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

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

Invasion versus Isolation: Trade-Offs in Managing Native Salmonids with Barriers to Upstream Movement

Abstract: Conservation biologists often face the trade‐off that increasing connectivity in fragmented landscapes to reduce extinction risk of native species can foster invasion by non‐native species that enter via the corridors created, which can then increase extinction risk. This dilemma is acute for stream fishes, especially native salmonids, because their populations are frequently relegated to fragments of headwater habitat threatened by invasion from downstream by 3 cosmopolitan non‐native salmonids. Managers often block these upstream invasions with movement barriers, but isolation of native salmonids in small headwater streams can increase the threat of local extinction. We propose a conceptual framework to address this worldwide problem that focuses on 4 main questions. First, are populations of conservation value present (considering evolutionary legacies, ecological functions, and socioeconomic benefits as distinct values)? Second, are populations vulnerable to invasion and displacement by non‐native salmonids? Third, would these populations be threatened with local extinction if isolated with barriers? And, fourth, how should management be prioritized among multiple populations? We also developed a conceptual model of the joint trade‐off of invasion and isolation threats that considers the opportunities for managers to make strategic decisions. We illustrated use of this framework in an analysis of the invasion‐isolation trade‐off for native cutthroat trout (Oncorhynchus clarkii) in 2 contrasting basins in western North America where invasion and isolation are either present and strong or farther away and apparently weak. These cases demonstrate that decisions to install or remove barriers to conserve native salmonids are often complex and depend on conservation values, environmental context (which influences the threat of invasion and isolation), and additional socioeconomic factors. Explicit analysis with tools such as those we propose can help managers make sound decisions in such complex circumstances.     

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.     

Chaos and pattern formation in a spatial tritrophic food chain

The model of Hastings and Powell describes a tritrophic food chain that exhibits chaotic dynamics. The model assumes that the populations are homogeneously mixed, so that the probability that any two individuals interact is uniform and space can be ignored. In this paper we propose a spatial version of the Hastings and Powell model in which predators seek their preys only in a finite neighborhood of their home location, breaking the mixing hypothesis. Treating both space and time as discrete variables we derive a set of coupled equations that describe the evolution of the populations at each site of the spatial domain. We show that the introduction of local predator–prey interactions result in qualitatively distinct dynamics of predator and prey populations. The evolution equations for the predators involve averages over the local density of preys, whereas the equations for the preys involve double averages, where the local density of both preys and predators appear. Our numerical simulations show that local predation also leads to spontaneous pattern formation and to qualitative changes in the global dynamics of the system. In particular, depending on the size of the predation neighborhoods, the chaotic strange attractor present in the original model of Hastings and Powell can be replaced by a stable fixed point or by an attractor of simpler topology.     

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.     

Invasive Microstegium populations consistently outperform native range populations across diverse environments

Plant species introduced into novel ranges may become invasive due to evolutionary change, phenotypic plasticity, or other biotic or abiotic mechanisms. Evolution of introduced populations could be the result of founder effects, drift, hybridization, or adaptation to local conditions, which could enhance the invasiveness of introduced species. However, understanding whether the success of invading populations is due to genetic differences between native and introduced populations may be obscured by origin x environment interactions. That is, studies conducted under a limited set of environmental conditions may show inconsistent results if native or introduced populations are differentially adapted to specific conditions. We tested for genetic differences between native and introduced populations, and for origin x environment interactions, between native (China) and introduced (U.S.) populations of the invasive annual grass Microstegium vimineum (stiltgrass) across 22 common gardens spanning a wide range of habitats and environmental conditions. On average, introduced populations produced 46% greater biomass and had 7.4% greater survival, and outperformed native range populations in every common garden. However, we found no evidence that introduced Microstegium exhibited greater phenotypic plasticity than native populations. Biomass of Microstegium was positively correlated with light and resident community richness and biomass across the common gardens. However, these relationships were equivalent for native and introduced populations, suggesting that the greater mean performance of introduced populations is not due to unequal responses to specific environmental parameters. Our data on performance of invasive and native populations suggest that post-introduction evolutionary changes may have enhanced the invasive potential of this species. Further, the ability of Microstegium to survive and grow across the wide variety of environmental conditions demonstrates that few habitats are immune to invasion.     

Local extinction of grassland plants: the landscape matrix is more important than patch attributes

Past studies of local extinctions in fragmented habitats most often tested the influence of fragment size and isolation while ignoring how differences in the surrounding landscape matrix may govern extinction. We assessed how both the spatial attributes of remnant patches (area and isolation) and landscape factors (extent of urbanization and maximum inter-fire interval) influence the persistence of native plant species in grasslands in western Victoria, Australia. Persistence was determined in 2001 by resurveying 30 remnants first surveyed in the 1980s, and correlates of extinction were assessed using Bayesian logistic regression models. On average, 26% of populations of native species became locally extinct over two decades. Area and isolation had little effect on the probability of local extinction, but urbanization and longer maximum inter-fire intervals increased extinction risk. These findings suggest that the native grasslands studied are relatively insensitive to area- and isolation-based fragmentation effects and that short-term persistence of plant populations requires the maintenance of habitat quality. The latter is strongly influenced by the landscape matrix surrounding remnant patches through changes in fire regimes and increased exogenous disturbance.     

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.     

Long-Term Population Changes of Native and Introduced Birds in the Alaka'i Swamp, Kaua'i

Abstract:  Within the last 30 years, five endemic bird species of the Alaka'i Swamp, Kaua'i, Hawai'i, have likely gone extinct. We documented population trends of the remaining avifauna in this time period to identify a common pattern in the Hawaiian Islands: decline of native species and expansion of introduced species. We conducted bird surveys over 100 km² of the Alaka'i and Kōke'e regions of Kaua'i in March–April 2000 to estimate population size, distribution, and range limits of seven native and six introduced forest birds. We compared the results with four previous surveys conducted over the last 30 years. Five of the seven native species we studied have fared well, maintaining sizeable populations (>20,000 individuals) and unchanged or increasing numbers. The endemic 'Akikiki ( Oreomystis bairdi ), however, declined from 6296 (SE ± 1374) to 1472 (SE ± 680) individuals and exhibited range contraction from 88 to 36 km². The 'I'iwi ( Vestiaria coccinea ) also experienced a decline and contraction, though not as severe. Populations of several introduced forest birds are increasing, but all species, excluding the Japanese White-eye ( Zosterops japonicus ), were at low numbers (<5,500 individuals in survey area). One introduced species, the Japanese Bush-Warbler ( Cettia diphone ) recently invaded, whereas another, the Red-billed Leiothrix ( Leiothrix lutea ), has been extirpated. Two hurricanes in the past 20 years appear to have most strongly affected nectarivores and may have contributed to the decline or extinction of several other species. Overall, native bird populations on Kaua'i have exhibited species-specific responses to limiting factors. Although most native populations appear stable, the extant native avifauna is vulnerable as a result of limited distributions and the potential for widespread habitat degradation.     

Population Persistence in Florida Torreya: Comparing Modeled Projections of a Declining Coniferous Tree

Abstract: The Florida torreya (   Torreya taxifolia ) is a coniferous tree endemic to a 35-km stretch of bluffs and ravines along the east side of the Apalachicola River in northern Florida and southern Georgia. This formerly locally abundant tree has declined as a result of disease during the 1950s and is on the U.S. endangered species list. With no seed production in the wild, this species is headed toward extinction. We conducted a survey on roughly 200 trees from 1988 to 1996 and used these data to estimate the likelihood of population persistence during the next several decades. We compared a stage-class transition matrix model ( RAMAS) and an individual-based stochastic model ( TORSIM) of growth and mortality to project future populations. Given the current lack of seed production in the wild, all models predict extinction. The question of concern is the imminence of this predicted extinction. Differing predicted times to extinction would suggest different immediate management recommendations. Both models predicted an over 90% likelihood of persistence during the next 50 years. Predictions differed in that the transition matrix model was less optimistic than the individual-based model regarding persistence. Model sensitivity analysis showed that the results were robust to significant decreases in growth and sprouting probabilities. Submodels identified different persistence likelihoods in different populations. Nonetheless, unless management of the population can facilitate maturation and seed production, extinction of this species in the wild is inevitable.     

A comparison of models for predicting population persistence

We consider a range of models that may be used to predict the future persistence of populations, particularly those based on discrete-state Markov processes. While the mathematical theory of such processes is very well-developed, they may be difficult to work with when attempting to estimate parameters or expected times to extinction. Hence, we focus on diffusion and other approximations to these models, presenting new and recent developments in parameter estimation for density dependent processes, and the calculation of extinction times for processes subject to catastrophes. We illustrate these and other methods using data from simulated and real time series. We give particular attention to a procedure, due to Ross et al. [Ross, J.V., Taimre, T., Pollett, P.K. On parameter estimation in population models, Theor. Popul. Biol., in press], for estimating the parameters of the stochastic SIS logistic model, and demonstrate ways in which these parameters may be used to estimate expected extinction times. Although the stochastic SIS logistic model is strictly density dependent and allows only for birth and death events, it nonetheless may be used to predict extinction times with some accuracy even for populations that are only weakly density dependent, or that are subject to catastrophes.     

Modeling biodiversity dynamics in countryside landscapes

The future of biodiversity hinges to a great extent on the conservation value of countryside, the growing fraction of Earth's surface heavily influenced by human activities. How many species, and which species, can persist in such landscapes (and analogous seascapes) are open questions. Here we explore two complementary theoretical frameworks to address these questions: species-area relationships and demographic models. We use the terrestrial mammal fauna of Central America to illustrate the application of both frameworks. We begin by proposing a multi-habitat species-area relationship, the countryside species-area relationship, to forecast species extinction rates. To apply it, we classify the mammal fauna by affinity to native and human-dominated habitats. We show how considering the conservation value of countryside habitats changes estimates derived from the classic species-area approach We also examine how the z value of the species-area relationship affects extinction estimates. Next, we present a framework for assessing the relative vulnerability of species to extinction in the countryside, based on the Skellam model of population dynamics. This model predicts the minimum area of contiguous native habitat required for persistence of a species, which we use as an indicator of vulnerability to habitat change. To apply the model, we use our habitat affinity classification of mammals and we estimate life-history parameters by species and habitat type. The resulting ranking of vulnerabilities is significantly correlated with the World Conservation Union (IUCN) Red List assessment.     

Hierarchical demographic approaches for assessing invasion dynamics of non-indigenous species: An example using northern snakehead (Channa argus)

Models of species’ demographic features are commonly used to understand population dynamics and inform management tactics. Hierarchical demographic models are ideal for the assessment of non-indigenous species because our knowledge of non-indigenous populations is usually limited, data on demographic traits often come from a species’ native range, these traits vary among populations, and traits are likely to vary considerably over time as species adapt to new environments. Hierarchical models readily incorporate this spatiotemporal variation in species’ demographic traits by representing demographic parameters as multi-level hierarchies. As is done for traditional non-hierarchical matrix models, sensitivity and elasticity analyses are used to evaluate the contributions of different life stages and parameters to estimates of population growth rate. We applied a hierarchical model to northern snakehead (Channa argus), a fish currently invading the eastern United States. We used a Monte Carlo approach to simulate uncertainties in the sensitivity and elasticity analyses and to project future population persistence under selected management tactics. We gathered key biological information on northern snakehead natural mortality, maturity and recruitment in its native Asian environment. We compared the model performance with and without hierarchy of parameters. Our results suggest that ignoring the hierarchy of parameters in demographic models may result in poor estimates of population size and growth and may lead to erroneous management advice. In our case, the hierarchy used multi-level distributions to simulate the heterogeneity of demographic parameters across different locations or situations. The probability that the northern snakehead population will increase and harm the native fauna is considerable. Our elasticity and prognostic analyses showed that intensive control efforts immediately prior to spawning and/or juvenile-dispersal periods would be more effective (and probably require less effort) than year-round control efforts. Our study demonstrates the importance of considering the hierarchy of parameters in estimating population growth rate and evaluating different management strategies for non-indigenous invasive species.     

Use of stochastic patch occupancy models in the California red-legged frog for Bayesian inference regarding past events and future persistence

Assessing causes of population decline is critically important to management of threatened species. Stochastic patch occupancy models (SPOMs) are popular tools for examining spatial and temporal dynamics of populations when presence–absence data in multiple habitat patches are available. We developed a Bayesian Markov chain method that extends existing SPOMs by focusing on past environmental changes that may have altered occupancy patterns prior to the beginning of data collection. Using occupancy data from 3 creeks, we applied the method to assess 2 hypothesized causes of population decline—in situ die-off and residual impact of past source population loss—in the California red-legged frog. Despite having no data for the 20–30 years between the hypothetical event leading to population decline and the first data collected, we were able to discriminate among hypotheses, finding evidence that in situ die-off increased in 2 of the creeks. Although the creeks had comparable numbers of occupied segments, owing to different extinction–colonization dynamics, our model predicted an 8-fold difference in persistence probabilities of their populations to 2030. Adding a source population led to a greater predicted persistence probability than did decreasing the in situ die-off, emphasizing that reversing the deleterious impacts of a disturbance may not be the most efficient management strategy. We expect our method will be useful for studying dynamics and evaluating management strategies of many species.     

Consumptive and nonconsumptive effects of predators on metacommunities of competing prey

Although predators affect prey both via consumption and by changing prey migration behavior, the interplay between these two effects is rarely incorporated into spatial models of predator-prey dynamics and competition among prey. We develop a model where generalist predators have consumptive effects (i.e., altering the likelihood of local prey extinction) as well as nonconsumptive effects (altering the likelihood of colonization) on spatially separated prey populations (metapopulations). We then extend this model to explore the effects of predators on competition among prey. We find that generalist predators can promote persistence of prey metapopulations by promoting prey colonization, but predators can also hasten system-wide extinction by either increasing local extinction or reducing prey migration. By altering rates of prey migration, predators in one location can exert remote control over prey dynamics in another location via predator-mediated changes in prey flux. Thus, the effect of predators may extend well beyond the proportion of patches they visit. In the context of prey metacommunities, predator-mediated shifts in prey migration and mortality can shift the competition-colonization trade-off among competing prey, leading to changes in the prey community as well as changes in the susceptibility of prey species to habitat loss. Consequently, native prey communities may be susceptible to invasion not only by exotic prey species that experience reduced amounts of mortality from resident predators, but also by exotic prey species that exhibit strong dispersal in response to generalist native predators. Ultimately, our work suggests that the consumptive and nonconsumptive effects of generalist predators may have strong, yet potentially cryptic, effects on competing prey capable of mediating coexistence, fostering invasion, and interacting with anthropogenic habitat alteration.     

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.     

Selecting for Tolerance against Pathogens and Herbivores to Enhance Success of Reintroduction and Translocation

Abstract: Some species have insufficient defenses against climate change, emerging infectious diseases, and non‐native species because they have not been exposed to these factors over their evolutionary history, and this can decrease their likelihood of persistence. Captive breeding programs are sometimes used to reintroduce individuals back into the wild; however, successful captive breeding and reintroduction can be difficult because species or populations often cannot coexist with non‐native pathogens and herbivores without artificial selection. In captive breeding programs, breeders can select for host defenses that prevent or reduce pathogen or herbivore burden (i.e., resistance) or traits that limit the effects of parasitism or herbivory on host fitness (i.e., tolerance). We propose that selection for host tolerance may enhance the success of reintroduction or translocation because tolerant hosts generally have neutral effects on introduced pathogens and herbivores. The release of resistant hosts would have detrimental effects on their natural enemies, promoting rapid evolution to circumvent the host resistance that may reduce the long‐term probability of persistence of the reintroduced or translocated species. We examined 2 case studies, one on the pathogenic amphibian chytrid fungus ( Batrachochytrium dendrobatidis [Bd]) and the other on the herbivorous cactus moth ( Cactoblastis cactorum) in the United States, where it is not native. In each case study, we provide recommendations for how captive breeders and managers could go about selecting for host tolerance. Selecting for tolerance may offer a promising tool to rescue hosts species from invasive natural enemies as well as new natural enemies associated with climate change‐induced range shifts.     

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.     

Minimise long-term loss or maximise short-term gain?: Optimal translocation strategies for threatened species

Translocation is a useful management option for conservation of threatened animal species. It can be used to increase the range of a species, augment the numbers in a critical population, or establish new populations and hence spread the risk of extinction through local catastrophes. As it is an important and expensive conservation tool, translocation management decisions must be carefully considered, with the objective of the translocation project in mind. By analysing the translocation problem within a decision-theory framework, we find optimal management decisions that are rational and transparent. We illustrate our approach using a case study of the bridled nailtail wallaby (Onychogalea fraenata). Our particular translocation question is: if we have a set number of wallabies to translocate in each time period and two translocation sites, how many wallabies should we put at each site given the state of each population to maximise the benefit to the species? We model the translocated populations with first-order Markov chain stochastic population models, and use stochastic dynamic programming to determine the optimal management decisions. We look at two sites with different growth rates – one increasing and one decreasing – and compare the optimal strategies for two different objective functions. The first is a long-term persistence objective function, which maximises the persistence of translocated populations a large number of time steps after the end of the translocation program. The second maximises total population size at the end of the translocation program. Although these objective functions are similar, they generate surprisingly different optimal translocation strategies. When maximising the long-term persistence of the translocated populations, translocation decisions are not important as long as an increasing population is established. This indicates that site quality – rather than the number and timing of translocations – primarily determines the long-term persistence of populations. When maximising total population size, the optimal strategy is to add to the increasing population unless it is above a size where it is likely to reach its carrying capacity over the planning timeframe. As translocation decisions are important in fulfilling the objective, this objective function is more useful in creating practical advice for translocation managers. The discrepancy between the optimal strategies given by the two objectives demonstrates the importance of careful consideration when specifying the goals of a project. This observation applies not only to translocation programs, but any project where clear decision-making is needed.     

Divergence in sink contributions to population persistence

Population sinks present unique conservation challenges. The loss of individuals in sinks can compromise persistence; but conversely, sinks can improve viability by improving connectivity and facilitating the recolonization of vacant sources. To assess the contribution of sinks to regional population persistence of declining populations, we simulated source–sink dynamics for 3 very different endangered species: Black‐capped Vireos (Vireo atricapilla) at Fort Hood, Texas, Ord's kangaroo rats (Dipodomys ordii) in Alberta, and Northern Spotted Owls (Strix occidentalis caurina) in the northwestern United States. We used empirical data from these case studies to parameterize spatially explicit individual‐based models. We then used the models to quantify population abundance and persistence with and without long‐term sinks. The contributions of sink habitats varied widely. Sinks were detrimental, particularly when they functioned as strong sinks with few emigrants in declining populations (e.g., Alberta's Ord's kangaroo rat) and benign in robust populations (e.g., Black‐capped Vireos) when Brown‐headed Cowbird (Molothrus ater) parasitism was controlled. Sinks, including ecological traps, were also crucial in delaying declines when there were few sources (e.g., in Black‐capped Vireo populations with no Cowbird control). Sink contributions were also nuanced. For example, sinks that supported large, variable populations were subject to greater extinction risk (e.g., Northern Spotted Owls). In each of our case studies, new context‐dependent sinks emerged, underscoring the dynamic nature of sources and sinks and the need for frequent re‐assessment. Our results imply that management actions based on assumptions that sink habitats are generally harmful or helpful risk undermining conservation efforts for declining populations.