferring extinction from sighting records difficult because extinction status depends on how the sighting rate varies. However, such methods allow rapid conservation prioritization of taxa that are poorly known and would otherwise go unassessed.     

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.     

Negative effects of nitrogen deposition on Swiss butterflies

Nitrogen (N) deposition from agriculture and combustion of fossil fuels is a major threat to plant diversity, but its effects on organisms at higher trophic levels are unclear. We investigated how N deposition may affect species richness and abundance (number of individuals per species) in butterflies. We reviewed the peer-reviewed literature on variables used to explain spatial variation in butterfly species richness and found that vegetation variables appeared to be as important as climate and habitat variables in explaining butterfly species richness. It thus seemed likely that increased N deposition could indirectly affect butterfly communities via its influence on plant communities. To test this prediction, we analyzed data from the Swiss biodiversity monitoring program for vascular plants and butterflies in 383 study sites of 1 km² that are evenly distributed throughout Switzerland. The area has a modeled N deposition gradient of 2–44 kg N ha⁻¹ year⁻¹. We used traditional linear models and structural equation models to infer the drivers of the spatial variation in butterfly species richness across Switzerland. High N deposition was consistently linked to low butterfly diversity, suggesting a net loss of butterfly diversity through increased N deposition. We hypothesize that at low elevations, N deposition may contribute to a reduction in butterfly species richness via microclimatic cooling due to increased plant biomass. At higher elevations, negative effects of N deposition on butterfly species richness may also be mediated by reduced plant species richness. In most butterfly species, abundance was negatively related to N deposition, but the strongest negative effects were found for species of conservation concern. We conclude that in addition to factors such as intensified agriculture, habitat fragmentation, and climate change, N deposition is likely to play a key role in negatively affecting butterfly diversity and abundance.     

Identifying and Managing Threatened Invertebrates through Assessment of Coextinction Risk

Abstract: Invertebrates with specific host species may have a high probability of extinction when their hosts have a high probability of extinction. Some of these invertebrates are more likely to go extinct than their hosts, and under some circumstances, specific actions to conserve the host may be detrimental to the invertebrate. A critical constraint to identifying such invertebrates is uncertainty about their level of host specificity. We used two host‐breadth models that explicitly incorporated uncertainty in the host specificity of an invertebrate species. We devised a decision protocol to identify actions that may increase the probability of persistence of a given dependent species. The protocol included estimates from the host‐breadth models and decision nodes to identify cothreatened species. We applied the models and protocol to data on 1055 insects (186 species) associated with 2 threatened (as designated by the Australian Government) plant species and 19 plant species that are not threatened to determine whether any insect herbivores have the potential to become extinct if the plant becomes extinct. According to the host‐breadth models, 18 species of insect had high host specificity to the threatened plant species. From these 18 insects, the decision protocol highlighted 6 species that had a high probability of extinction if their hosts were to become extinct (3% of all insects examined). The models and decision protocol have added objectivity and rigor to the process of deciding which dependent invertebrates require conservation action, particularly when dealing with largely unknown and speciose faunas.     

The Potential Conservation Value of Non‐Native Species

Abstract: Non‐native species can cause the loss of biological diversity (i.e., genetic, species, and ecosystem diversity) and threaten the well‐being of humans when they become invasive. In some cases, however, they can also provide conservation benefits. We examined the ways in which non‐native species currently contribute to conservation objectives. These include, for example, providing habitat or food resources to rare species, serving as functional substitutes for extinct taxa, and providing desirable ecosystem functions. We speculate that non‐native species might contribute to achieving conservation goals in the future because they may be more likely than native species to persist and provide ecosystem services in areas where climate and land use are changing rapidly and because they may evolve into new and endemic taxa. The management of non‐native species and their potential integration into conservation plans depends on how conservation goals are set in the future. A fraction of non‐native species will continue to cause biological and economic damage, and substantial uncertainty surrounds the potential future effects of all non‐native species. Nevertheless, we predict the proportion of non‐native species that are viewed as benign or even desirable will slowly increase over time as their potential contributions to society and to achieving conservation objectives become well recognized and realized.     

Effects of past and present livestock grazing on herpetofauna in a landscape‐scale experiment

Livestock grazing is the most widespread land use on Earth and can have negative effects on biodiversity. Yet, many of the mechanisms by which grazing leads to changes in biodiversity remain unresolved. One reason is that conventional grazing studies often target broad treatments rather than specific parameters of grazing (e.g., intensity, duration, and frequency) or fail to account for historical grazing effects. We conducted a landscape‐scale replicated grazing experiment (15,000 km², 97 sites) to examine the impact of past grazing management and current grazing regimes (intensity, duration, and frequency) on a community of ground‐dwelling herpetofauna (39 species). We analyzed community variables (species richness and composition) for all species and built multiseason patch‐occupancy models to predict local colonization and extinction for the 7 most abundant species. Past grazing practices did not influence community richness but did affect community composition and patch colonization and extinction for 4 of 7 species. Present grazing parameters did not influence community richness or composition, but 6 of the 7 target species were affected by at least one grazing parameter. Grazing frequency had the most consistent influence, positively affecting 3 of 7 species (increased colonization or decreased extinction). Past grazing practice affected community composition and population dynamics in some species in different ways, which suggests that conservation planners should examine the different grazing histories of an area. Species responded differently to specific current grazing practices; thus, incentive programs that apply a diversity of approaches rather than focusing on a change such as reduced grazing intensity should be considered. Based on our findings, we suggest that determining fine‐scale grazing attributes is essential for advancing grazing as a conservation strategy.