Using Ecological Function to Develop Recovery Criteria for Depleted Species: Sea Otters and Kelp Forests in the Aleutian Archipelago

Abstract: Recovery criteria for depleted species or populations normally are based on demographic measures, the goal being to maintain enough individuals over a sufficiently large area to assure a socially tolerable risk of future extinction. Such demographically based recovery criteria may be insufficient to restore the functional roles of strongly interacting species. We explored the idea of developing a recovery criterion for sea otters (Enhydra lutris) in the Aleutian archipelago on the basis of their keystone role in kelp forest ecosystems. We surveyed sea otters and rocky reef habitats at 34 island‐time combinations. The system nearly always existed in either a kelp‐dominated or deforested phase state, which was predictable from sea otter density. We used a resampling analysis of these data to show that the phase state at any particular island can be determined at 95% probability of correct classification with information from as few as six sites. When sea otter population status (and thus the phase state of the kelp forest) was allowed to vary randomly among islands, just 15 islands had to be sampled to estimate the true proportion that were kelp dominated (within 10%) with 90% confidence. We conclude that kelp forest phase state is a more appropriate, sensitive, and cost‐effective measure of sea otter recovery than the more traditional demographically based metrics, and we suggest that similar approaches have broad potential utility in establishing recovery criteria for depleted populations of other functionally important species.     

Bald eagles and sea otters in the Aleutian Archipelago: indirect effects of trophic cascades

Because sea otters (Enhydra lutris) exert a wide array of direct and indirect effects on coastal marine ecosystems throughout their geographic range, we investigated the potential influence of sea otters on the ecology of Bald Eagles (Haliaeetus leucocephalus) in the Aleutian Islands, Alaska, USA. We studied the diets, productivity, and density of breeding Bald Eagles on four islands during 1993-1994 and 2000-2002, when sea otters were abundant and scarce, respectively. Bald Eagles depend on nearshore marine communities for most of their prey in this ecosystem, so we predicted that the recent decline in otter populations would have an indirect negative effect on diets and demography of Bald Eagles. Contrary to our predictions, we found no effects on density of breeding pairs on four islands from 1993-1994 to 2000-2002. In contrast, diets and diet diversity of Bald Eagles changed considerably between the two time periods, likely reflecting a change in prey availability resulting from the increase and subsequent decline in sea otter populations. The frequency of sea otter pups, rock greenling (Hexagammus lagocephalus), and smooth lumpsuckers (Aptocyclus ventricosus) in the eagle's diet declined with corresponding increases in Rock Ptarmigan (Lagopus mutus), Glaucous-winged Gulls (Larus glaucescens), Atka mackerel (Pleurogrammus monopterygius), and various species of seabirds during the period of the recent otter population decline. Breeding success and productivity of Bald Eagles also increased during this time period, which may be due to the higher nutritional quality of avian prey consumed in later years. Our results provide further evidence of the wide-ranging indirect effects of sea otter predation on nearshore marine communities and another apex predator, the Bald Eagle. Although the indirect effects of sea otters are widely known, this example is unique because the food-web pathway transcended five species and several trophic levels in linking one apex predator to another.     

Developing a Criterion for Delisting the Southern Sea Otter under the U.S. Endangered Species Act

Recent surveys of recovery plans indicate that criteria, such as population sizes, for delisting species from the U.S. Endangered Species Act (ESA) are often unrealistically low by scientific standards. We describe the delisting criterion for the threatened southern sea otter (Enhydra lutris nereis) developed by the Southern Sea Otter Recovery Team. A major oil spill is the most serious threat to this sea otter population. After extensive modeling of oil spills, the recovery team concluded that it was not scientifically defensible to develop a delisting criterion in terms of a single probability of extinction over a specified time period. Instead, the team decided to define a size at which it would consider the population endangered and to consider the population threatened as long as a major oil spill might reduce it to that size. The effective population size (N_e) for endangered status was set at 500, estimated to be about 1850 otters. Using a spill the size of the Exxon Valdez spill (250,000 bbl), the oil spill model was iterated to generate a frequency distribution of the number of sea otters contacted by oil, from which the team estimated that less than 800 otters would be killed by 90% of the simulated spills. Thus, the delisting criterion was set at 1850 + 800 = 2650 individuals. There have been several proposals to improve the Endangered Species Act by providing quantitative guidance, in the form of specific probabilities of extinction within some time frame or specific criteria like those used by the World Conservation Union as to the levels of extinction risk represented by the terms "threatened" and "endangered." Experiences of the Sea Otter Recovery Team indicate that guidelines should not be overly rigid and should allow flexibility for dealing with specific situations. The most important consideration is to appoint a recovery team that is both technically well qualified and unconstrained by pressures from management agencies.     

Evaluating mortality rates with a novel integrated framework for nonmonogamous species

The conservation of wildlife requires management based on quantitative evidence, and especially for large carnivores, unraveling cause‐specific mortalities and understanding their impact on population dynamics is crucial. Acquiring this knowledge is challenging because it is difficult to obtain robust long‐term data sets on endangered populations and, usually, data are collected through diverse sampling strategies. Integrated population models (IPMs) offer a way to integrate data generated through different processes. However, IPMs are female‐based models that cannot account for mate availability, and this feature limits their applicability to monogamous species only. We extended classical IPMs to a two‐sex framework that allows investigation of population dynamics and quantification of cause‐specific mortality rates in nonmonogamous species. We illustrated our approach by simultaneously modeling different types of data from a reintroduced, unhunted brown bear (Ursus arctos) population living in an area with a dense human population. In a population mainly driven by adult survival, we estimated that on average 11% of cubs and 61% of adults died from human‐related causes. Although the population is currently not at risk, adult survival and thus population dynamics are driven by anthropogenic mortality. Given the recent increase of human‐bear conflicts in the area, removal of individuals for management purposes and through poaching may increase, reversing the positive population growth rate. Our approach can be generalized to other species affected by cause‐specific mortality and will be useful to inform conservation decisions for other nonmonogamous species, such as most large carnivores, for which data are scarce and diverse and thus data integration is highly desirable.     

Sea otter foraging on deep-burrowing bivalves in a California coastal lagoon

Sea otter, Enhydra lutris, predation had no detectable effect on abundance and size distribution of deep-burrowing bivalve prey in the Elkhorn Slough, California, USA. Up to 23 otters were present for 6 mo of the study period (March 1984 through April 1985). This is in contrast to previous studies of sea otter predation, especially on the shallow-burrowing Pismo clam Tivela stultorum, which can be found along the wave-exposed coast near the slough. The deep-burrowing clams Tresus nuttallii and Saxidomus nuttalli made up 61% of the prey taken in the slough, and are more difficult for otters to excavate than Pismo clams. The occurrence of foraging otters was highest in an area where the two bivalve prey were extremely abundant (18 individuals m^–2). However, the otters did not selectively prey on the largest clams available within the study sight, but foraged preferentially in a patch of smaller individuals where bivalve burrow depth was restricted by the presence of a dense clay layer. This foraging strategy maximized the amount of prey biomass obtained per unit volume of sediment excavated. Our findings suggest that in soft-sediment habitats deep-burrowing bivalves may be more resistant to otter predation than shallower burrowers.     

Exploiting sea otter populations: A simulation analysis

We used a Leslie matrix population model to investigate the impact of a range of harvest rates proposed for Alaskan sea otters (Enhydra lutris). The simulation included an analysis of several population mechanisms that might be important in the natural regulation of sea otter populations or in their reactions to harvesting. Significant differences in equilibrium population levels were found between compensatory mechanisms when fixed harvest rates were applied for 25-year periods. Adult harvests set at 2 and 4% of the total population showed that new stable population levels were rapidly attained. Harvest rates of 8 and 10%, however, resulted in marked population declines in simulated harvests. This analysis demonstrates that limited harvesting can be sustained by the population and that otter population compensation responses will be a critical determinant of sustainable harvest rates of sea otter populations.     

Rapid Increases and Time‐Lagged Declines in Amphibian Occupancy after Wildfire

Climate change is expected to increase the frequency and severity of drought and wildfire. Aquatic and moisture‐sensitive species, such as amphibians, may be particularly vulnerable to these modified disturbance regimes because large wildfires often occur during extended droughts and thus may compound environmental threats. However, understanding of the effects of wildfires on amphibians in forests with long fire‐return intervals is limited. Numerous stand‐replacing wildfires have occurred since 1988 in Glacier National Park (Montana, U.S.A.), where we have conducted long‐term monitoring of amphibians. We measured responses of 3 amphibian species to fires of different sizes, severity, and age in a small geographic area with uniform management. We used data from wetlands associated with 6 wildfires that burned between 1988 and 2003 to evaluate whether burn extent and severity and interactions between wildfire and wetland isolation affected the distribution of breeding populations. We measured responses with models that accounted for imperfect detection to estimate occupancy during prefire (0–4 years) and different postfire recovery periods. For the long‐toed salamander (Ambystoma macrodactylum) and Columbia spotted frog (Rana luteiventris), occupancy was not affected for 6 years after wildfire. But 7–21 years after wildfire, occupancy for both species decreased ≥25% in areas where >50% of the forest within 500 m of wetlands burned. In contrast, occupancy of the boreal toad (Anaxyrus boreas) tripled in the 3 years after low‐elevation forests burned. This increase in occupancy was followed by a gradual decline. Our results show that accounting for magnitude of change and time lags is critical to understanding population dynamics of amphibians after large disturbances. Our results also inform understanding of the potential threat of increases in wildfire frequency or severity to amphibians in the region. Incrementos Rápidos y Declinaciones Desfasadas en la Ocupación de Anfibios Después de un Incendio     

Factors Mediating Co‐Occurrence of an Economically Valuable Introduced Fish and Its Native Frog Prey

Habitat characteristics mediate predator–prey coexistence in many ecological systems but are seldom considered in species introductions. When economically important introduced predators are stocked despite known negative impacts on native species, understanding the role of refuges, landscape configurations, and community interactions can inform habitat management plans. We measured these factors in basins with introduced trout (Salmonidae) and the Cascades frog (Rana cascadae) to determine, which are responsible for observed patterns of co‐occurrence of this economically important predator and its native prey. Large, vegetated shallows were strongly correlated to co‐occurrence, and R. cascadae larvae occur in shallower water when fish are present, presumably to escape predation. The number of nearby breeding sites of R. cascadae was also correlated to co‐occurrence, but only when the western toad (Anaxyrus boreas) was present. Because A. boreas larvae are unpalatable to fish and resemble R. cascadae, they may provide protection from trout via Batesian mimicry. Although rescue‐effect dispersal from nearby populations may maintain co‐occurrence, within‐lake factors proved more important for predicting co‐occurrence. Learning which factors allow co‐occurrence between economically important introduced species and their native prey enables managers to make better‐informed stocking decisions. Factores que Median la Co‐Ocurrencia de un Pez Introducido con Valor Económico y su Presa, una Rana Nativa     

Drivers of Seabird Population Recovery on New Zealand Islands after Predator Eradication

Eradication of introduced mammalian predators from islands has become increasingly common, with over 800 successful projects around the world. Historically, introduced predators extirpated or reduced the size of many seabird populations, changing the dynamics of entire island ecosystems. Although the primary outcome of many eradication projects is the restoration of affected seabird populations, natural population responses are rarely documented and mechanisms are poorly understood. We used a generic model of seabird colony growth to identify key predictor variables relevant to recovery or recolonization. We used generalized linear mixed models to test the importance of these variables in driving seabird population responses after predator eradication on islands around New Zealand. The most influential variable affecting recolonization of seabirds around New Zealand was the distance to a source population, with few cases of recolonization without a source population ≤25 km away. Colony growth was most affected by metapopulation status; there was little colony growth in species with a declining status. These characteristics may facilitate the prioritization of newly predator‐free islands for active management. Although we found some evidence documenting natural recovery, generally this topic was understudied. Our results suggest that in order to guide management strategies, more effort should be allocated to monitoring wildlife response after eradication. Conductores de la Recuperación de Poblaciones de Aves Marinas en Islas de Nueva Zelanda después de la Erradicación de Depredadores     

Predicting Recovery Criteria for Threatened and Endangered Plant Species on the Basis of Past Abundances and Biological Traits

Recovery plans for species listed under the U.S. Endangered Species Act are required to specify measurable criteria that can be used to determine when the species can be delisted. For the 642 listed endangered and threatened plant species that have recovery plans, we applied recursive partitioning methods to test whether the number of individuals or populations required for delisting can be predicted on the basis of distributional and biological traits, previous abundance at multiple time steps, or a combination of traits and previous abundances. We also tested listing status (threatened or endangered) and the year the recovery plan was written as predictors of recovery criteria. We analyzed separately recovery criteria that were stated as number of populations and as number of individuals (population‐based and individual‐based criteria, respectively). Previous abundances alone were relatively good predictors of population‐based recovery criteria. Fewer populations, but a greater proportion of historically known populations, were required to delist species that had few populations at listing compared with species that had more populations at listing. Previous abundances were also good predictors of individual‐based delisting criteria when models included both abundances and traits. The physiographic division in which the species occur was also a good predictor of individual‐based criteria. Our results suggest managers are relying on previous abundances and patterns of decline as guidelines for setting recovery criteria. This may be justifiable in that previous abundances inform managers of the effects of both intrinsic traits and extrinsic threats that interact and determine extinction risk. Predicción de Criterios de Recuperación para Especies de Plantas en Peligro y Amenazadas con Base en Abundancias Pasadas y Atributos Biológicos     

Effects of Forest Fragmentation on Seed Dispersal and Seedling Establishment in Ornithochorous Trees

Abstract: Habitat fragmentation increases seed dispersal limitation across the landscape and may also affect subsequent demographic stages such as seedling establishment. Thus, the development of adequate plans for forest restoration requires an understanding of mechanisms by which fragmentation hampers seed delivery to deforested areas and knowledge of how fragmentation affects the relationship between seed‐deposition patterns and seedling establishment. We evaluated the dispersal and recruitment of two bird‐dispersed, fleshy‐fruited tree species (Crataegus monogyna and Ilex aquifolium) in fragmented secondary forests of northern Spain. Forest fragmentation reduced the probability of seed deposition for both trees because of decreased availability of woody perches and fruit‐rich neighborhoods for seed dispersers, rather than because of reductions in tree cover by itself. The effects of fragmentation went beyond effects on the dispersal stage in Crataegus because seedling establishment was proportional to the quantities of bird‐dispersed seeds arriving at microsites. In contrast, postdispersal mortality in Ilex was so high that it obscured the seed‐to‐seedling transition. These results suggest that the effects of fragmentation are not necessarily consistent across stages of recruitment across species. Habitat management seeking to overcome barriers to forest recovery must include the preservation, and even the planting, of fleshy‐fruited trees in the unforested matrix as a measure to encourage frugivorous birds to enter into open and degraded areas. An integrative management strategy should also explicitly consider seed‐survival expectancies at microhabitats to preserve plant‐population dynamics and community structure in fragmented landscapes.     

The Capacity of Australia's Protected‐Area System to Represent Threatened Species

Abstract: The acquisition or designation of new protected areas is usually based on criteria for representation of different ecosystems or land‐cover classes, and it is unclear how well‐threatened species are conserved within protected‐area networks. Here, we assessed how Australia's terrestrial protected‐area system (89 million ha, 11.6% of the continent) overlaps with the geographic distributions of threatened species and compared this overlap against a model that randomly placed protected areas across the continent and a spatially efficient model that placed protected areas across the continent to maximize threatened species’ representation within the protected‐area estate. We defined the minimum area needed to conserve each species on the basis of the species’ range size. We found that although the current configuration of protected areas met targets for representation of a given percentage of species’ ranges better than a random selection of areas, 166 (12.6%) threatened species occurred entirely outside protected areas and target levels of protection were met for only 259 (19.6%) species. Critically endangered species were among those with the least protection; 12 (21.1%) species occurred entirely outside protected areas. Reptiles and plants were the most poorly represented taxonomic groups, and amphibians the best represented. Spatial prioritization analyses revealed that an efficient protected‐area system of the same size as the current protected‐area system (11.6% of the area of Australia) could meet representation targets for 1272 (93.3%) threatened species. Moreover, the results of these prioritization analyses showed that by protecting 17.8% of Australia, all threatened species could reach target levels of representation, assuming all current protected areas are retained. Although this amount of area theoretically could be protected, existing land uses and the finite resources available for conservation mean land acquisition may not be possible or even effective for the recovery of threatened species. The optimal use of resources must balance acquisition of new protected areas, where processes that threaten native species are mitigated by the change in ownership or on‐ground management jurisdiction, and management of threatened species inside and outside the existing protected‐area system.     

Actual and Potential Use of Population Viability Analyses in Recovery of Plant Species Listed under the U.S. Endangered Species Act

Use of population viability analyses (PVAs) in endangered species recovery planning has been met with both support and criticism. Previous reviews promote use of PVA for setting scientifically based, measurable, and objective recovery criteria and recommend improvements to increase the framework's utility. However, others have questioned the value of PVA models for setting recovery criteria and assert that PVAs are more appropriate for understanding relative trade‐offs between alternative management actions. We reviewed 258 final recovery plans for 642 plants listed under the U.S. Endangered Species Act to determine the number of plans that used or recommended PVA in recovery planning. We also reviewed 223 publications that describe plant PVAs to assess how these models were designed and whether those designs reflected previous recommendations for improvement of PVAs. Twenty‐four percent of listed species had recovery plans that used or recommended PVA. In publications, the typical model was a matrix population model parameterized with ≤5 years of demographic data that did not consider stochasticity, genetics, density dependence, seed banks, vegetative reproduction, dormancy, threats, or management strategies. Population growth rates for different populations of the same species or for the same population at different points in time were often statistically different or varied by >10%. Therefore, PVAs parameterized with underlying vital rates that vary to this degree may not accurately predict recovery objectives across a species’ entire distribution or over longer time scales. We assert that PVA, although an important tool as part of an adaptive‐management program, can help to determine quantitative recovery criteria only if more long‐term data sets that capture spatiotemporal variability in vital rates become available. Lacking this, there is a strong need for viable and comprehensive methods for determining quantitative, science‐based recovery criteria for endangered species with minimal data availability. Uso Actual y Potencial del Análisis de Viabilidad Poblacional para la Recuperación de Especies de Plantas Enlistadas en el Acta de Especies En Peligro de E.U.A     

Accounting for enforcement costs in the spatial allocation of marine zones

Marine fish stocks are in many cases extracted above sustainable levels, but they may be protected through restricted‐use zoning systems. The effectiveness of these systems typically depends on support from coastal fishing communities. High management costs including those of enforcement may, however, deter fishers from supporting marine management. We incorporated enforcement costs into a spatial optimization model that identified how conservation targets can be met while maximizing fishers’ revenue. Our model identified the optimal allocation of the study area among different zones: no‐take, territorial user rights for fisheries (TURFs), or open access. The analysis demonstrated that enforcing no‐take and TURF zones incurs a cost, but results in higher species abundance by preventing poaching and overfishing. We analyzed how different enforcement scenarios affected fishers’ revenue. Fisher revenue was approximately 50% higher when territorial user rights were enforced than when they were not. The model preferentially allocated area to the enforced‐TURF zone over other zones, demonstrating that the financial benefits of enforcement (derived from higher species abundance) exceeded the costs. These findings were robust to increases in enforcement costs but sensitive to changes in species’ market price. We also found that revenue under the existing zoning regime in the study area was 13–30% lower than under an optimal solution. Our results highlight the importance of accounting for both the benefits and costs of enforcement in marine conservation, particularly when incurred by fishers. Justificación de los Costos de Aplicación en la Asignación Espacial de Zonas Marinas     

Dependence of the Endangered Black‐Capped Vireo on Sustained Cowbird Management

Conservation‐reliant species depend on active management, even after surpassing recovery goals, for protection from persistent threats. Required management may include control of another species, habitat maintenance, or artificial recruitment. Sometimes, it can be difficult to determine whether sustained management is required. We used nonspatial stochastic population projection matrix simulation and a spatially explicit population model to estimate the effects of parasitism by a brood parasite, the Brown‐headed Cowbird (Moluthrus ater), on a population of endangered Black‐capped Vireos (Vireo atricapilla). We simulated parasitism as a percentage of breeding vireo pairs experiencing decreased fecundity due to cowbirds. We estimated maximum sustainable parasitism (i.e., highest percentage of parasitized vireo breeding pairs for which population growth is ≥1) with the nonspatial model under multiple scenarios designed to assess sensitivity to assumptions about population growth rate, demographic effects of parasitism, and spatial distribution of parasitism. We then used the spatially explicit model to estimate cumulative probabilities of the population falling below the population recovery target of 1000 breeding pairs for a range of parasitism rates under multiple scenarios. We constructed our models from data on vireos collected on the Fort Hood Military Reservation, Texas (U.S.A.). Estimates of maximum sustainable parasitism rates ranged from 9–12% in scenarios with a low (6%) vireo population growth rate to 49–60% in scenarios with a high (24%) growth rate. Sustained parasitism above 45–85%, depending on the scenario, would likely result in the Fort Hood Vireo population dropping below its recovery goal within the next 25 years. These estimates suggest that vireos, although tolerant of low parasitism rates, are a conservation‐reliant species dependent on cowbird management. Dependencia de Vireo atricapilla, Especie en Peligro, hacia el Manejo Sostenido de Moluthurs ater     

Ecological Trap for Desert Lizards Caused by Anthropogenic Changes in Habitat Structure that Favor Predator Activity

Abstract: Anthropogenic habitat perturbation is a major cause of population decline. A standard practice managers use to protect populations is to leave portions of natural habitat intact. We describe a case study in which, despite the use of this practice, the critically endangered lizard Acanthodactylus beershebensis was locally extirpated from both manipulated and natural patches within a mosaic landscape of an afforestation project. We hypothesized that increased structural complexity in planted patches favors avian predator activity and makes these patches less suitable for lizards due to a heightened risk of predation. Spatial rarity of natural perches (e.g., trees) in arid scrublands may hinder the ability of desert lizards to associate perches with low‐quality habitat, turning planted patches into ecological traps for such species. We erected artificial trees in a structurally simple arid habitat (similar to the way trees were planted in the afforestation project) and compared lizard population dynamics in plots with these structures and without. Survival of lizards in the plots with artificial trees was lower than survival in plots without artificial trees. Hatchlings dispersed into plots with artificial trees in a manner that indicated they perceived the quality of these plots as similar to the surrounding, unmanipulated landscape. Our results showed that local anthropogenic changes in habitat structure that seem relatively harmless may have a considerable negative effect beyond the immediate area of the perturbation because the disturbed habitat may become an ecological trap.     

Geography and Recovery under the U.S. Endangered Species Act

Abstract: The U.S. Endangered Species Act (ESA) defines an endangered species as one “at risk of extinction throughout all or a significant portion of its range.” The prevailing interpretation of this phrase, which focuses exclusively on the overall viability of listed species without regard to their geographic distribution, has led to development of listing and recovery criteria with fundamental conceptual, legal, and practical shortcomings. The ESA's concept of endangerment is broader than the biological concept of extinction risk in that the “esthetic, ecological, educational, historical, recreational, and scientific” values provided by species are not necessarily furthered by a species mere existence, but rather by a species presence across much of its former range. The concept of “significant portion of range” thus implies an additional geographic component to recovery that may enhance viability, but also offers independent benefits that Congress intended the act to achieve. Although the ESA differs from other major endangered‐species protection laws because it acknowledges the distinct contribution of geography to recovery, it resembles the “representation, resiliency, and redundancy” conservation‐planning framework commonly referenced in recovery plans. To address representation, listing and recovery standards should consider not only what proportion of its former range a species inhabits, but the types of habitats a species occupies and the ecological role it plays there. Recovery planning for formerly widely distributed species (e.g., the gray wolf [Canis lupus]) exemplifies how the geographic component implicit in the ESA's definition of endangerment should be considered in determining recovery goals through identification of ecologically significant types or niche variation within the extent of listed species, subspecies, or “distinct population segments.” By linking listing and recovery standards to niche and ecosystem concepts, the concept of ecologically significant type offers a scientific framework that promotes more coherent dialogue concerning the societal decisions surrounding recovery of endangered species.     

Resembling a Viper: Implications of Mimicry for Conservation of the Endangered Smooth Snake

The phenomenon of Batesian mimicry, where a palatable animal gains protection against predation by resembling an unpalatable model, has been a core interest of evolutionary biologists for 150 years. An extensive range of studies has focused on revealing mechanistic aspects of mimicry (shared education and generalization of predators) and the evolutionary dynamics of mimicry systems (co‐operation vs. conflict) and revealed that protective mimicry is widespread and is important for individual fitness. However, according to our knowledge, there are no case studies where mimicry theories have been applied to conservation of mimetic species. Theoretically, mimicry affects, for example, frequency dependency of predator avoidance learning and human induced mortality. We examined the case of the protected, endangered, nonvenomous smooth snake (Coronella austriaca) that mimics the nonprotected venomous adder (Vipera berus), both of which occur in the Åland archipelago, Finland. To quantify the added predation risk on smooth snakes caused by the rarity of vipers, we calculated risk estimates from experimental data. Resemblance of vipers enhances survival of smooth snakes against bird predation because many predators avoid touching venomous vipers. Mimetic resemblance is however disadvantageous against human predators, who kill venomous vipers and accidentally kill endangered, protected smooth snakes. We found that the effective population size of the adders in Åland is very low relative to its smooth snake mimic (28.93 and 41.35, respectively).Because Batesian mimicry is advantageous for the mimic only if model species exist in sufficiently high numbers, it is likely that the conservation program for smooth snakes will fail if adders continue to be destroyed. Understanding the population consequences of mimetic species may be crucial to the success of endangered species conservation. We suggest that when a Batesian mimic requires protection, conservation planners should not ignore the model species (or co‐mimic in Mullerian mimicry rings) even if it is not itself endangered. Implications of mimicry for Conservation of the endangered smooth snake     

Conservation Strategies for Species Affected by Apparent Competition

Apparent competition is an indirect interaction between 2 or more prey species through a shared predator, and it is increasingly recognized as a mechanism of the decline and extinction of many species. Through case studies, we evaluated the effectiveness of 4 management strategies for species affected by apparent competition: predator control, reduction in the abundances of alternate prey, simultaneous control of predators and alternate prey, and no active management of predators or alternate prey. Solely reducing predator abundances rapidly increased abundances of alternate and rare prey, but observed increases are likely short‐lived due to fast increases in predator abundance following the cessation of control efforts. Substantial reductions of an abundant alternate prey resulted in increased predation on endangered huemul (Hippocamelus bisulcus) deer in Chilean Patagonia, which highlights potential risks associated with solely reducing alternate prey species. Simultaneous removal of predators and alternate prey increased survival of island foxes (Urocyon littoralis) in California (U.S.A.) above a threshold required for population recovery. In the absence of active management, populations of rare woodland caribou (Rangifer tarandus caribou) continued to decline in British Columbia, Canada. On the basis of the cases we examined, we suggest the simultaneous control of predators and alternate prey is the management strategy most likely to increase abundances and probabilities of persistence of rare prey over the long term. Knowing the mechanisms driving changes in species’ abundances before implementing any management intervention is critical. We suggest scientists can best contribute to the conservation of species affected by apparent competition by clearly communicating the biological and demographic forces at play to policy makers responsible for the implementation of proposed management actions. Estrategias de Conservación para Especies Afectadas por Competencia Aparente     

Using empirical models of species colonization under multiple threatening processes to identify complementary threat‐mitigation strategies

Approaches to prioritize conservation actions are gaining popularity. However, limited empirical evidence exists on which species might benefit most from threat mitigation and on what combination of threats, if mitigated simultaneously, would result in the best outcomes for biodiversity. We devised a way to prioritize threat mitigation at a regional scale with empirical evidence based on predicted changes to population dynamics—information that is lacking in most threat‐management prioritization frameworks that rely on expert elicitation. We used dynamic occupancy models to investigate the effects of multiple threats (tree cover, grazing, and presence of an hyperaggressive competitor, the Noisy Miner (Manorina melanocephala) on bird‐population dynamics in an endangered woodland community in southeastern Australia. The 3 threatening processes had different effects on different species. We used predicted patch‐colonization probabilities to estimate the benefit to each species of removing one or more threats. We then determined the complementary set of threat‐mitigation strategies that maximized colonization of all species while ensuring that redundant actions with little benefit were avoided. The single action that resulted in the highest colonization was increasing tree cover, which increased patch colonization by 5% and 11% on average across all species and for declining species, respectively. Combining Noisy Miner control with increasing tree cover increased species colonization by 10% and 19% on average for all species and for declining species respectively, and was a higher priority than changing grazing regimes. Guidance for prioritizing threat mitigation is critical in the face of cumulative threatening processes. By incorporating population dynamics in prioritization of threat management, our approach helps ensure funding is not wasted on ineffective management programs that target the wrong threats or species.