A model for behavioral regulation of metapopulation dynamics

Habitat fragmentation can decrease local population persistence by reducing connectivity, which is a function of dispersal of individuals among habitat fragments. Dispersal is often treated as diffusion in population models, even though for many species it is a result of a series of behavioral decisions. We developed a metapopulation model to explore the potential importance of dispersal behaviors in driving metapopulation dynamics. We incorporated types of behavior that affect dispersal—colonization inhibiting, colonization enhancing, extinction inhibiting, extinction enhancing, rescue enhancing, rescue inhibiting—into Levins’ (1969) metapopulation model and projected occupancy rates for a variety of parameter values. Examples from the literature of behaviors associated with each of these parameters are provided. Our model simplifies into previously published metapopulation models that incorporate only a single behavior, and we present a density-dependent rescue function that leads to multiple non-zero equilibria. We found a variety of behavioral effects on metapopulations. Rescue enhancement fills patches faster than does colonization enhancement or extinction inhibition, and declines in patch occupancy are moderate with extinction enhancement, but colonization inhibition causes metapopulation extinction. We also found that with colonization and extinction inhibitions, equilibrium patch occupancy is inversely related to patch turnover rate. With density-dependent rescue, persistence depends not only on the strength of the strong rescue effect, but also on having a sufficient initial fraction of patches occupied; the stronger the rescue effect, the lower this fraction can be. This study suggests that dispersal behavior can have strong influences on metapopulation dynamics. It confirms the importance of understanding the relationship between landscape structure and dispersal behavior in understanding population persistence.     

Examining the relationship between local extinction risk and position in range

Over half of globally threatened animal species have experienced rapid geographic range loss. Identifying the parts of species’ distributions most vulnerable to local extinction would benefit conservation planning. However, previous studies give little consensus on whether ranges decline to the core or edge. We built on previous work by using empirical data to examine the position of recent local extinctions within species’ geographic ranges, address range position as a continuum, and explore the influence of environmental factors. We aggregated point‐locality data for 125 Galliform species from across the Palearctic and Indo‐Malaya into equal‐area half‐degree grid cells and used a multispecies dynamic Bayesian occupancy model to estimate rates of local extinctions. Our model provides a novel approach to identify loss of populations from within species ranges. We investigated the relationship between extinction rates and distance from range edge by examining whether patterns were consistent across biogeographic realm and different categories of land use. In the Palearctic, local extinctions occurred closer to the range edge than range core in both unconverted and human‐dominated landscapes. In Indo‐Malaya, no pattern was found for unconverted landscapes, but in human‐dominated landscapes extinctions tended to occur closer to the core than the edge. Our results suggest that local and regional factors override general spatial patterns of recent local extinction within species’ ranges and highlight the difficulty of predicting the parts of a species’ distribution most vulnerable to threat.     

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.     

Consistent scaling of persistence time in metapopulations

Recent theory and experimental work in metapopulations and metacommunities demonstrates that long-term persistence is maximized when the rate at which individuals disperse among patches within the system is intermediate; if too low, local extinctions are more frequent than recolonizations, increasing the chance of regional-scale extinctions, and if too high, dynamics exhibit region-wide synchrony, and local extinctions occur in near unison across the region. Although common, little is known about how the size and topology of the metapopulation (metacommunity) affect this bell-shaped relationship between dispersal rate and regional persistence time. Using a suite of mathematical models, we examined the effects of dispersal, patch number, and topology on the regional persistence time when local populations are subject to demographic stochasticity. We found that the form of the relationship between regional persistence time and the number of patches is consistent across all models studied; however, the form of the relationship is distinctly different among low, intermediate, and high dispersal rates. Under low and intermediate dispersal rates, regional persistence times increase logarithmically and exponentially (respectively) with increasing numbers of patches, whereas under high dispersal, the form of the relationship depends on local dynamics. Furthermore, we demonstrate that the forms of these relationships, which give rise to the bell-shaped relationship between dispersal rate and persistence time, are a product of recolonization and the region-wide synchronization (or lack thereof) of population dynamics. Identifying such metapopulation attributes that impact extinction risk is of utmost importance for managing and conserving the earth's evermore fragmented populations.     

Amphibian Declines: Judging Stability, Persistence, and Susceptibility of Populations to Local and Global Extinctions

Extinctions are normal biological phenomena. Both mass extinctions in geological time and local extinctions in ecological time are well documented, but rates of extinction have increased in recent years—especially in vertebrates, including amphibians—as illustrated by recent reports of their population declines and range reductions. We suggest that long-term population data are necessary for rigorously evaluating the significance of the amphibian declines. Due to the physiological constraints, relatively low mobility, and site fidelity of amphibians, we suggest that many amphibian populations may be unable to recolonize areas after local extinction.     

Objectives for Multiple-Species Conservation Planning

Abstract:  The first step in conservation planning is to identify objectives. Most stated objectives for conservation, such as to maximize biodiversity outcomes, are too vague to be useful within a decision-making framework. One way to clarify the issue is to define objectives in terms of the risk of extinction for multiple species. Although the assessment of extinction risk for single species is common, few researchers have formulated an objective function that combines the extinction risks of multiple species. We sought to translate the broad goal of maximizing the viability of species into explicit objectives for use in a decision-theoretic approach to conservation planning. We formulated several objective functions based on extinction risk across many species and illustrated the differences between these objectives with simple examples. Each objective function was the mathematical representation of an approach to conservation and emphasized different levels of threat. Our objectives included minimizing the joint probability of one or more extinctions, minimizing the expected number of extinctions, and minimizing the increase in risk of extinction from the best-case scenario. With objective functions based on joint probabilities of extinction across species, any correlations in extinction probabilities had to be known or the resultant decisions were potentially misleading. Additive objectives, such as the expected number of extinctions, did not produce the same anomalies. We demonstrated that the choice of objective function is central to the decision-making process because alternative objective functions can lead to a different ranking of management options. Therefore, decision makers need to think carefully in selecting and defining their conservation goals.     

Body Size and Risk of Extinction in Australian Mammals

Abstract: The link between body size and risk of extinction has been the focus of much recent attention. For Australian terrestrial mammals this link is of particular interest because it is widely believed that species in the intermediate size range of 35–5500 g (the "critical weight range") have been the most prone to recent extinction. But the relationship between body size and extinction risk in Australian mammals has never been subject to a robust statistical analysis. Using a combination of randomization tests and phylogenetic comparative analyses, we found that Australian mammal extinctions and declines have been nonrandom with respect to body size, but we reject the hypothesis of a critical weight range at intermediate sizes. Small species appear to be the least prone to extinction, but extinctions have not been significantly clustered around intermediate sizes. Our results suggest that hypotheses linking intermediate body size with high risk of extinction in Australian mammals are misguided and that the focus of future research should shift to explaining why the smallest species are the most resistant to extinction.     

A multispecies test of source–sink indicators to prioritize habitat for declining populations

For species at risk of decline or extinction in source–sink systems, sources are an obvious target for habitat protection actions. However, the way in which source habitats are identified and prioritized can reduce the effectiveness of conservation actions. Although sources and sinks are conceptually defined using both demographic and movement criteria, simplifications are often required in systems with limited data. To assess the conservation outcomes of alternative source metrics and resulting prioritizations, we simulated population dynamics and extinction risk for 3 endangered species. Using empirically based habitat population models, we linked habitat maps with measured site‐ or habitat‐specific demographic conditions, movement abilities, and behaviors. We calculated source–sink metrics over a range of periods of data collection and prioritized consistently high‐output sources for conservation. We then tested whether prioritized patches identified the habitats that most affected persistence by removing them and measuring the population response. Conservation decisions based on different source–sink metrics and durations of data collection affected species persistence. Shorter time series obscured the ability of metrics to identify influential habitats, particularly in temporally variable and slowly declining populations. Data‐rich source–sink metrics that included both demography and movement information did not always identify the habitats with the greatest influence on extinction risk. In some declining populations, patch abundance better predicted influential habitats for short‐term regional persistence. Because source–sink metrics (i.e., births minus deaths; births and immigrations minus deaths and emigration) describe net population conditions and cancel out gross population counts, they may not adequately identify influential habitats in declining populations. For many nonequilibrium populations, new metrics that maintain the counts of individual births, deaths, and movement may provide additional insight into habitats that most influence persistence.     

Response of complex food webs to realistic extinction sequences

Although an ecosystem's response to biodiversity loss depends on the order in which species are lost, the extinction sequences generally used to explore such responses in food webs have been ecologically unrealistic. We investigate how several extinction orders affect the minimum number of secondary extinctions expected within pelagic food webs from 34 temperate freshwater lakes. An ecologically plausible extinction order is derived from the geographically nested pattern of species composition among the lakes and is corroborated by species' pH tolerances. Simulations suggest that lake communities are remarkably robust to this realistic extinction order and highly sensitive to the reverse sequence of species loss. This sensitivity is not well explained by the known sensitivity of networks to the loss of highly connected species but appears to be better explained by our observation that trophic specialists preferentially consume widely distributed species at low risk of extinction. Our results highlight an important aspect of community organization that may help to maintain biodiversity amidst changing environments.     

Rates of Movement of Threatened Bird Species between IUCN Red List Categories and toward Extinction

Abstract:  In recent centuries bird species have been deteriorating in status and becoming extinct at a rate that may be 2–3 orders of magnitude higher than in prehuman times. We examined extinction rates of bird species designated critically endangered in 1994 and the rate at which species have moved through the IUCN (World Conservation Union) Red List categories of extinction risk globally for the period 1988–2004 and regionally in Australia from 1750 to 2000. For Australia we drew on historical accounts of the extent and condition of species habitats, spread of invasive species, and changes in sighting frequencies. These data sets permitted comparison of observed rates of movement through the IUCN Red List categories with novel predictions based on the IUCN Red List criterion E, which relates to explicit extinction probabilities determined, for example, by population viability analysis. The comparison also tested whether species listed on the basis of other criteria face a similar probability of moving to a higher threat category as those listed under criterion E. For the rate at which species moved from vulnerable to endangered, there was a good match between observations and predictions, both worldwide and in Australia. Nevertheless, species have become extinct at a rate that, although historically high, is 2 (Australia) to 10 (globally) times lower than predicted. Although the extinction probability associated with the critically endangered category may be too high, the shortfall in realized extinctions can also be attributed to the beneficial impact of conservation intervention. These efforts may have reduced the number of global extinctions from 19 to 3 and substantially slowed the extinction trajectory of 33 additional critically endangered species. Our results suggest that current conservation action benefits species on the brink of extinction, but is less targeted at or has less effect on moderately threatened species.     

Conserving the abundance of nonthreatened species

Human modification of the environment is driving declines in population size and distributional extent of much of the world's biota. These declines extend to many of the most abundant and widespread species, for which proportionally small declines can result in the loss of vast numbers of individuals, biomass, and interactions. These losses could have major localized effects on ecological and cultural processes and services without elevating a species’ global extinction risk. Although most conservation effort is directed at species threatened with extinction in the very near term, the value of retaining abundance regardless of global extinction risk is justifiable based on many biodiversity or ecosystem service metrics, including cultural services, at scales from local to global. The challenges of identifying conservation priorities for widespread and abundant species include quantifying the effects of species’ abundance on services and understanding how these effects are realized as populations decline. Negative effects of population declines may be disconnected from the threat processes driving declines because of species movements and environment flows (e.g., hydrology). Conservation prioritization for these species shares greater similarity with invasive species risk assessments than extinction risk assessments because of the importance of local context and per capita effects of abundance on other species. Because conservation priorities usually focus on preventing the extinction of threatened species, the rationale and objectives for incorporating declines of nonthreatened species must be clearly articulated, going beyond extinction risk to encompass the range of likely harmful effects (e.g., secondary extinctions, loss of ecosystem services) if declines persist or are not reversed. Research should focus on characterizing the effects of local declines in species that are not threatened globally across a range of ecosystem services and quantifying the spatial distribution of these effects through the distribution of abundance. The case for conserving abundance in nonthreatened species can be made most powerfully when the costs of losing this abundance are better understood.     

A population viability analysis for the Island Fox on Santa Catalina Island,California

The island fox (Urocyon littoralis) on Santa Catalina Island is among the most imperiled species on the Channel Islands due to a recent outbreak of canine distemper virus (CDV). The western subpopulation, which was not exposed to CDV, is a crucial element in the recovery of foxes by providing a source of animals for translocation and captive breeding. Using the program VORTEX, we developed a population viability analysis for the Santa Catalina Island fox to (1) address the likelihood of population persistence, (2) estimate the current susceptibility of the population to catastrophic events, and (3) evaluate the efficacy of current restoration strategies of releasing captive bred foxes and transplanting wild animals. Overall, we found the population to be susceptible to catastrophic events; a 50% increase in mortality every 20 years was sufficient to elevate the extinction risk above 5%. Current management activities entail the transplanting of 12 juvenile foxes annually, which may reduce the viability of the western subpopulation. A minimum population size of at least 150 foxes should be maintained in each subpopulation to reduce the risk of extinction due to demographic stochasticity. Releases of translocated and captive bred animals affect the speed of recovery on the eastern half of Catalina Island, but not the probability of extinction, which is near zero under current conditions. We conducted a sensitivity analysis for demographic parameters by incrementally varying survival, fecundity and density-dependence parameters, while holding all other parameters constant. Sensitivity analyses identified mortality and mean litter size as the most sensitive parameters, while the implementation of density-dependence and environmental variation of model parameters did not seem to affect population performance. We conclude that the population of island foxes on Santa Catalina is currently at a critically low population level, but recovery of the species appears possible.     

Modern Insect Extinctions, the Neglected Majority

Abstract:  Most extinctions estimated to have occurred in the historical past, or predicted to occur in the future, are of insects. Despite this, the study of insect extinctions has been neglected. Only 70 modern insect extinctions have been documented, although thousands are estimated to have occurred. By focusing on some of the 70 documented extinctions as case studies, I considered ways in which insect extinctions may differ from those of other taxa. These case studies suggested that two types of extinction might be common for insects but rare for other taxa: extinction of narrow habitat specialists and coextinctions of affiliates with the extinctions of their hosts. Importantly, both of these forms of extinction are often ignored by conservation programs focused on vertebrates and plants. Anecdotal evidence and recent simulations suggest that many insect extinctions may have already occurred because of loss of narrow habitat specialists from restricted habitats and the loss of hosts. If we are serious about insect conservation, we need to spend more time and money documenting such extinctions. To neglect such extinctions is to ignore the majority of species that are or were in need of conservation.     

Two Hundred Years of Local Avian Extinctions in Eastern Amazonia

Local, regional, and global extinctions caused by habitat loss, degradation, and fragmentation have been widely reported for the tropics. The patterns and drivers of this loss of species are now increasingly well known in Amazonia, but there remains a significant gap in understanding of long‐term trends in species persistence and extinction in anthropogenic landscapes. Such a historical perspective is critical for understanding the status and trends of extant biodiversity as well as for identifying priorities to halt further losses. Using extensive historical data sets of specimen records and results of contemporary surveys, we searched for evidence of local extinctions of a terra firma rainforest avifauna over 200 years in a 2500 km² eastern Amazonian region around the Brazilian city of Belém. This region has the longest history of ornithological fieldwork in the entire Amazon basin and lies in the highly threatened Belém Centre of Endemism. We also compared our historically inferred extinction events with extensive data on species occurrences in a sample of catchments in a nearby municipality (Paragominas) that encompass a gradient of past forest loss. We found evidence for the possible extinction of 47 species (14% of the regional species pool) that were unreported from 1980 to 2013 (80% last recorded between 1900 and 1980). Seventeen species appear on the International Union for Conservation of Nature Red List, and many of these are large‐bodied. The species lost from the region immediately around Belém are similar to those which are currently restricted to well‐forested catchments in Paragominas. Although we anticipate the future rediscovery or recolonization of some species inferred to be extinct by our calculations, we also expect that there are likely to be additional local extinctions, not reported here, given the ongoing loss and degradation of remaining areas of native vegetation across eastern Amazonia. Doscientos Años de Extinciones Locales de Aves en la Amazonia Oriental     

Global population collapse in a superabundant migratory bird and illegal trapping in China

Persecution and overexploitation by humans are major causes of species extinctions. Rare species, often confined to small geographic ranges, are usually at highest risk, whereas extinctions of superabundant species with very large ranges are rare. The Yellow‐breasted Bunting (Emberiza aureola) used to be one of the most abundant songbirds of the Palearctic, with a very large breeding range stretching from Scandinavia to the Russian Far East. Anecdotal information about rapid population declines across the range caused concern about unsustainable trapping along the species’ migration routes. We conducted a literature review and used long‐term monitoring data from across the species’ range to model population trend and geographical patterns of extinction. The population declined by 84.3–94.7% between 1980 and 2013, and the species’ range contracted by 5000 km. Quantitative evidence from police raids suggested rampant illegal trapping of the species along its East Asian flyway in China. A population model simulating an initial harvest level of 2% of the population, and an annual increase of 0.2% during the monitoring period produced a population trajectory that matched the observed decline. We suggest that trapping strongly contributed to the decline because the consumption of Yellow‐breasted Bunting and other songbirds has increased as a result of economic growth and prosperity in East Asia. The magnitude and speed of the decline is unprecedented among birds with a comparable range size, with the exception of the Passenger Pigeon (Ectopistes migratorius), which went extinct in 1914 due to industrial‐scale hunting. Our results demonstrate the urgent need for an improved monitoring of common and widespread species’ populations, and consumption levels throughout East Asia.     

Erosion of Heterozygosity in Fluctuating Populations

Abstract: Demographic, environmental, and genetic stochasticity threaten the persistence of isolated populations. The relative importance of these intertwining factors remains unresolved, but a common view is that random demographic and environmental events will usually drive small populations to the brink of extinction before genetic deterioration poses a serious threat. To evaluate the potential importance of genetic factors, we analyzed a model linking demographic and environmental conditions to the loss of genetic diversity in isolated populations undergoing natural levels of fluctuation. Nongenetic processes—environmental stochasticity and population demography—were modeled according to a bounded diffusion process. Genetic processes were modeled by quantifying the rate of drift according to the effective population size, which was predicted from the same parameters used to describe the nongenetic processes. We combined these models to predict the heterozygosity remaining at the time of extinction, as predicted by the nongenetic portion of the model. Our model predicts that many populations will lose most or all of their neutral genetic diversity before nongenetic random events lead to extinction. Given the abundant evidence for inbreeding depression and recent evidence for elevated extinction rates of inbred populations, our findings suggest that inbreeding may be a greater general threat to population persistence than is generally recognized. Therefore, conservation biologists should not ignore the genetic component of extinction risk when assessing species endangerment and developing recovery plans.     

A Behaviorally Explicit Demographic Model Integrating Habitat Selection and Population Dynamics in California Sea Lions

Abstract: Although there has been a call for the integration of behavioral ecology and conservation biology, there are few tools currently available to achieve this integration. Explicitly including information about behavioral strategies in population viability analyses may enhance the ability of conservation biologists to understand and estimate patterns of extinction risk. Nevertheless, most behavioral‐based PVA approaches require detailed individual‐based data that are rarely available for imperiled species. We present a mechanistic approach that incorporates spatial and demographic consequences of behavioral strategies into population models used for conservation. We developed a stage‐structured matrix model that includes the costs and benefits of movement associated with 2 habitat‐selection strategies (philopatry and direct assessment). Using a life table for California sea lions (Zalophus californianus), we explored the sensitivity of model predictions to the inclusion of these behavioral parameters. Including behavioral information dramatically changed predicted population sizes, model dynamics, and the expected distribution of individuals among sites. Estimated population sizes projected in 100 years diverged up to 1 order of magnitude among scenarios that assumed different movement behavior. Scenarios also exhibited different model dynamics that ranged from stable equilibria to cycles or extinction. These results suggest that inclusion of behavioral data in viability models may improve estimates of extinction risk for imperiled species. Our approach provides a simple method for incorporating spatial and demographic consequences of behavioral strategies into population models and may be easily extended to other species and behaviors to understand the mechanisms of population dynamics for imperiled populations.     

Risk of Local Extinction of Odonata Freshwater Habitat Generalists and Specialists

Understanding the risk of a local extinction in a single population relative to the habitat requirements of a species is important in both theoretical and applied ecology. Local extinction risk depends on several factors, such as habitat requirements, range size of species, and habitat quality. We studied the local extinctions among 31 dragonfly and damselfly species from 1930 to 1975 and from 1995 to 2003 in Central Finland. We tested whether habitat specialists had a higher local extinction rate than generalist species. Approximately 30% of the local dragonfly and damselfly populations were extirpated during the 2 study periods. The size of the geographical range of the species was negatively related to extinction rate of the local populations. In contrast to our prediction, the specialist species had lower local extinction rates than the generalist species, probably because generalist species occurred in both low‐ and high‐quality habitat. Our results are consistent with source–sink theory. Riesgo de Extinción Local de Odonatos de Agua Dulce Generalistas y Especialistas de Hábitat     

Association of Coloration Mode with Population Declines and Endangerment in Australian Frogs

Abstract: Successful protection of biodiversity requires increased understanding of the ecological characteristics that predispose some species to endangerment. Theory posits that species with polymorphic or variable coloration should have larger distributions, use more diverse resources, and be less vulnerable to population declines and extinctions, compared with taxa that do not vary in color. We used information from literature on 194 species of Australian frogs to search for associations of coloration mode with ecological variables. In general, species with variable or polymorphic color patterns had larger ranges, used more habitats, were less prone to have a negative population trend, and were estimated as less vulnerable to extinction compared with nonvariable species. An association of variable coloration with lower endangerment was also evident when we controlled statistically for the effects of range size. Nonvariable coloration was not a strong predictor of endangerment, and information on several characteristics is needed to reliably identify and protect species that are prone to decline and may become threatened by extinction in the near future. Analyses based on phylogenetic‐independent contrasts did not support the hypothesis that evolutionary transitions between nonvariable and variable or polymorphic coloration have been accompanied by changes in the ecological variables we examined. Irrefutable demonstration of a role of color pattern variation in amphibian decline and in the dynamics and persistence of populations in general will require a manipulative experimental approach.     

The control of emigration and its consequences for the survival of populations

Dispersal is the key process enhancing the long-term persistence of metapopulations in heterogeneous and dynamic landscapes. However, any individual emigrating from a occupied patch also increases the risk of local population extinction. The consequences of this increase for metapopulation persistence likely depend on the control of emigration. In this paper, we present results of individual-based simulations to compare the consequences of density-independent (DIE) and density-dependent (DDE) emigration on the extinction risk of local populations and a two-patch metapopulation. (1) For completely isolated patches extinction risk increases linearly with realised emigration rates in the DIE scenario. (2) For the DDE scenario extinction risk is nearly insensitive to emigration as longs as emigration probabilities remain below ≈0.2. Survival chances are up to half an order of magnitude larger than for populations with DIE. (3) For low dispersal mortality both modes of emigration increase survival of a metapopulation by ca. one order of magnitude. (4) For high dispersal mortality only DDE can improve the global survival chances of the metapopulation. (5) With DDE individuals are only removed from a population at high population density and the risk of extinction due to demographic stochasticity is thus much smaller compared to the DIE scenario.With density-dependent emigration prospects of metapopulations survival may thus be much higher compared to a system with density-independent emigration. Consequently, the knowledge about the factors driving emigration may significantly affect our conclusions concerning the conservation status of species.