Role of Patch Size, Disease, and Movement in Rapid Extinction of Bighorn Sheep

Abstract: The controversy (  Berger 1990, 1999 ; Wehausen 1999 ) over rapid extinction in bighorn sheep ( Ovis canadensis ) has focused on population size alone as a correlate to persistence time. We report on the persistence and population performance of 24 translocated populations of bighorn sheep. Persistence in these sheep was strongly correlated with larger patch sizes, greater distance to domestic sheep, higher population growth rates, and migratory movements, as well as to larger population sizes. Persistence was also positively correlated with larger average home-range size ( p = 0.058, n = 10 translocated populations) and home-range size of rams ( p = 0.087, n = 8 translocated populations). Greater home-range size and dispersal rates of bighorn sheep were positively correlated to larger patches. We conclude that patch size and thus habitat carrying capacity, not population size per se, is the primary correlate to both population performance and persistence. Because habitat carrying capacity defines the upper limit to population size, clearly the amount of suitable habitat in a patch is ultimately linked to population size. Larger populations (250+ animals) were more likely to recover rapidly to their pre-epizootic survey number following an epizootic ( p = 0.019), although the proportion of the population dying in the epizootic also influenced the probability of recovery ( p = 0.001). Expensive management efforts to restore or increase bighorn sheep populations should focus on large habitat patches located ≥23 km from domestic sheep, and less effort should be expended on populations in isolated, small patches of habitat.     

Habitat Change and Plant Demography: Assessing the Extinction Risk of a Formerly Common Grassland Perennial

Abstract:  An important aim of conservation biology is to understand how habitat change affects the dynamics and extinction risk of populations. We used matrix models to analyze the effect of habitat degradation on the demography of the declining perennial plant Trifolium montanum in 9 calcareous grasslands in Germany over 4 years and experimentally tested the effect of grassland management. Finite population growth rates (λ) decreased with light competition, measured as leaf-area index above T. montanum plants. At unmanaged sites λ was <1 due to lower recruitment and lower survival and flowering probability of large plants. Nevertheless, in stochastic simulations, extinction of unmanaged populations of 100 flowering plants was delayed for several decades. Clipping as a management technique rapidly increased population growth because of higher survival and flowering probability of large plants in managed than in unmanaged plots. Transition-matrix simulations from these plots indicated grazing or mowing every second year would be sufficient to ensure a growth rate ≥1 if conditions stayed the same. At frequently grazed sites, the finite growth rate was approximately 1 in most populations of T. montanum . In stochastic simulations, the extinction risk of even relatively small grazed populations was low, but about half the extant populations of T. montanum in central Germany are smaller than would be sufficient for a probability of survival of >95% over 100 years. We conclude that habitat change after cessation of management strongly reduces recruitment and survival of established individuals of this perennial plant. Nevertheless, our results suggest extinction processes may take a long time in perennial plants, resulting in an extinction debt. Even if management is frequent, many remnant populations of T. montanum may be at risk because of their small size, but even a slight increase in size could considerably reduce their extinction risk.     

Importance of Habitat Quality and Landscape Connectivity for the Persistence of Endangered Natterjack Toads

Abstract: The natterjack toad (Bufo calamita) is endangered in several parts of its distribution, including Belgium, where it occurs mainly in artificial habitats. We parameterized a general model for natterjack population viability analysis (PVA) and tested its sensitivity to changes in the values of basic parameters. Then we assessed the relative efficiency of various conservation measures in 2 situations: a small isolated population and a system of 4 populations connected by rare dispersal movements. We based the population viability analysis on a stage‐structured model of natterjack population dynamics. We parameterized the model in the RAMAS GIS platform with vital rates obtained from our own field experience and from published studies. Simulated natterjack populations were highly sensitive to habitat quality (particularly pond drying), to dispersal from surrounding local populations, and to a lesser extent to values of fecundity and survival of terrestrial stages. Population trajectories were nearly insensitive to initial abundances, carrying capacities, and the frequency of extreme climatic conditions. The simulations showed that in habitats with highly ephemeral ponds, where premetamorphosis mortality was high, natterjack populations nearly always had a very high extinction risk. We also illustrated how low dispersal rates (<1 dispersing individual/generation) efficiently rescued declining local populations. Such source‐sink dynamics demonstrate that the identification and management of source populations should be a high priority.     

Demographic Viability of a Relict Population of the Critically Endangered Plant Borderea chouardii

Abstract:   In addition to human-caused changes in the environment, natural stochasticity may threaten species persistence, and its impact must be taken into account when priorities are established and management plans are designed. Borderea chouardii is a Tertiary relict at risk of extinction that occurs in only one location in the world, where the probability of human disturbance is low. Its persistence, therefore, is mainly linked to its response to natural threats such as stochasticity. Over 8 years I monitored up to 25% of this rupicolous small geophyte. The population had an unbalanced size structure and 90% failure in seed arrival at appropriate microhabitats, which suggests a problem with recruitment. I used matrix models to describe its population dynamics, conducted hand sowings, and performed stochastic simulations to investigate the effect of environmental stochasticity on population trend and viability. I modeled several scenarios to represent a variety of ecological situations, such as population reduction, episodic or persistent disease, and enhancement or decrease of recruitment. Population growth rate (λ) was never significantly different from unity over the study period. The risk of extinction was null over the next five centuries under current conditions. Increase of mortality and decrease of recruitment reduced stochastic population growth rate, but no factor except a persistent increase of 10% mortality resulted in extinction. These results are the consequence of the plant's extremely long life span (over 300 years) and low temporal variability of key vital rates. Even though hand sowing significantly increased the stochastic population growth rate, other approaches may be more important for the persistence of this species. The extremely slow capacity for recovery following disturbances renders habitat preservation essential. In addition, the founding of new populations would reduce the risk associated with habitat destruction.     

Negative relationship between dispersal distance and demography in butterfly metapopulations

Little is known about the connection between demography and dispersal in metapopulations. The meta-analysis of the population time series of five butterfly species indicated that (meta)population dynamics are driven by density-dependent factors. Inter-specific comparison reveals a significant inverse relationship between population growth rate and the magnitude of dispersal distance. As the range of dispersal distances is constrained by the patch system, dispersing individuals moving too far away would (probably) get lost. This generates selective pressures on individuals with a high dispersal propensity, but favors individuals investing more in reproduction and results in a higher (meta)population growth rate. From a conservation perspective, individuals from (meta)populations and species sacrificing dispersal for the sake of reproductive performances are most vulnerable because of their higher sensitivity to stochastic events: the temporal variation of growth rate was much higher in the two metapopulations where dispersal was limited.     

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.     

Integral projection models for finite populations in a stochastic environment

Continuous types of population structure occur when continuous variables such as body size or habitat quality affect the vital parameters of individuals. These structures can give rise to complex population dynamics and interact with environmental conditions. Here we present a model for continuously structured populations with finite size, including both demographic and environmental stochasticity in the dynamics. Using recent methods developed for discrete age-structured models we derive the demographic and environmental variance of the population growth as functions of a continuous state variable. These two parameters, together with the expected population growth rate, are used to define a one-dimensional diffusion approximation of the population dynamics. Thus, a substantial reduction in complexity is achieved as the dynamics of the complex structured model can be described by only three population parameters. We provide methods for numerical calculation of the model parameters and demonstrate the accuracy of the diffusion approximation by computer simulation of specific examples. The general modeling framework makes it possible to analyze and predict future dynamics and extinction risk of populations with various types of structure, and to explore consequences of changes in demography caused by, e.g., climate change or different management decisions. Our results are especially relevant for small populations that are often of conservation concern.     

From population sources to sieves: the matrix alters host-parasitoid source-sink structure

Field experiments that examine the impact of immigration, emigration, or landscape structure (e.g., the composition of the matrix) on the source sink dynamics of fragmented populations are scarce. Here, planthoppers (Prokelisia crocea) and egg parasitoids (Anagrus columbi) were released among host-plant patches that varied in structural (caged, isolated, or in a network of other patches) and functional (mudflat matrix that impedes dispersal vs. brome-grass matrix that facilitates dispersal) connectivity. Planthoppers and parasitoids on caged patches exhibited density-dependent growth rates, achieved high equilibrium densities, and rarely went extinct. Therefore, experimental cordgrass patches were classified as population sources. Because access to immigrants did not result in elevated population densities, source populations were not also pseudosinks, i.e., patches whose densities occur above carrying capacity due to high immigration. Planthoppers and parasitoids in open patches in mudflat had dynamics similar to those in caged patches, but went extinct in 4-5 generations in open patches in brome. Brome-embedded patches leaked emigrants at a rate that exceeded the gains from reproduction and immigration; populations of this sort are known as population sieves. For species whose suitable patches are becoming smaller and more isolated as a result of increased habitat fragmentation, emigration losses are likely to become paramount, a condition favoring the formation of population sieves. An increase in the proportion of patches that are sieves is predicted to destabilize regional population dynamics.     

Extinction Rate Estimates for Plant Populations in Revisitation Studies: Importance of Detectability

Abstract:  Many researchers have obtained extinction-rate estimates for plant populations by comparing historical and current records of occurrence. A population that is no longer found is assumed to have gone extinct. Extinction can then be related to characteristics of these populations, such as habitat type, size, or species, to test ideas about what factors may affect extinction. Such studies neglect the fact that a population may be overlooked, however, which may bias estimates of extinction rates upward. In addition, if populations are unequally detectable across groups to be compared, such as habitat type or population size, comparisons become distorted to an unknown degree. To illustrate the problem, I simulated two data sets, assuming a constant extinction rate, in which populations occurred in different habitats or habitats of different size and these factors affected their detectability. The conventional analysis implicitly assumed that detectability equalled 1 and used logistic regression to estimate extinction rates. It wrongly identified habitat and population size as factors affecting extinction risk. In contrast, with capture-recapture methods, unbiased estimates of extinction rates were recovered. I argue that capture-recapture methods should be considered more often in estimations of demographic parameters in plant populations and communities.     

Population size affects vital rates but not population growth rate of a perennial plant

Negative effects of habitat fragmentation on individual performance have been widely documented, but relatively little is known about how simultaneous effects on multiple vital rates translate into effects on population viability in long-lived species. In this study, we examined relationships between population size, individual growth, survival and reproduction, and population growth rate in the perennial plant Phyteuma spicatum. Population size positively affected the growth of seedlings, the survival of juveniles, the proportion of adults flowering, and potential seed production. Analyses with integral projection models, however, showed no relationship between population size and population growth rate. This was due to the fact that herbivores and pathogens eliminated the relationship between population size and seed production, and that population growth rate was not sensitive to changes in the vital rates that varied with population size. We conclude that effects of population size on vital rates must not translate into effects on population growth rate, and that populations of long-lived organisms may partly be able to buffer negative effects of small population size on vital rates that have a relatively small influence on population growth rate. Our study illustrates that we need to be cautious when assessing the consequences of habitat fragmentation for population viability based on effects on only one or a few vital rates.     

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     

How do dispersal costs and habitat selection influence realized population connectivity?

Despite the importance of dispersal for population connectivity, dispersal is often costly to the individual. A major impediment to understanding connectivity has been a lack of data combining the movement of individuals and their survival to reproduction in the new habitat (realized connectivity). Although mortality often occurs during dispersal (an immediate cost), in many organisms costs are paid after dispersal (deferred costs). It is unclear how such deferred costs influence the mismatch between dispersal and realized connectivity. Through a series of experiments in the field and laboratory, we estimated both direct and indirect deferred costs in a marine bryozoan (Bugula neritina). We then used the empirical data to parameterize a theoretical model in order to formalize predictions about how dispersal costs influence realized connectivity. Individuals were more likely to colonize poor-quality habitat after prolonged dispersal durations. Individuals that colonized poor-quality habitat performed poorly after colonization because of some property of the habitat (an indirect deferred cost) rather than from prolonged dispersal per se (a direct deferred cost). Our theoretical model predicted that indirect deferred costs could result in nonlinear mismatches between spatial patterns of potential and realized connectivity. The deferred costs of dispersal are likely to be crucial for determining how well patterns of dispersal reflect realized connectivity. Ignoring these deferred costs could lead to inaccurate predictions of spatial population dynamics.     

Metapopulation Extinction Risk under Spatially Autocorrelated Disturbance

Abstract:  Recent extinction models generally show that spatial aggregation of habitat reduces overall extinction risk because sites emptied by local extinction are more rapidly recolonized. We extended such an investigation to include spatial structure in the disturbance regime. A spatially explicit metapopulation model was developed with a wide range of dispersal distances. The degree of aggregation of both habitat and disturbance pattern could be varied from a random distribution, through the intermediate case of a fractal distribution, all the way to complete aggregation (single block). Increasing spatial aggregation of disturbance generally increased extinction risk. The relative risk faced by populations in different landscapes varied greatly, depending on the disturbance regime. With random disturbance, the spatial aggregation of habitat reduced extinction risk, as in earlier studies. Where disturbance was spatially autocorrelated, however, this advantage was eliminated or reversed because populations in aggregated habitats are at risk of mass extinction from coarse-scale disturbance events. The effects of spatial patterns on extinction risk tended to be reduced by long-distance dispersal. Given the high levels of spatial correlation in natural and anthropogenic disturbance processes, population vulnerability may be greatly underestimated both by classical (nonspatial) models and by those that consider spatial structure in habitat alone.     

Habitat complexity modifies post-settlement mortality and recruitment dynamics of a marine fish

For species that have an open population structure, local population size may be strongly influenced by a combination of propagule supply and post-settlement survival. While it is widely recognized that supply of larvae (or recruits) is variable and that variable recruitment may affect the relative contribution of pre- and post-settlement factors, less effort has been made to quantify how variation in the strength of post-settlement mortality (particularly density-dependent mortality) will affect the importance of processes that determine population size. In this study, I examined the effects of habitat complexity on mortality of blue rockfish (Sebastes mystinus) within nearshore reefs off central California. I first tested whether variation in habitat complexity (measured as three-dimensional complexity of rocky substrate) affected the magnitude of both density-independent and density-dependent mortality. I then used limitation analysis to quantify how variation in habitat complexity alters the relative influence of recruitment, density-independent mortality, and density-dependent mortality in determining local population size. Increased habitat complexity was associated with a reduction in both density-independent and density-dependent mortality. At low levels of habitat complexity, limitation analysis revealed that mortality was strong and recruitment had relatively little influence on population size. However, as habitat complexity increased, recruitment became more important. At the highest levels of habitat complexity, limitation by recruitment was substantial, although density-dependent mortality was ultimately the largest constraint on population size. In high-complexity habitats, population dynamics may strongly reflect variation in recruitment even though fluctuations may be dampened by density-dependent mortality. By affecting both density-independent and density-dependent mortality, variation in habitat complexity may result in qualitative changes in the dynamics of populations. These findings suggest that the relative importance of pre- vs. post-settlement factors may be determined by quantifiable habitat features, rather than ambient recruitment level alone. Because the magnitude of recruitment fluctuations can affect species coexistence and the persistence of populations, habitat-driven changes in population dynamics may have important consequences for both community structure and population viability.     

The Role of Behavior in Recent Avian Extinctions and Endangerments

Abstract: Understanding patterns of differential extinction and predicting the relative risks of extinction among extant species are among the most important problems in conservation biology. Although recent studies reveal that behavior can be a critical component in many species' extinctions or endangerments, current approaches to the problem of predicting extinction patterns largely ignore behavior. I reviewed how behavior can affect population persistence and then used recent avian extinctions and endangerments to illustrate behaviors relevant to extinction risk. Behaviors that affect population persistence can be grouped as aggregation, interspecific responses, dispersal, habitat selection, intraspecific behavior, and maladaptive behavior. Behavior that can affect extinction risk is not limited to birds; for example, in many taxonomic groups (vertebrate and invertebrate) there is evidence of socially facilitated reproduction in colonial species, Allee effects on reproductive success and survival, behavioral regulation of population size, and conspecific attraction to breeding sites. Incorporating specific behaviors into models predicting extinction probabilities and patterns should improve their predictions.     

Floater mortality within settlement areas can explain the Allee effect in breeding populations

The Allee effect (the positive relationship between population growth rate and population size) is a constraint of some animal populations at low numbers, which increases their likelihood of extinction because of a decrease in reproduction and/or survival. We were able to demonstrate that the Allee effect can be the result of a mortality increase affecting floaters (i.e. dispersing individuals able to enter as breeders in the reproductive population when a breeding territory or a potential mate – owner of a suitable breeding territory – becomes available). Previously, potential mechanisms underlying Allee effects were always related to the breeding portion of a population only. In contrast, our understanding of or solutions to population declines due to the Allee effects can reside elsewhere, away from breeding territories.     

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.     

Modeling the Effects of Habitat Fragmentation on Source and Sink Demography of Neotropical Migrant Birds

Many songbird populations in the midwestem United States are structured as a network of sources and sinks that are linked by dispersal. We used a modeling approach to examine explicitly how populations respond to incremental fragmentation of source habitat and how this response may vary depending upon two life-history attributes: fidelity to natal habitat type and reproductive strength of the source. Fragmentation of source habitat led to a predictable decline in population for both attributes examined, but the manner in which populations declined varied depending upon the reproductive strength of the source and the level of fidelity. When the source was weak and produced few excess individuals, fragmentation of source habitats resulted in a predictable and parallel population decline of adults in both the source and the sink. In this situation high fidelity to natal habitats was important for maintenance of population size and structure. Low fidelity to weak sources resulted in population extinction; populations experienced a demographic cost by dispersing from high quality source habitat to low quality sink habitat. In contrast, when the source was strong and produced many excess individuals, fragmentation of the source led to population declines in both the source and the sink, but this decline was more abrupt in sink habitats. When the source was strong and produced a large excess of individuals, nonfidelity to natal habitats had little effect on metapopulation size and structure.     

Modeling Dynamics of Habitat-Interior Bird Populations in Fragmented Landscapes

Abstract: A stochastic computer model was used to examine the effects of varying degrees of habitat fragmentation on the dynamics of a hypothetical population of forest-interior bid. The primary demographic parameter that influenced the population's dynamics was fecundity, which varied as a function of how far a birds territory was from an ecological edge. As our model landscape became more fragmented the proportion of forest habitat that was near edges increased geometrically, and the population's overall fecundity dropped as a result. The model demonstrates that impaired reproduction in a fragmented landscape is, by itself a sufficient disruption of the population's dynamics to generate population declines and shifts in distribution similar to those observed in the fragmented forests of southern Wisconsin. Without immigration of recruits from other regions where reproduction is better, habitat-interior populations in a severely fragmented landscape can become locally extinct.     

Populations in small, ephemeral habitat patches may drive dynamics in a Daphnia magna metapopulation

Migration is the key process to understand the dynamics and persistence of a metapopulation. Many metapopulation models assume a positive correlation between habitat patch size or stability and the number of emigrants. However, few empirical data exist, and habitat patch size and habitat stability may affect dispersal differently than they affect local persistence. Here, we studied the production of the migration stage (i.e., resting eggs called ephippia) of the cladoceran Daphnia magna in a metapopulation consisting of 530 rock pool habitat patches over 25 years. Earlier, the functioning of this metapopulation was explained with a Levins-type metapopulation model or with a mainland-island metapopulation model, based on local extinction and colonization data or time series data, respectively. We used pool volume, hydroperiod length, and number of desiccation events to calculate per-pool production of ephippia (i.e., migration stages). We estimated that populations in small and ephemeral habitat patches produced more than half of the 250 000 to 1 million ephippia that were produced in the metapopulation as a whole per year between 1982 and 2006. Furthermore, these small populations contributed approximately 90% of the ephippia exposed during desiccation events, while the contribution of the long-lived populations in large pools was minimal. We term this an "inverse mainland-island" type metapopulation and propose that populations in small, ephemeral habitat patches may also be the driving force for metapopulation dynamics in other systems.