Human demography and reserve size predict wildlife extinction in West Africa.

Species-area models have become the primary tool used to predict baseline extinction rates for species in isolated habitats, and have influenced conservation and land-use planning worldwide. In particular, these models have been used to predict extinction rates following the loss or fragmentation of natural habitats in the absence of direct human influence on species persistence. Thus, where direct human influences, such as hunting, put added pressure on species in remnant habitat patches, we should expect to observe extinction rates higher than those predicted by simple species-area models. Here, we show that extinction rates for 41 species of large mammals in six nature reserves in West Africa are 14-307 times higher than those predicted by models based on reserve size alone. Human population and reserve size accounted for 98% of the observed variation in extinction rates between reserves. Extinction occurred at higher rates than predicted by species-area models for carnivores, primates and ungulates, and at the highest rates overall near reserve borders. Our results indicate that, where the harvest of wildlife is common, conservation plans should focus on increasing the size of reserves and reducing the rate of hunting.     

Comparison of reintroduction and enhancement effects on metapopulation viability

Metapopulation viability depends upon a balance of extinction and colonization of local habitats by a species. Mechanisms that can affect this balance include physical characteristics related to natural processes (e.g. succession) as well as anthropogenic actions. Plant restorations can help to produce favorable metapopulation dynamics and consequently increase viability; however, to date no studies confirm this is true. Population viability analysis (PVA) allows for the use of empirical data to generate theoretical future projections in the form of median time to extinction and probability of extinction. In turn, PVAs can inform and aid the development of conservation, recovery, and management plans. Pitcher's thistle (Cirsium pitcheri) is a dune endemic that exhibited metapopulation dynamics. We projected viability of three natural and two restored populations with demographic data spanning 15–23 years to determine the degree the addition of reintroduced population affects metapopulation viability. The models were validated by comparing observed and projected abundances and adjusting parameters associated with demographic and environmental stochasticity to improve model performance. Our chosen model correctly predicted yearly population abundance for 60% of the population‐years. Using that model, 50‐year projections showed that the addition of reintroductions increases metapopulation viability. The reintroduction that simulated population performance in early‐successional habitats had the maximum benefit. In situ enhancements of existing populations proved to be equally effective. This study shows that restorations can facilitate and improve metapopulation viability of species dependent on metapopulation dynamics for survival with long‐term persistence of C. pitcheri in Indiana likely to depend on continued active management.     

Using population viability analysis to predict the effect of fire on the extinction risk of an endangered shrub <Emphasis Type="Italic">Verticordia fimbrilepis</Emphasis> subsp. <Emphasis Type="Italic">fimbrilepis</Emphasis> in a fragmented landscape

The fragmentation of mediterranean climate landscapes where fire is an important landscape process may lead to unsuitable fire regimes for many species, particularly rare species that occur as small isolated populations. We investigate the influence of fire interval on the persistence of population fragments of the endangered shrub Verticordia fimbrilepis Turcz. subsp. fimbrilepis in mediterranean climate south-west of Western Australia. We studied the population biology of the species over 5 years. While the species does recruit sporadically without fire this occurs only in years with above average rainfall, so fire seems to be the main environmental factor producing extensive recruitment. Transition matrix models were constructed to describe the shrub’s population dynamics. As the species is killed by fire and relies on a seed bank stimulated to germinate by smoke, stochastic simulations to compare different fire frequencies on population viability were completed. Extinction risk increased with increasing average fire interval. Initial population size was also important, with the lowest extinction risk in the largest population. For populations in small reserves where fire is generally excluded, inevitable plant senescence will lead to local extirpation unless fires of suitable frequency can be used to stimulate regeneration. While a suitable fire regime reduces extinction risk small populations are still prone to extinction due to stochastic influences, and this will be exacerbated by a projected drying climate increasing rates of adult mortality and also seedling mortality in the post-fire environment.     

Logic for designing nature reserves for multiple species

We examine the logic of designing nature reserves to understand better how to integrate the concepts of representativeness and persistence. Simple models of viability are used to evaluate how the expected number of species in the reserve system changes with variation in the risk of extinction among species, their rate of occurrence, and the distribution of species. The optimal size of individual reserves increased with the mean and variance of the probability of extinction among species and with the rate at which the risk of extinction declines with the cost of each reserve. In contrast, the rate of occurrence of species within reserves and their rate of accumulation with increasing reserve area had a relatively minor influence on the optimal size of reserves. Patterns of endemism were most important for the location of reserves. Including differences among species in the analysis reduced the optimal number of individual reserves (and increased the size of each) when operating under a fixed budget compared with reserve designs based on single species. A case study in the city of Melbourne, Australia, demonstrates the conservation value of small (approximately 1 ha) grassland reserves and the underrepresentation of Melbourne's volcanic plains in the region's conservation network.     

种群生存力分析研究进展和趋势

种群生存力分析（PVA）是正在迅速发展的新方法,已成为保护生物学研究的热点。它主要研究随机干扰对小种群绝灭的影响,其目的是制定最小可存活种群（MVP）,把绝灭减少到可接受的水平。随机干扰可分四类;统计随机性,环境随机性,自然灾害和遗传随机性。确定MVP的方法有三种：理论模型,模拟模型,模拟模型和岛屿生物地理学方法。理论模型主要研究理想或特定条件下随机因素对种群的影响;模拟模型是利用计算机模拟种群绝灭过程;岛屿生物地理学方法主要分析岛屿物种的分布和存活,证实分析模型和模拟模型。已有大量的文献研究统计随机性,环境随机性和自然灾害的行为特征,但遗传因素与种群生存力之间的关系还不清楚。建立包括四种随机性的综合性模型,广泛地检验PVA模型,系统地研制目标种的遗传和生态特性以及MVP的实际应用是PVA的发展趋势。     

Designing nature reserves in the face of uncertainty

Conservation reserves are a fundamental tool for managing biodiversity. The so-called SLOSS debate--should we have a Single Large Or Several Small reserves - is central to conservation theory. Population dynamic models suggest that the design that minimizes the risk of extinction of a species is case-specific, with the optimal number of reserves ranging between one and very many. Uncertainty is pervasive in ecology, but, the previous analyses of the SLOSS debate have not considered how uncertainty in the model of extinction risk might influence the optimal design. Herein, we show that when uncertainty is considered, the SLOSS problem is simplified and driven more by the aspirations of the manager than the population dynamics of the species. In this case, the optimal solution is to have in the order of twenty or fewer reserves for any species. This result shows counter-intuitively that considering uncertainty actually simplifies rather than complicates decisions about designing nature reserves.     

Variation Thresholds for Extinction and Their Implications for Conservation Strategies

We examine the degree to which fitting simple dynamic models to time series of population counts can predict extinction probabilities. This is both an active branch of ecological theory and an important practical topic for resource managers. We introduce an approach that is complementary to recently developed techniques for estimating extinction risks (e.g., diffusion approximations) and, like them, requires only count data rather than the detailed ecological information available for traditional population viability analyses. Assuming process error, we use four different models of population growth to generate snapshots of population dynamics via time series of the lengths commonly available to ecologists. We then ask to what extent we can identify which of several broad classes of population dynamics is evident in the time series snapshot. Along the way, we introduce the idea of "variation thresholds," which are the maximum amount of process error that a population may withstand and still have a specified probability of surviving for a given length of time. We then show how these thresholds may be useful to both ecologists and resource managers, particularly when dealing with large numbers of poorly understood species, a common problem faced by those designing biodiversity reserves.     

Demographic effects of temperature‐dependent sex determination: will tuatara survive global warming?

Global climate change is of particular concern for small and isolated populations of reptiles with temperature-dependent sex determination because low genetic variation can limit adaptive response in pivotal temperatures, leading to skewed sex ratios. We explore the demographic consequences of skewed sex ratios on the viability of a tuatara population characterized by low genetic diversity. We studied the rare species of tuatara ( Sphenodon guntheri ) on the 4 ha North Brother Island in New Zealand over two nesting seasons and captured 477 individuals, with a 60% male bias in the adult population. Females first breed at 15 years and have extremely low rates of gravidity, producing clutches of three to eight eggs every 9 years. Simulations of the population using population viability analysis showed that the current population is expected to persist for at least 2000 years at hatchling sex ratios of up to 75% male, but populations with 85% male hatchlings are expected to become extinct within approximately 300 years (some eight generations). Incorporation of inbreeding depression increased the probability of extinction under male biased sex ratios, with no simulated populations surviving at hatchling sex ratios >75% male. Because recent models have predicted that climate change could lead to the production of all male S. guntheri hatchlings by 2085, we examined whether periodic intervention to produce mixed or female biased sex ratios would allow the population to survive if only males were produced in natural nests. We show that intervention every 2–3 years could buffer the effects of climate change on population sex ratios, but translocation to cooler environs might be more cost-effective. Climate change threatens tuatara populations because neither modified nesting behaviour nor adaptive response of the pivotal temperature can modify hatchling sex ratios fast enough in species with long generation intervals.     

Spatially-correlated extinction in a metapopulation model of Leadbeater's Possum

The importance of considering spatially-correlated extinction in metapopulation viability analyses was investigated using a model of the population dynamics of Gymnobelideus leadbeateri McCoy (Leadbeater's Possum). Fire caused local extinction of G. leadbeateri and induced changes in the suitability of the habitat over a period of decades and centuries. Spatially-correlated fires, in which the correlation between the incidence of fire declines with distance, and uniformly-correlated fires were simulated. The predicted risk of metapopulation extinction increased: (i) as the variance in the number of fires each year increased, (ii) as the mean fire interval decreased, and (iii) as the mean dispersal distance decreased. Incorporating spatial correlation in the incidence of fires between patches had little effect on the results, provided the variance in the number of fires per year remained the same and fires modified habitat quality. The predicted risk of metapopulation extinction was greater for spatially-correlated fires than for uniformly-correlated fires when fires only caused local extinction but did not change habitat suitability. Incorporating spatial correlation in the incidence of fire within patches, which allowed partial burning of patches, reduced the predicted risk of extinction. This effect was only slight when patches were smaller than about 50 ha. The results of our simulations demonstrate the importance of considering correlations in disturbance regimes in metapopulation models, especially if these models are used to assist the design of nature reserves.     

Life-history models of extinction: a test with island spiders

This study analyzes extinction patterns for two species of orb spiders monitored annually on 77 islands over a continuous 20-yr period. One species, Argiope argentata, has large populations sometimes crashing quickly to extinction and a much weaker relation of extinction likelihood to population size than does the other species, Metepeira datona. Demographic models were built for both species and matched against observations. Differences between the species in life-history traits-estimated with measurements from the field-together with incorporation of demographic stochasticity, a population ceiling, and environmental stochasticity, were necessary to fit the observed extinction curves. As predicted from life-history patterns, long-term population growth rates (and hence predicted extinction probabilities) are relatively very sensitive to values of juvenile survivorship. Models are also sensitive to variation in the population ceiling and environmental noise, which tend to act in a complementary manner. A simple model with no age structure was able to fit the data on large initial population sizes but not on small initial population sizes, showing that life cycle characteristics interact with the various sources of stochasticity and hence have to be taken into account to produce a precise model of the extinction process.     

Sheltered from the storm? Population viability analysis of a rare endemic under periodic catastrophe regimes

Rare species are important targets for biodiversity conservation efforts because rarity often equates to small populations and increased endangerment. Rare species are prone to stochastic extinction events and may be particularly susceptible to catastrophes. Therefore, understanding how rare species respond to disturbances is critical for evaluating extinction risk and guiding conservation managers. Population viability analyses (PVAs) are essential for assessing rare species' status yet they seldom consider catastrophic events. Accordingly, we present a PVA of a rare tropical epiphyte, Lepanthes caritensis (Orchidaceae), under simulated disturbance regimes to evaluate its demographics and extinction risk. We aimed to test how demographic models incorporating catastrophes affect population viability estimates. Our goal was to better guide management of these orchids and other rare plants. Results revealed L. caritensis numbers have declined recently, but projected growth rates indicated that most subpopulations should increase in size if undisturbed. Still, projection models show that moderate catastrophes reduce growth rates, increase stochasticity in subpopulation sizes, and elevate extinction risk. Severe catastrophes had a more pronounced effect in simulations; growth rates fell below replacement level, there was greater variation in projected population sizes, and extinction risk was significantly higher. PVAs incorporating periodic catastrophes indicate that rare species may have greater extinction probabilities than standard models suggest. Thus, precautionary conservation measures should be taken in disturbance prone settings and we encourage careful monitoring after environmental catastrophes. Future rare plant PVAs should incorporate catastrophes and aim to determine if rescue and reintroduction efforts are necessary after disturbances to insure long-term population viability.     

Time to extinction in deteriorating environments

Habitat degradation and destruction are the predominant drivers of population extinction, but there is little theory to guide the analysis of population viability in deteriorating environments. To address this gap, we investigated extinction times in time-varying, demographically stochastic versions of the logistic model for population dynamics. A property of these models is the “extinction delay,” a quantitative measure of the time lag in extinction created by species-specific extinction debt. For completeness, three models were constructed to represent the different demographic routes by which deterioration may affect population dynamics. Numerical analysis for two notional life histories indicated that the demographic response to environmental deterioration had a large effect on extinction delay, but a third analysis showed that the trajectory of the decline in carrying capacity ultimately characterized its magnitude. A concave decline in carrying capacity produced a large extinction delay while a small delay occurred with a convex decline. Furthermore, our results explore the non-monotonicity of extinction debt with respect to the speed of deterioration. A peak is present at low levels of deterioration, and the height of the peak and the asymptote of delay are affected by both life history parameterizations and the rate of change of the carrying capacity. The results suggest that population viability analyses must consider not only environmental deterioration, but also the effects of deterioration on the trajectory of the decline in carrying capacity.     

Considering threats to population viability of the endangered Korean long-tailed goral (Naemorhedus caudatus) using VORTEX

Population viability analysis (PVA) has frequently been used in conservation biology to predict extinction rates for threatened or endangered species. In this study, we used VORTEX to model Korean long-tailed goral (Naemorhedus caudatus) using previously collected ecological data. We focused on modelling population extinction, mean population size and heterozygosity. The minimum viable population size was found to be at least 50 gorals for 100 years, regardless of carrying capacity. However, populations with fewer than 50 gorals could not remain successful in the model. Inbreeding depression, catastrophes and supplementation also affected patterns of population extinction, mean population size and heterozygosity. Supplementation with new individuals had the strongest effect on extinction, mean population size and heterozygosity, followed by initial population size, inbreeding, catastrophes and carrying capacity. These results suggest that a supplementation by extra goral individuals from goral proliferation facilities would be the most helpful means for the restoration programme. More Korean goral-specific information regarding demographic and habitat parameters is needed for further PVA of the species.     

Habitat destruction and the extinction debt revisited: The Allee effect

Habitat destruction, often caused by anthropogenic disturbance, can lead to the extinction of species at an unprecedented rate. It is important, therefore, to consider habitat destruction when assessing population viability. Another factor often ignored in population viability analysis, is the Allee effect that adds to the risk of populations already on the verge of extinction. Understanding the Allee effect on species dynamics and response to habitat destruction has intrinsic value in conservation prioritization. Here, the Allee effect was considered in a multi-species hierarchical competition model. Results showed that species persistence declines dramatically due to the Allee effect, and certain species become more susceptible to habitat destruction than others. Two extinction orders emerged under habitat destruction: either the best competitor becomes extinct first or the best colonizer first. The extinction debt and order, as well as the time lag between habitat destruction and species extinction, were found to be determined by species abundance and the intensity of the Allee effect.     

Density-dependent demographic variation determines extinction rate of experimental populations

Understanding population extinctions is a chief goal of ecological theory. While stochastic theories of population growth are commonly used to forecast extinction, models used for prediction have not been adequately tested with experimental data. In a previously published experiment, variation in available food was experimentally manipulated in 281 laboratory populations of Daphnia magna to test hypothesized effects of environmental variation on population persistence. Here, half of those data were used to select and fit a stochastic model of population growth to predict extinctions of populations in the other half. When density-dependent demographic stochasticity was detected and incorporated in simple stochastic models, rates of population extinction were accurately predicted or only slightly biased. However, when density-dependent demographic stochasticity was not accounted for, as is usual when forecasting extinction of threatened and endangered species, predicted extinction rates were severely biased. Thus, an experimental demonstration shows that reliable estimates of extinction risk may be obtained for populations in variable environments if high-quality data are available for model selection and if density-dependent demographic stochasticity is accounted for. These results suggest that further consideration of density-dependent demographic stochasticity is required if predicted extinction rates are to be relied upon for conservation planning.     

Climate change and bet-hedging: interactions between increased soil temperatures and seed bank persistence

In order to predict the long-term consequences of climate change, it is necessary to link future environmental changes to mechanisms that control plant population processes. This information can then be incorporated into strategies to more accurately model climate change impacts on species or to estimate future extinction risks. We examined the impact of increased temperatures on the longevity and dynamics of the persistent soil seed banks of eight ephemeral species from arid Australia. We found that the predicted global temperature increases under climate change will be reflected in increased soil temperatures, and that seeds in the soil seed bank will be exposed to long durations of high temperatures over the summer months. Three of the eight species studied had significantly greater levels of germination after exposure to predicted increased soil temperatures. Another species displayed a dramatic decrease in seed viability after such exposure. The capacity of such species to use the seed bank to bet hedge against rainfall events that cause germination but are insufficient to allow plant maturation, is compromised by increased germinability and subsequent loss or reduction of seed bank persistence. These predicted changes in the dynamics of soil seed banks increase the risk of local extinctions of these species, while the composition of the community may be altered by changes in species abundance. Our results show that the risk spreading mechanism provided by persistent seed banks could be compromised by the mechanistic impact of forecast temperature increases in arid habitats, and highlight the need to understand mechanisms that control population dynamics when attempting to address likely future impacts of climate change on biodiversity.     

Theory for designing nature reserves for single species

We examine the question of the optimal number of reserves that should be established to maximize the persistence of a species. We assume that the mean time to extinction of a single population increases as a power of the habitat area, that there is a certain amount of habitat to be reserved, and that the aim is to determine how this habitat is most efficiently divided. The optimal configuration depends on whether the management objective is to maximize the mean time to extinction or minimize the risk of extinction. When maximizing the mean time to extinction, the optimal number of independent reserves does not depend on the amount of available habitat for the reserve system. In contrast, the risk of extinction is minimized when individual reserves are equal to the optimal patch size, making the optimal number of reserves linearly proportional to the amount of available habitat. A model that includes dispersal and correlation in the incidence of extinction demonstrates the importance of considering the relative rate at which these two factors decrease with distance between reserves. A small number of reserves is optimal when the mean time to extinction increases rapidly with habitat area or when risks of extinction are high.     

Population Viability, Nature Reserves, and the Outlook for Gray Wolf Conservation in North America

Theoretical work on population viability and extinction probabilities, empirical data from Canis lupus (gray wolf) populations, and expert opinion provide only general and conflicting conclusions about the number of wolves and the size of areas needed for conservation of wolf populations. There is no threshold population size or proven reserve design that guarantees long-term (century or more) survival for a gray wolf population. Most theoretical analyses of population viability have assumed a single, isolated population and lack of management intervention, neither of which is likely for wolves. Data on survival of actual wolf populations suggest greater resiliency than is indicated by theory. In our view, the previous theoretical treatments of population viability have not been appropriate to wolves, have contributed little to their conservation, and have created unnecessary dilemmas for wolf recovery programs by overstating the required population size. Nonetheless, viability as commonly understood may be problematic for small populations at the fringe of or outside the contiguous species range, unless they are part of a metapopulation. The capability of existing nature reserves to support viable wolf populations appears related to a variety of in situ circumstances, including size, shape and topography of the reserve; productivity, numbers, dispersion, and seasonal movement of prey; extent of poaching inside; degree of persecution outside; exposure to enzootica; attitudes of local people; and proximity to other wolf populations. We estimate that a population of 100 or more wolves and a reserve of several thousand square kilometers may be necessary to maintain a viable population in complete isolation, although 3000 km² or even 500–1000 km² may be adequate under favorable circumstances. In most cases, management intervention is probably necessary to assure the viability of relatively small, isolated populations. Because most reserves may be inadequate by themselves to ensure the long-term survival of wolf populations, favorable human attitudes toward the species and its management must be recognized as paramount, and cooperation of neighboring management jurisdictions will be increasingly important.     

Viability analysis of a threatened amphibian population: modelling the past,present and future

Conservation actions that have maintained populations in the past may not necessarily do so in the future. Population viability analysis provides one tool for exploring the impact of management actions on large temporal scales. However, there are relatively few long‐term data sets that provide the demographic and environmental data demanded by such models. Using a 37‐yr data set, we used RAMAS Metapop to model the persistence of natterjack toads Epidalea (Bufo) calamita on a heathland in southern Britain. A retrospective analysis showed that the best fit between the predicted population trajectories and the real population was when the management carried out was modelled as an increase in K of 150 toads yr^?1. However, even if ongoing management continues to improve K by a further 40–60 toads yr^?1 over the next 50 yr, the population still has an extinction risk of at least 60% if other factors remain unchanged. Sensitivity analyses and simulated management scenarios indicated that the population was most sensitive to changes in the survival of juvenile (i.e. 1–2 yr old) toads. In addition, if the frequency and severity of pond desiccation increases, the risk of extinction was predicted to increase as a result of reduced recruitment. Low levels of extinction risk occurred irrespective of K when juvenile survival was enhanced in combination with low frequency and severity of pond desiccation. The models suggest that populations that are responding to management against a background of natural fluctuations may remain vulnerable to extinction for several decades. These extinction risks may increase if habitat management fails to offset reductions in recruitment and juvenile survival caused by environmental change.     

Plant extinction risk under climate change: are forecast range shifts alone a good indicator of species vulnerability to global warming?

Models that couple habitat suitability with demographic processes offer a potentially improved approach for estimating spatial distributional shifts and extinction risk under climate change. Applying such an approach to five species of Australian plants with contrasting demographic traits, we show that: (i) predicted climate‐driven changes in range area are sensitive to the underlying habitat model, regardless of whether demographic traits and their interaction with habitat patch configuration are modeled explicitly; and (ii) caution should be exercised when using predicted changes in total habitat suitability or geographic extent to infer extinction risk, because the relationship between these metrics is often weak. Measures of extinction risk, which quantify threats to population persistence, are particularly sensitive to life‐history traits, such as recruitment response to fire, which explained approximately 60% of the deviance in expected minimum abundance. Dispersal dynamics and habitat patch structure have the strongest influence on the amount of movement of the trailing and leading edge of the range margin, explaining roughly 40% of modeled structural deviance. These results underscore the need to consider direct measures of extinction risk (population declines and other measures of stochastic viability), as well as measures of change in habitat area, when assessing climate change impacts on biodiversity. Furthermore, direct estimation of extinction risk incorporates important demographic and ecosystem processes, which potentially influence species’ vulnerability to extinction due to climate change.