Human Influence on the Spatial Structure of Threatened Pacific Salmon Metapopulations

Abstract: To remain viable, populations must be resilient to both natural and human‐caused environmental changes. We evaluated anthropogenic effects on spatial connections among populations of Chinook salmon (Oncorhynchus tshawytscha) and steelhead (O. mykiss) (designated as threatened under the U.S. Endangered Species Act) in the lower Columbia and Willamette rivers. For several anthropogenic‐effects scenarios, we used graph theory to characterize the spatial relation among populations. We plotted variance in population size against connectivity among populations. In our scenarios, reduced habitat quality decreased the size of populations and hydropower dams on rivers led to the extirpation of several populations, both of which decreased connectivity. Operation of fish hatcheries increased connectivity among populations and led to patchy or panmictic populations. On the basis of our results, we believe recolonization of the upper Cowlitz River by fall and spring Chinook and winter steelhead would best restore metapopulation structure to near‐historical conditions. Extant populations that would best conserve connectivity would be those inhabiting the Molalla (spring Chinook), lower Cowlitz, or Clackamas (fall Chinook) rivers and the south Santiam (winter steelhead) and north fork Lewis rivers (summer steelhead). Populations in these rivers were putative sources; however, they were not always the most abundant or centrally located populations. This result would not have been obvious if we had not considered relations among populations in a metapopulation context. Our results suggest that dispersal rate strongly controls interactions among the populations that comprise salmon metapopulations. Thus, monitoring efforts could lead to understanding of the true rates at which wild and hatchery fish disperse. Our application of graph theory allowed us to visualize how metapopulation structure might respond to human activity. The method could be easily extended to evaluations of anthropogenic effects on other stream‐dwelling populations and communities and could help prioritize among competing conservation measures.     

Techno-Arrogance and Halfway Technologies: Salmon Hatcheries on the Pacific Coast of North America

Humankind has adopted an arrogant and ultimately self-defeating attitude toward nature that places technological mastery over nature at the forefront of our approach to many environmental problems. This "techno-arrogance" fails to recognize limitations on, and ramifications of, attempted control of nature. An example of techno-arrogance is the flawed attempt to recover Pacific salmonid fisheries through technological application in the form of hatcheries. Countless salmon stocks have declined precipitously over the last century as a result of overfishing and widespread habitat destruction. A central feature of recovery efforts has been to build many hatcheries to produce large quantities of fish to restock streams. This approach addresses the symptoms but not the causes of the declines (an example of a halfway technology), because the habitats remain largely unsuitable for salmon. There are at least six reasons why the hatchery approach will ultimately fail: (1) data demonstrate that hatcheries are not solving the problem—salmon continue to decline despite decades of hatchery production; (2) hatcheries are costly to run, and divert resources from other efforts, such as habitat restoration; (3) hatcheries are not sustainable in the long term, requiring continual input of money and energy, (4) hatcheries are a genetically unsound approach to management that can adversely affect wild populations; (5) hatchery production leads to increased harvest of declining wild populations of salmon; and (6) hatcheries conceal from the public the truth of real salmon decline. I recommend that salmonid management turn from the symptoms to the causes of decline. Overharvest and habitat destruction must be directly addressed in a major, landscape-level effort, on a scale comparable to the hatchery program, if salmonid fisheries are to remain a part of the ecological recreational, commercial and asthetic arenas in the long term.     

The Interplay between Climate Variability and Density Dependence in the Population Viability of Chinook Salmon

Abstract:  The viability of populations is influenced by driving forces such as density dependence and climate variability, but most population viability analyses (PVAs) ignore these factors because of data limitations. Additionally, simplified PVAs produce limited measures of population viability such as annual population growth rate (λ) or extinction risk. Here we developed a "mechanistic" PVA of threatened Chinook salmon ( Oncorhynchus tshawytscha ) in which, based on 40 years of detailed data, we related freshwater recruitment of juveniles to density of spawners, and third-year survival in the ocean to monthly indices of broad-scale ocean and climate conditions. Including climate variability in the model produced important effects: estimated population viability was very sensitive to assumptions of future climate conditions and the autocorrelation contained in the climate signal increased mean population abundance while increasing probability of quasi extinction. Because of the presence of density dependence in the model, however, we could not distinguish among alternative climate scenarios through mean λ values, emphasizing the importance of considering multiple measures to elucidate population viability. Our sensitivity analyses demonstrated that the importance of particular parameters varied across models and depended on which viability measure was the response variable. The density-dependent parameter associated with freshwater recruitment was consistently the most important, regardless of viability measure, suggesting that increasing juvenile carrying capacity is important for recovery.     

Evaluating Alternative Strategies for Minimizing Unintended Fitness Consequences of Cultured Individuals on Wild Populations

Artificial propagation strategies often incur selection in captivity that leads to traits that are maladaptive in the wild. For propagation programs focused on production rather than demographic contribution to wild populations, effects on wild populations can occur through unintentional escapement or the need to release individuals into natural environments for part of their life cycle. In this case, 2 alternative management strategies might reduce unintended fitness consequences on natural populations: (1) reduce selection in captivity as much as possible to reduce fitness load (keep them similar), or (2) breed a separate population to reduce captive‐wild interactions as much as possible (make them different). We quantitatively evaluate these 2 strategies with a coupled demographic–genetic model based on Pacific salmon hatcheries that incorporates a variety of relevant processes and dynamics: selection in the hatchery relative to the wild, assortative mating based on the trait under selection, and different life cycle arrangements in terms of hatchery release, density dependence, natural selection, and reproduction. Model results indicate that, if natural selection only occurs between reproduction and captive release, the similar strategy performs better. However, if natural selection occurs between captive release and reproduction, the different and similar strategies present viable alternatives to reducing unintended fitness consequences because of the greater opportunity to purge maladaptive individuals. In this case, the appropriate approach depends on the feasibility of each strategy and the demographic goal (e.g., increasing natural abundance, or ensuring that a high proportion of natural spawners are naturally produced). In addition, the fitness effects of hatchery release are much greater if hatchery release occurs before (vs. after) density‐dependent interactions. Given the logistical challenges to achieving both the similar and different strategies, evaluation of not just the preferred strategy but also the consequences of failing to achieve the desired target is critical. Evaluación de Estrategias Alternativas para Minimizar las Consecuencias No Inesperadas en la Adecuación de Individuos Criados en Cautiverio sobre Poblaciones Silvestres     

Interacting Effects of Translocation,Artificial Propagation,and Environmental Conditions on the Marine Survival of Chinook Salmon from the Columbia River,Washington, U.S.A.

Abstract: Captive rearing and translocation are often used concurrently for species conservation, yet the effects of these practices can interact and lead to unintended outcomes that may undermine species’ recovery efforts. Controls in translocation or artificial‐propagation programs are uncommon; thus, there have been few studies on the interacting effects of these actions and environmental conditions on survival. The Columbia River basin, which drains 668,000 km² of the western United States and Canada, has an extensive network of hydroelectric and other dams, which impede and slow migration of anadromous Pacific salmon (Oncorhynchus spp.) and can increase mortality rates. To mitigate for hydrosystem‐induced mortality during juvenile downriver migration, tens of millions of hatchery fish are released each year and a subset of wild‐ and hatchery‐origin juveniles are translocated downstream beyond the hydropower system. We considered how the results of these practices interact with marine environmental conditions to affect the marine survival of Chinook salmon (O. tshawytscha). We analyzed data from more than 1 million individually tagged fish from 1998 through 2006 to evaluate the probability of an individual fish returning as an adult relative to its rearing (hatchery vs. wild) and translocation histories (translocated vs. in‐river migrating fish that traveled downriver through the hydropower system) and a suite of environmental variables. Except during select periods of very low river flow, marine survival of wild translocated fish was approximately two‐thirds less than survival of wild in‐river migrating fish. For hatchery fish, however, survival was roughly two times higher for translocated fish than for in‐river migrants. Competition and predator aggregation negatively affected marine survival, and the magnitude of survival depended on rearing and translocation histories and biological and physical conditions encountered during their first few weeks of residence in the ocean. Our results highlight the importance of considering the interacting effects of translocation, artificial propagation, and environmental variables on the long‐term viability of species.     

Bayesian Decision Analysis for Evaluating Management Options to Promote Recovery of a Depleted Salmon Population

Abstract:  The endangered population of sockeye salmon (Oncorhynchus nerka) in Cultus Lake, British Columbia, Canada, migrates through commercial fishing areas along with other, much more abundant sockeye salmon populations, but it is not feasible to selectively harvest only the latter, abundant populations. This situation creates controversial trade-offs between recovery actions and economic revenue. We conducted a Bayesian decision analysis to evaluate options for recovery of Cultus Lake sockeye salmon. We used a stochastic population model that included 2 sources of uncertainty that are often omitted from such analyses: structural uncertainty in the magnitude of a potential Allee effect and implementation uncertainty (the deviation between targets and actual outcomes of management actions). Numerous state-dependent, time-independent management actions meet recovery objectives. These actions prescribe limitations on commercial harvest rates as a function of abundance of Cultus Lake sockeye salmon. We also quantified how much reduction in economic value of commercial harvests of the more abundant sockeye salmon populations would be expected for a given increase in the probability of recovery of the Cultus population. Such results illustrate how Bayesian decision analysis can rank options for dealing with conservation risks and can help inform trade-off discussions among decision makers and among groups that have competing objectives.     

Seasonal density dependence, timing of mortality, and sustainable harvesting

Birth-pulse populations are often characterized with discrete-time models, that use a single function to relate the post-breeding population size to the post-breeding size of the previous year. Recently, models of seasonal density dependence have been constructed that emphasize interactions during shorter time periods also. Here, we study two very simple forms of density-dependent mortality, that lead to Ricker and Beverton-Holt type population dynamics when viewed over the whole year. We explore the consequences of harvest timing to equilibrium population sizes under such density dependence. Whether or not individual mortality compensates for the harvested quota, the timing of harvesting has a strong impact on the sustainability of a harvesting quota. Further, we show that careless discretization of a continuous mortality scheme may seriously underestimate the reduction in population size caused by hunting and overestimate the sustainable yield. We also introduce the concept of the demographic value of an individual, which reflects the expected contribution to population size over time in the presence of density dependence. Finally, we discuss the possibility of calculating demographic values as means of optimizing harvest strategies. Here, a Pareto optimal harvest strategy will minimize the loss of demographic value from the population for a given yield.     

Cost-Effective Management Alternatives for Snake River Chinook Salmon: a Biological-Economic Synthesis

Abstract: The mandate to increase endangered salmon populations in the Columbia River Basin of North America has created a complex, controversial resource‐management issue. We constructed an integrated assessment model as a tool for analyzing biological‐economic trade‐offs in recovery of Snake River spring‐ and summer‐run chinook salmon (Oncorhynchus tshawytscha). We merged 3 frameworks: a salmon‐passage model to predict migration and survival of smolts; an age‐structured matrix model to predict long‐term population growth rates of salmon stocks; and a cost‐effectiveness analysis to determine a set of least‐cost management alternatives for achieving particular population growth rates. We assessed 6 individual salmon‐management measures and 76 management alternatives composed of one or more measures. To reflect uncertainty, results were derived for different assumptions of effectiveness of smolt transport around dams. Removal of an estuarine predator, the Caspian Tern (Sterna caspia), was cost‐effective and generally increased long‐term population growth rates regardless of transport effectiveness. Elimination of adult salmon harvest had a similar effect over a range of its cost estimates. The specific management alternatives in the cost‐effective set depended on assumptions about transport effectiveness. On the basis of recent estimates of smolt transport effectiveness, alternatives that discontinued transportation or breached dams were prevalent in the cost‐effective set, whereas alternatives that maximized transportation dominated if transport effectiveness was relatively high. More generally, the analysis eliminated 80–90% of management alternatives from the cost‐effective set. Application of our results to salmon management is limited by data availability and model assumptions, but these limitations can help guide research that addresses critical uncertainties and information. Our results thus demonstrate that linking biology and economics through integrated models can provide valuable tools for science‐based policy and management.     

Range-Wide Selection of Catchments for Pacific Salmon Conservation

Abstract:  Freshwater ecosystems are declining in quality globally, but a lack of data inhibits identification of areas valuable for conservation across national borders. We developed a biological measure of conservation value for six species of Pacific salmon ( Oncorhynchus spp.) in catchments of the northern Pacific across Canada, China, Japan, Russia, and the United States. We based the measure on abundance and life-history richness and a model-based method that filled data gaps. Catchments with high conservation value ranged from California to northern Russia and included catchments in regions that are strongly affected by human development (e.g., Puget Sound). Catchments with high conservation value were less affected by agriculture and dams than other catchments, although only 1% were within biodiversity reserves. Our set of high-value areas was largely insensitive to simulated error, although classification remained uncertain for 3% of catchments. Although salmon face many threats, we propose they will be most likely to exhibit resilience into the future if a complementary mosaic of conservation strategies can be proactively adopted in catchments with healthy salmon populations. Our analysis provides an initial map of where these catchments are likely to be located.     

Use of Linkage Disequilibrium Data to Estimate Effective Size of Hatchery and Natural Fish Populations

A primary parameter in the assessment of the viability of a population is its effective population size ( Ne ). Allozyme analysis of four groups of fishes provided data on linkage disequilibrium, which were then used to estimate Ne . The groups included hatchery samples of juvenile white seabass, Atractoscion nobilis , juvenile rainbow trout, Oncorhynchus mykiss , from the Shasta Hatchery, and juvenile chinook salmon, O. tshawytscha , from the Coleman National Fish Hatchery. The fourth sample consisted of juvenile chinook salmon from the threatened winter run in the upper Sacramento River. The groups of fish were chosen to represent different applications of the methodology to conservation of fishes. For a variety of reasons. Ne may be considerably lower than census counts of fish present in the parental populations. The Ne of the hatchery broodstock that produced the sample of juvenile white seabass was estimated to be approximately 10, although 25 adult white seabass were present in a mass spawning tank. Ne estimates for the parental populations of the Shasta and Coleman Hatchery samples were 35.8 and 132.5, respectively. The actual number of fish spawned at the Shasta Hatchery was approximately 40, whereas nearly 10,000 salmon were spawned at the Coleman Hatchery. The threatened winter run of chinook salmon had an estimated Ne of 85.5 and an approximate run size of 2000 salmon. The method of estimating effective population size from linkage disequilibrium data appears to result in realistic estimates of effective population size when adequate sample size and a sufficient number of polymorphic loci are available.     

The effect of summer drought on the predictability of local extinctions in a butterfly metapopulation

The ecological impacts of extreme climatic events on population dynamics and community composition are profound and predominantly negative. Using extensive data of an ecological model system, we tested whether predictions from ecological models remain robust when environmental conditions are outside the bounds of observation. We observed a 10-fold demographic decline of the Glanville fritillary butterfly (Melitaea cinxia) metapopulation on the Åland islands, Finland in the summer of 2018 and used climatic and satellite data to demonstrate that this year was an anomaly with low climatic water balance values and low vegetation productivity indices across Åland. Population growth rates were strongly associated with spatiotemporal variation in climatic water balance. Covariates shown previously to affect the extinction probability of local populations in this metapopulation were less informative when populations were exposed to severe drought during the summer months. Our results highlight the unpredictable responses of natural populations to extreme climatic events.     

Effects of Harvest of Nontimber Forest Products and Ecological Differences between Sites on the Demography of African Mahogany

Abstract: The demographic impacts of harvesting nontimber forest products (NTFP) have been increasingly studied because of reports of potentially unsustainable harvest. Nevertheless, our understanding of how plant demographic response to harvest is altered by variation in ecological conditions, which is critical for developing realistic sustainable‐use plans, is limited. We built matrix population models to test whether and how variation in ecological conditions affects population responses to harvest. In particular, we examined the effect of bark and foliage harvest on the demography of populations of African mahogany (Khaya senegalensis) in two contrasting ecological regions of Benin, West Africa. K. senegalensis bark and foliage harvest significantly reduced its stochastic population growth rates, but ecological differences between regions had a greater effect on population growth rates than did harvest. The effect of harvest on population growth rates (Δλ) was slightly stronger in the moist than in the drier region. Life‐table response experiments revealed that the mechanism by which harvesting reduced λ differed between ecological regions. Lowered stasis (persistence) of larger life stages lead to a reduction in λ in the drier region, whereas lowered growth of all life stages lowered λ in moist region. Potential strategies to increase population growth rates should include decreasing the proportion of individuals harvested, promoting harvester‐owned plantations of African mahogany, and increasing survival and growth by promoting no‐fire zones in gallery forests. Our results show how population responses to harvest of NTFP may be altered by ecological differences across sites and emphasize the importance of monitoring populations over the climatic range in which they occur to develop more realistic recommendations for conservation.     

Colonization of the Southern Patagonia Ocean by Exotic Chinook Salmon

Abstract:  Anadromous salmonids have been particularly successful at establishing wild populations in southern Patagonia, in contrast to their limited success elsewhere outside their native ranges. The most recent such discovery is a spawning population of Chinook salmon in the Santa Cruz River, which flows into the Atlantic Ocean from Argentina. We used mitochondrial DNA analysis to discriminate between alternative potential sources of this population and were able to discard in situ introductions of fish imported directly from California in the early twentieth century. Our results showed that the fish most likely came from Puget Sound, Washington, imported into southern Chile for salmon-ranching experiments in the 1980s. This finding provides concrete evidence of colonization of Atlantic rivers from Pacific locations. The southern Pacific and Atlantic oceans provide a favorable marine environment for the success of invading salmon. In particular, the waters associated with fjords, southern channels, and the inshore portion of the Patagonian shelf provide a rather bounded, continuous waterway for exotic anadromous salmonids, rich in diverse forage species.     

Estimating the functional form for the density dependence from life history data

Two contrasting approaches to the analysis of population dynamics are currently popular: demographic approaches where the associations between demographic rates and statistics summarizing the population dynamics are identified; and time series approaches where the associations between population dynamics, population density, and environmental covariates are investigated. In this paper, we develop an approach to combine these methods and apply it to detailed data from Soay sheep (Ovis aries). We examine how density dependence and climate contribute to fluctuations in population size via age- and sex-specific demographic rates, and how fluctuations in demographic structure influence population dynamics. Density dependence contributes most, followed by climatic variation, age structure fluctuations and interactions between density and climate. We then simplify the density-dependent, stochastic, age-structured demographic model and derive a new phenomenological time series which captures the dynamics better than previously selected functions. The simple method we develop has potential to provide substantial insight into the relative contributions of population and individual-level processes to the dynamics of populations in stochastic environments.     

Salmon, wildlife, and wine: marine-derived nutrients in human-dominated ecosystems of central California.

Pacific salmon transfer large quantities of marine-derived nutrients to adjacent forest ecosystems with profound effects on plant and wildlife production. We investigated this process for two highly modified California wine country rivers, one with consistent salmon runs (Mokelumne River) and one without (Calaveras River). Mokelumne River Chinook salmon transported biomass and N comparable to Pacific Northwest salmon streams. Calaveras River levels were much less. Scavenger numbers correlated with salmon carcass counts over time on the Mokelumne River but not the Calaveras River. Likewise, salmon carcasses were consumed significantly faster on the Mokelumne River. Native riparian vegetation as well as cultivated wine grapes adjacent to Mokelumne River spawning sites received 18-25% of foliar N from marine sources, significantly higher than vegetation along the Calaveras River. These data suggest that robust salmon runs continue to provide important ecological services with high economic value, even in impaired watersheds. Loss of Pacific salmon can not only negatively affect stream and riparian ecosystem function, but can also affect local economies where agriculture and salmon streams coexist.     

A roadmap for ladybird conservation and recovery

Ladybirds (Coleoptera: Coccinellidae) provide services that are critical to food production, and they fulfill an ecological role as a food source for predators. The richness, abundance, and distribution of ladybirds, however, are compromised by many anthropogenic threats. Meanwhile, a lack of knowledge of the conservation status of most species and the factors driving their population dynamics hinders the development and implementation of conservation strategies for ladybirds. We conducted a review of the literature on the ecology, diversity, and conservation of ladybirds to identify their key ecological threats. Ladybird populations are most affected by climate factors, landscape composition, and biological invasions. We suggest mitigating actions for ladybird conservation and recovery. Short-term actions include citizen science programs and education, protective measures for habitat recovery and threatened species, prevention of the introduction of non-native species, and the maintenance and restoration of natural areas and landscape heterogeneity. Mid-term actions involve the analysis of data from monitoring programs and insect collections to disentangle the effect of different threats to ladybird populations, understand habitat use by taxa on which there is limited knowledge, and quantify temporal trends of abundance, diversity, and biomass along a management-intensity gradient. Long-term actions include the development of a worldwide monitoring program based on standardized sampling to fill data gaps, increase explanatory power, streamline analyses, and facilitate global collaborations.     

A case study of bats and white‐nose syndrome demonstrating how to model population viability with evolutionary effects

Ecological factors generally affect population viability on rapid time scales. Traditional population viability analyses (PVA) therefore focus on alleviating ecological pressures, discounting potential evolutionary impacts on individual phenotypes. Recent studies of evolutionary rescue (ER) focus on cases in which severe, environmentally induced population bottlenecks trigger a rapid evolutionary response that can potentially reverse demographic threats. ER models have focused on shifting genetics and resulting population recovery, but no one has explored how to incorporate those findings into PVA. We integrated ER into PVA to identify the critical decision interval for evolutionary rescue (DIER) under which targeted conservation action should be applied to buffer populations undergoing ER against extinction from stochastic events and to determine the most appropriate vital rate to target to promote population recovery. We applied this model to little brown bats (Myotis lucifugus) affected by white‐nose syndrome (WNS), a fungal disease causing massive declines in several North American bat populations. Under the ER scenario, the model predicted that the DIER period for little brown bats was within 11 years of initial WNS emergence, after which they stabilized at a positive growth rate (λ = 1.05). By comparing our model results with population trajectories of multiple infected hibernacula across the WNS range, we concluded that ER is a potential explanation of observed little brown bat population trajectories across multiple hibernacula within the affected range. Our approach provides a tool that can be used by all managers to provide testable hypotheses regarding the occurrence of ER in declining populations, suggest empirical studies to better parameterize the population genetics and conservation‐relevant vital rates, and identify the DIER period during which management strategies will be most effective for species conservation.     

Relating chronic toxicity responses to population-level effects: A comparison of population-level parameters for three salmon species as a function of low-level toxicity

Standard laboratory toxicity tests assess the physiological responses of individual organisms to exposure to toxic substances under controlled conditions. Time and space restrictions often prevent the assessment of population-level responses to a toxic substance. Contaminants can affect various biological functions (e.g. growth, fecundity or behavior), which may alter different demographic traits, leading to population-level impacts. In this study, immune suppression, reproductive dysfunction and somatic growth impairment were examined using life history matrix models for coho salmon (Oncorhynchus kisutch), sockeye salmon (Oncorhynchus nerka) and chinook salmon (Oncorhynchus tshawytscha). Our intent was to gauge the relative magnitude of response to toxic effects among species and between life history stages, not provide a specific estimate of population growth rate or abundance. Effects due to immune suppression were modeled as reductions in age-specific survival. Toxic impacts on reproductive function were modeled as a 10% reduction in reproductive contribution for all reproductively mature age groups. Model runs that examined the effect of somatic growth reduction on population parameters incorporated both survival and reproductive impacts. All impacts were modeled as 10% reductions in the affected population demographic parameters. First-year survival and reproductive impacts produced similar population growth rates (λ), but resulted in different sensitivity and stable age distributions. Modeled somatic growth reduction produced additive effects on survival and reproduction. Toxic stressors producing similar changes in λ did not necessarily produce similar changes in the age distributions. Sensitivity and elasticity analyses demonstrated that changes to the first-year survival rate produced the greatest per-unit effect on λ for each species. Alteration in abundance of mature females also strongly influenced λ. Differences observed between species showed that the number of reproductive ages and time to reproductive maturity were important components for population-level responses. These results emphasize the importance of linking toxicity responses at low concentrations to the demographic traits they affect, and help to highlight the toxicity tests that are more suitable for assessing impacts on the focal species. Additionally, life history modeling is a useful tool for developing testable hypotheses regarding impacts on specific populations as well as for conducting comparisons between populations.     

Effects of Population Size on Seed Production and Germinability in an Endangered, Fragmented Grassland Plant

Abstract: Fragmentation and isolation of plant populations are thought to affect demographic processes such as seed production and cause reductions in fitness. I followed seed set over a 3-year period in eight populations of the endangered Rutidosis leptorrhynchoides (Asteraceae) that differed in population size from 13 to over 5000 flowering plants. Germinability of the resultant seed was also examined to determine whether small populations had lower fitness than large populations. Seed set was significantly associated with population size in 2 of the 3 years. Small populations (<30 flowering plants) produced significantly fewer seeds per head in 1994 and 1995 than did large populations (500 to over 5000 flowering plants), which did not differ significantly from one another. There was, however, substantial variation within populations. In 1993 seed production did not follow any simple relationship with population size, possibly because environmental stress from low rainfall had an overriding impact. Differences in seed germinability between populations were largely not evident, suggesting that this aspect of fitness has not declined substantially in small populations relative to large populations. This study suggests that nongenetic, demographic factors are of immediate importance to the persistence of small populations of R. leptorrhynchoides because of their potential impacts on seedling recruitment.     

Critically evaluating best management practices for preventing freshwater turtle extinctions

Ex situ conservation tools, such as captive breeding for reintroduction, are considered a last resort to recover threatened or endangered species, but they may also help reduce anthropogenic threats where it is difficult or impossible to address them directly. Headstarting, or captive rearing of eggs or neonate animals for subsequent release into the wild, is controversial because it treats only a symptom of a larger conservation problem; however, it may provide a mechanism to address multiple threats, particularly near population centers. We conducted a population viability analysis of Australia's most widespread freshwater turtle, Chelodina longicollis, to determine the effect of adult roadkill (death by collision with motor vehicles), which is increasing, and reduced recruitment through nest predation from introduced European red foxes (Vulpes vulpes). We also modeled management scenarios to test the effectiveness of headstarting, fox management, and measures to reduce mortality on roads. Only scenarios with headstarting from source populations eliminated all risks of extinction and allowed population growth. Small increases in adult mortality (2%) had the greatest effect on population growth and extinction risk. Where threats simultaneously affected other life‐history stages (e.g., recruitment), eliminating harvest pressures on adult females alone did not eliminate the risk of population extinction. In our models, one source population could supply enough hatchlings annually to supplement 25 other similar‐sized populations such that extinction was avoided. Based on our results, we believe headstarting should be a primary tool for managing freshwater turtles for which threats affect multiple life‐history stages. We advocate the creation of source populations for managing freshwater turtles that are greatly threatened at multiple life‐history stages, such as depredation of eggs by invasive species and adult mortality via roadkill.