Key Role of European Rabbits in the Conservation of the Western Mediterranean Basin Hotspot

Abstract: The Mediterranean Basin is a global hotspot of biodiversity. Hotspots are said to be experiencing a major loss of habitat, but an added risk could be the decline of some species having a special role in ecological relationships of the system. We reviewed the role of European rabbits (Oryctolagus cuniculus) as a keystone species in the Iberian Peninsula portion of the Mediterranean hotspot. Rabbits conspicuously alter plant species composition and vegetation structure through grazing and seed dispersal, which creates open areas and preserves plant species diversity. Moreover, rabbit latrines have a demonstrable effect on soil fertility and plant growth and provide new feeding resources for many invertebrate species. Rabbit burrows provide nest sites and shelter for vertebrates and invertebrates. In addition, rabbits serve as prey for a number of predators, including the critically endangered Iberian lynx (Lynx pardinus) and Spanish Imperial Eagle (Aquila adalberti). Thus, the Mediterranean ecosystem of the Iberian Peninsula should be termed “the rabbit's ecosystem.” To our knowledge, this is the first empirical support for existence of a multifunctional keystone species in a global hotspot of biodiversity. Rabbit populations have declined drastically on the Iberian Peninsula, with potential cascading effects and serious ecological and economic consequences. From this perspective, rabbit recovery is one of the biggest challenges for conservation of the Mediterranean Basin hotspot.     

Assessment of the Conservation Efforts to Prevent Extinction of the Iberian Lynx

Abstract: The Iberian lynx (Lynx pardinus) may be the first charismatic felid to become extinct in a high‐income country, despite decades of study and much data that show extinction is highly probable. The International Union for Conservation of Nature categorizes it as critically endangered; about 200 free‐ranging individuals remain in two populations in southern Spain. Conservation measures aimed at averting extirpation have been extensively undertaken with 4 of the former 10 Iberian lynx populations recorded 25 years ago. Two of the four populations have been extirpated. The number of individuals in the third population have declined by 83%, and in the fourth the probability of extirpation has increased from 34% to 95%. Major drivers of the pending extinction are the small areas to which conservation measures have been applied; lack of incorporation of evidence‐based conservation, scientific monitoring, and adaptive management into conservation efforts; a lack of continuity in recovery efforts, and distrust by conservation agencies of scientific information. In contrast to situations in which conservation and economic objectives conflict, in the case of the Iberian lynx all stakeholders desire the species to be conserved.     

Iberian Lynx in a Fragmented Landscape: Predispersal, Dispersal, and Postdispersal Habitats

Abstract: Applied conservation biology must provide solutions for the conservation of species in modern landscapes, where prime habitats are being continuously fragmented and altered and animals are restricted to small, nonviable populations. We studied habitat selection in a fragmented population of endangered Iberian lynx (   Lynx pardinus ) by examining 14 years of radiotracking data obtained from lynx trapped in two different source areas. Habitat selection was studied independently for predispersal lynx in the source areas, for dispersing individuals through the region, and for postdispersing animals, most of which settled far from their point of origin. A multivariate analysis of variance showed that habitat use differed significantly among these phases and between area of origin, but not between sexes. The habitat type most used, and best represented within home ranges, was the mediterranean scrubland. Pine plantations were also important during and after dispersal. The rest of the habitats were either avoided (open habitats) or used according to availability ( pine and eucalyptus plantations) by dispersing lynx. Differences due to lynx origin were detected only during predispersal and dispersal and were observed because animals from each area had different habitat availability. Lynx with established territories did not use areas at random. They occupied patches of mediterranean scrubland more often than would be expected from scrubland availability during predispersal; the rest of the habitats were included within home ranges less than would be expected from their availability in the landscape. Results indicate that dispersing animals may use habitats of lower quality than habitats used by resident individuals, which suggests that conservation strategies applied across regions might be a viable objective.     

Effect of Weekend Road Traffic on the Use of Space by Raptors

Abstract:  The traffic load near large cities may show dramatic cyclical changes induced by weekend tourism, and this could induce cyclical changes in the activity patterns of wildlife. We studied a 19-km-long section of a road that crossed a high-use raptor area near a large city in Spain. We observed 18 raptor species along this segment of the road, including some threatened species, such as the Spanish Imperial Eagle ( Aquila adalberti ). The number of cars increased dramatically on Saturdays and Sundays, so we assessed the effect of varying traffic loads on raptor behavior by recording all birds of prey as close or distant to the road during working days and weekend days. On weekends, the occurrence of Spanish Imperial Eagles and vultures decreased near the road. The occurrence of other species did not change between working days and weekend days. The activity decrease on weekends by Imperial Eagles and other large raptors suggests that there are weekly cycles in raptor activity. Weekly cycles in wildlife caused by human activity are a source of concern in conservation biology.     

Positive Effects on Game Species of Top Predators by Controlling Smaller Predator Populations: An Example with Lynx, Mongooses, and Rabbits

Top predators have often been persecuted because of their supposed negative effects on species of economic concern on which they feed. In some cases, however, they may actually benefit their prey through intraguild predation on other smaller predators that share the prey. In each of two representative situations, in one of which lynx were present and in the other absent, we (1) estimated gross number of rabbits taken by lynx and Egyptian mongooses (smaller predators that are themselves preyed upon by lynx; (2) simulated size-structured rabbit populations of different densities, taking into account the reproductive value of the individuals taken by predators; and (3) estimated actual rabbit densities. Numbers of rabbits taken by predators during a year were found to be between 4.8 and 9.5 times greater when lynx were not present. After a year, rabbit population growth for an initial rabbit density of 15/ha was between 12% and 22% lower when lynx were not present. For lower initial rabbit densities, the positive effect of lynx presence on rabbits was greater. Actual rabbit densities in the areas used by lynx were 2–4 times higher than in areas not used by lynx, even though these areas were similar or identical in habitat composition. These results support the suggestion that removal of top predators may sometimes have a negative effect on prey populations of human economic concern.     

Geographical gradients in diet affect population dynamics of Canada lynx

Geographical gradients in the stability of cyclic populations of herbivores and their predators may relate to the degree of specialization of predators. However, such changes are usually associated with transition from specialist to generalist predator species, rather than from geographical variation in dietary breadth of specialist predators. Canada lynx (Lynx canadensis) and snowshoe hare (Lepus americanus) populations undergo cyclic fluctuations in northern parts of their range, but cycles are either greatly attenuated or lost altogether in the southern boreal forest where prey diversity is higher. We tested the influence of prey specialization on population cycles by measuring the stable carbon and nitrogen isotope ratios in lynx and their prey, estimating the contribution of hares to lynx diet across their range, and correlating this degree of specialization to the strength of their population cycles. Hares dominated the lynx diet across their range, but specialization on hares decreased in southern and western populations. The degree of specialization correlated with cyclic signal strength indicated by spectral analysis of lynx harvest data, but overall variability of lynx harvest (the standard deviation of natural-log-transformed harvest numbers) did not change significantly with dietary specialization. Thus, as alternative prey became more important in the lynx diet, the fluctuations became decoupled from a regular cycle but did not become less variable. Our results support the hypothesis that alternative prey decrease population cycle regularity but emphasize that such changes may be driven by dietary shifts among dominant specialist predators rather than exclusively through changes in the predator community.     

Landscape evaluation in conservation: molecular sampling and habitat modeling for the Iberian lynx.

Conservation of endangered species requires comprehensive understanding of their distribution and habitat requirements, in order to implement better management strategies. Unfortunately, this understanding is often difficult to gather at the short term required by rapidly declining populations of many rare vertebrates. We present a spatial habitat modeling approach that integrates a molecular technique for species detection with landscape information to assess habitat requirements of a critically endangered mammalian carnivore, the Iberian lynx (Lynx pardinus), in a poorly known population in Spain. We formulated a set of model hypotheses for habitat selection at the spatial scale of home ranges, based on previous information on lynx requirements of space, vegetation, and prey. To obtain the required data for model selection, we designed a sampling protocol based on surveys of feces and their molecular analysis for species identification. After comparing candidate models, we selected a parsimonious one that allowed (1) reliable assessment of lynx habitat requirements at the scale of home ranges, (2) prediction of lynx distribution and potential population size, and (3) identification of landscape management priorities for habitat conservation. This model predicted that the species was more likely to occur in landscapes with a higher percentage of rocky areas and higher cover of bushes typical of mature mediterranean shrubland mosaics. Its accuracy for discriminating lynx presence was approximately 85%, indicating high predictive performance. Mapping model predictions showed that only 16% of the studied areas constitute potential habitat for lynx, even though the region is dominated by large extents of well-preserved native vegetation with low human interference. Habitat was mostly clumped in two nearby patches connected by vegetation adequate for lynx dispersal and had a capacity for 28-62 potential breeding territories. The lynx population in Sierra Morena is probably the largest persisting today, but it is still critically small for optimism about its long-term persistence. Model results suggest habitat conservation and restoration actions needed for preserving the species, including reconciliation of hunting management with preservation of mature shrubland over large areas (particularly in rocky landscapes). The approach presented here can be applied to many other species for which the ecological information needed to develop sound habitat conservation strategies is lacking.     

Possible Extinction Vortex for a Population of Iberian Lynx on the Verge of Extirpation

Abstract: Theory suggests that demographic and genetic traits deteriorate (i.e., fitness and genetic diversity decrease) when populations become small, and that such deterioration could precipitate positive feedback loops called extinction vortices. We examined whether demographic attributes and genetic traits have changed over time in one of the 2 remaining small populations of the highly endangered Iberian lynx (Lynx pardinus) in Doñana, Spain. From 1983 to 2008, we recorded nontraumatic mortality rates, litter size, offspring survival, age at territory acquisition, and sex ratio. We combined these demographic attributes with measures of inbreeding and genetic diversity at neutral loci (microsatellites) and genes subjected to selection (major histocompatibility complex). Data on demographic traits were obtained through capture and radio tracking, checking dens during breeding, track surveys, and camera trapping. For genetic analyses, we obtained blood or tissue samples from captured or necropsied individuals or from museum specimens. Over time a female‐biased sex ratio developed, age of territory acquisition decreased, mean litter size decreased, and rates of nontraumatic mortality increased, but there were no significant changes in overall mortality rates, standardized individual heterozygosity declined steadily, and allelic diversity of exon 2 of class II major histocompatibility complex DRB genes remained constant (2 allelic variants present in all individuals analyzed). Changes in sex ratio and age of territory acquisition may have resulted from demographic stochasticity, whereas changes in litter size and nontraumatic mortality may be related to observed increases in inbreeding. Concomitant deterioration of both demographic attributes and genetic traits is consistent with an extinction vortex. The co‐occurrence, with or without interaction, of demographic and genetic deterioration may explain the lack of success of conservation efforts with the Doñana population of Iberian lynx.     

Developing Classification Criteria under the U.S. Endangered Species Act: Bowhead Whales as a Case Study

Abstract: Under the U.S. Endangered Species Act, a species is classified as endangered, threatened, or recovered based on the extent to which its survival is affected by one or more of five subjective factors. A key criticism of the act is that it makes no reference to quantitative or even qualitative parameters of what constitutes "danger of extinction." Without objective standards to guide decisionmakers, classification decisions fall prey to political and social influences. We recommend the development of species-specific, status-determining criteria as a means to rationalize and expedite the listing process and reclassification decisions, independent of the requirement for delisting criteria in recovery plans. Such criteria should (1) clearly define levels of vulnerability, (2) identify gaps in information on life-history parameters, and (3) address uncertainty in existing data. As a case study, we developed preliminary criteria for bowhead whales (    Balaena mysticetus ). Thresholds for endangered and threatened status were based on World Conservation Union ( IUCN) Red List criteria and population viability analyses. Our analysis indicates that particular attention must be focused on population structure within the species to appropriately classify the degree to which one or more components of a species are vulnerable to extinction. A similar approach could be used in the classification of other species. According to our application of the IUCN criteria and those developed for similar species by Gerber and DeMaster (1999) , the Bering Sea population of bowhead whales should be delisted, whereas the other four populations of bowheads should continue to be considered endangered.     

Heterozygosity‐Fitness Correlations and Inbreeding Depression in Two Critically Endangered Mammals

The relation among inbreeding, heterozygosity, and fitness has been studied primarily among outbred populations, and little is known about these phenomena in endangered populations. Most researchers conclude that the relation between coefficient of inbreeding estimated from pedigrees and fitness traits (inbreeding‐fitness correlations) better reflects inbreeding depression than the relation between marker heterozygosity and fitness traits (heterozygosity‐fitness correlations). However, it has been suggested recently that heterozygosity‐fitness correlations should only be expected when inbreeding generates extensive identity disequilibrium (correlations in heterozygosity and homozygosity across loci throughout the genome). We tested this hypothesis in Mohor gazelle (Gazella dama mhorr) and Iberian lynx (Lynx pardinus). For Mohor gazelle, we calculated the inbreeding coefficient and measured heterozygosity at 17 microsatellite loci. For Iberian lynx, we measured heterozygosity at 36 microsatellite loci. In both species we estimated semen quality, a phenotypic trait directly related to fitness that is controlled by many loci and is affected by inbreeding depression. Both species showed evidence of extensive identity disequilibrium, and in both species heterozygosity was associated with semen quality. In the Iberian lynx the low proportion of normal sperm associated with low levels of heterozygosity was so extreme that it is likely to limit the fertility of males. In Mohor gazelle, although heterozygosity was associated with semen quality, inbreeding coefficient was not. This result suggests that when coefficient of inbreeding is calculated on the basis of a genealogy that begins after a long history of inbreeding, the coefficient of inbreeding fails to capture previous demographic information because it is a poor estimator of accumulated individual inbreeding. We conclude that among highly endangered species with extensive identity disequilibrium, examination of heterozygosity‐fitness correlations may be an effective way to detect inbreeding depression, whereas inbreeding‐fitness correlations may be poor indicators of inbreeding depression if the pedigree does not accurately reflect the history of inbreeding. Correlaciones Heterocigosidad‐ Adaptabilidad y Depresión Endogámica en Dos Especies de Mamíferos Críticamente en Peligro     

The noble cats and the big bad scavengers: effects of dominant scavengers on solitary predators

Scavenging is an important but poorly understood ecological process. Dominant scavengers can impose a selection pressure that alters the predator’s fitness, morphology, behavior, and ecology. Interactions between ursids, likely the most important dominant scavengers in the Holarctic region, and solitary felids, which are characterized by long feeding times, provide a good opportunity for studying the effects of kleptoparasitism by dominant scavengers. We analyzed the effects of scavenging by brown bears Ursus arctos on Eurasian lynx Lynx lynx and predator’s response to kleptoparasitism in a European temperate forest ecosystem. Bears found 32?% of lynx prey remains and 15?% of all biomass of large prey killed by lynx was lost to bears. In response, lynx increased their kill rate by 23?% but were able to compensate for only 59?% of the losses. The frequency of bear scavenging was strongly dependent on bear activity patterns and was highest during the lynx pregnancy and lactation period, when up to half of lynx kills were usurped by bears. We suggest that ursid scavenging, by promoting the hunting of smaller prey, may have played an important role in the evolution of the Lynx genus as well as other predators in the Holarctic. Our study indicates that prey loss to dominant scavengers is a widespread phenomenon among felids worldwide, including forest habitats. We highlight several implications of scavenging that could considerably improve our understanding of the ecology of vertebrate communities and the evolution of predators as well as benefit the future management and conservation of endangered predators.     

Four Factors Modifying the Effect of Competition on Carnivore Population Dynamics as Illustrated by African Wild Dogs

Abstract: Limitation of predator populations by prey availability and the effects of predators on prey populations are widely recognized as important ecological processes that affect carnivore conservation. Interspecific competition can also be a strong limiting factor for carnivore populations, and the effects of competition help explain why some carnivore species are prone to extinction. Competition among carnivores is unusual in some ways, so some predictions from traditional models of competition do not hold. For example, an increase in the density of prey can increase the effect of competition among carnivores, rather than weakening it. I used published data from African wild dogs (    Lycaon pictus ) to highlight four complexities that can modify the effects of competition on the population dynamics of carnivores: habitat fragmentation, counterintuitive effects of prey density, predator-prey size ratios, and habitat type.     

When conservation management becomes contraindicated: impact of food supplementation on health of endangered wildlife

Understanding the conditions that force the implementation of management actions and their efficiency is crucial for conservation of endangered species. Wildlife managers are widely and increasingly using food supplementation for such species because the potentially immediate benefits may translate into rapid conservation improvements. Supplementary feeding can also pose risks eventually promoting undesired, unexpected, subtle, or indirect, and often unnoticed, effects that are generally poorly understood. For two decades, intensive food supplementation has been used in attempting to improve the breeding productivity of the Spanish Imperial Eagle, Aquila adalberti, one of the most endangered birds of prey in the world. Here, we examined the impact of this intensive management action on nestling health, including contamination, immunodepression, and acquisition of disease agents derived from supplementation techniques and provisioned food. Contrary to management expectations, we found that fed individuals were often inadvertently "medicated" with pharmaceuticals (antibiotics and antiparasitics) contained in supplementary food (domestic rabbits). Individuals fed with medicated rabbits showed a depressed immune system and a high prevalence and richness of pathogens compared with those with no or safe supplementary feeding using non-medicated wild rabbits. A higher presence of antibiotics (fluoroquinolones) was found in sick as opposed to healthy individuals among eaglets with supplementary feeding, which points directly toward a causal effect of these drugs in disease and other health impairments. This study represents a telling example of well-meaning management strategies not based on sound scientific evidence becoming a "contraindicated" action with detrimental repercussions undermining possible beneficial effects by increasing the impact of stochastic factors on extinction risk of endangered wildlife.     

Modeling the Effect of Population Dynamics on the Impact of Rabbit Hemorrhagic Disease

Abstract:  The European wild rabbit ( Oryctolagus cuniculus ) is a staple prey species in Mediterranean ecosystems. The arrival and subsequent spread of rabbit hemorrhagic disease throughout southwestern Europe, however, has caused a decline in rabbit numbers, leading to considerable efforts to enhance wild rabbit populations, especially through habitat management. Because rabbit population dynamics depend on habitat suitability and changes in habitat structure and composition subsequent to habitat management, I evaluated the effects of population dynamics on the long-term impact of rabbit hemorrhagic disease on rabbit populations. I used an age-structured model with varying degrees of population productivity and turnover and different habitat carrying capacities, and I assumed the existence of a unique, highly pathogenic virus. My results suggest that disease impact may be highly dependent on habitat carrying capacity and rabbit population dynamics, and the model provided some insight into the current abundance of wild rabbits in different locations in southwestern Europe. The highest disease impact was estimated for populations located in habitats with low to medium carrying capacity. In contrast, disease impact was lower in high-density populations in habitats with high carrying capacity, corresponding to a lower mean age of rabbit infection and a resulting lower mortality from rabbit hemorrhagic disease. The outcomes of the model suggest that management strategies to help rabbit populations recover should be based on improving habitats to their maximum carrying capacity and increasing rabbit population productivity. In contrast, the use of strategies based on temporary increases in rabbit density, including vaccination campaigns, translocations, and temporal habitat improvements at medium carrying capacities, may increase disease impact, resulting in short-term decreases in rabbit population density.     

Native predators reduce harvest of reindeer by Sámi pastoralists

Contemporary efforts to protect biological diversity recognize the importance of sustaining traditional human livelihoods, particularly uses of the land that are compatible with intact landscapes and ecologically complete food webs. However, these efforts often confront conflicting goals. For example, conserving native predators may harm pastoralist economies because predators consume domestic livestock that sustain people. This potential conflict must be reconciled by policy, but such reconciliation requires a firm understanding of the effects of predators on the prey used by people. We used a long-term, large-scale database and Bayesian models to estimate the impacts of lynx (Lynx lynx), wolverine (Gulo gulo), and brown bear (Ursus arctos) on harvest of semi-domesticated reindeer (Rangifer tarandus) by Sami pastoralists in Sweden. The average annual harvest of reindeer averaged 25% of the population (95% credible interval = 19, 31). Annual harvest declined by 96.6 (31, 155) reindeer for each lynx family group (the surveyed segment of the lynx population) in a management unit and by 94.3 (20, 160) for each wolverine reproduction (the surveyed segment of the wolverine population). We failed to detect effects of predation by brown bear. The mechanism for effects of predation on harvest was reduced population growth rate. The rate of increase of reindeer populations declined with increasing abundance of lynx and wolverine. The density of reindeer, latitude, and weather indexed by the North Atlantic Oscillation also influenced reindeer population growth rate. We conclude that there is a biological basis for compensating the Sámi reindeer herders for predation on reindeer.     

Chemical composition of Eurasian lynx urine conveys information on reproductive state,individual identity,and urine age

In mammals, the chemical profiles of individuals are complex and variable mixtures, and animals perceive information based on variation in the overall quality of these mixtures. A variety of compounds potentially involved in chemical communication have been characterized in the urine of different felid species, but little is known about the information content of felid scent marks. In this study, we investigated whether chemical composition of Eurasian lynx Lynx lynx urine was related to sex, reproductive state, and individual identity. We further analysed if elemental sulphur in lynx urine could serve as a dietary cue or as an indicator for the freshness of a scent mark. We collected urine from captive and wild Eurasian lynx, and analysed volatile constituents of urine by means of solid phase microextraction and gas chromatography-mass spectrometry. Our results show that lynx scent profiles contain sex-specific information on reproductive state, as well as individual identity cues. Urine marks are, therefore, well-suited to fulfil a role in reproductive behaviour and social organisation of wild lynx populations. Relative sulphur content was unrelated to time since last feeding, but decreased with age of the urine sample. The influence of diet and body condition on scent profiles should be further investigated by means of experimental studies, and may shed more light on the messages encoded in carnivore scent-marks.     

Small Populations and Offspring Sex-Ratio Deviations in Eagles

Abstract:  Stochastic variation of sex ratio has long been appreciated as a potential factor driving small populations to extinction, but it is not the only source of sex-ratio bias in small populations. We examined whether some consequences of sex allocation could affect extinction risk in small populations of size-dimorphic birds such as eagles. We report variations in sex ratio at fledging from a long-term study of a declining population of Spanish Imperial Eagles ( Aquila adalberti ). Nestling sex-ratio deviation apparently was mediated by age of breeders, whereas territory quality had no obvious effect. Adult–adult pairs produced the same proportion of both sexes in high- or low-density situations, but pairs with at least one member in nonadult plumage class produced more males. As the population declined over a period of years, the proportion of breeders with immature plumage increased; consequently, the proportion of fledgling males increased. However, when population density was high, the proportion of breeders with immature plumage decreased and more female offspring were produced. This relationship between population density, composition of breeder age, and fledgling sex ratios allowed us to make predictions of extinction risk due to nonstochastic deviations of sex ratio in small, declining populations. In the study population, on the basis of the Vortex simulation results, an estimated reduction of 42.5% in predicted mean time to extinction was attributed solely to biased sex ratio.     

Use of ambiguous detections to improve estimates from species distribution models

As large carnivores recover throughout Europe, their distribution needs to be studied to determine their conservation status and assess the potential for human-carnivore conflicts. However, efficient monitoring of many large carnivore species is challenging due to their rarity, elusive behavior, and large home ranges. Their monitoring can include opportunistic sightings from citizens in addition to designed surveys. Two types of detection errors may occur in such monitoring schemes: false negatives and false positives. False-negative detections can be accounted for in species distribution models (SDMs) that deal with imperfect detection. False-positive detections, due to species misidentification, have rarely been accounted for in SDMs. Generally, researchers use ad hoc data-filtering methods to discard ambiguous observations prior to analysis. These practices may discard valuable ecological information on the distribution of a species. We investigated the costs and benefits of including data types that may include false positives rather than discarding them for SDMs of large carnivores. We used a dynamic occupancy model that simultaneously accounts for false negatives and positives to jointly analyze data that included both unambiguous detections and ambiguous detections. We used simulations to compare the performances of our model with a model fitted on unambiguous data only. We tested the 2 models in 4 scenarios in which parameters that control false-positive detections and true detections varied. We applied our model to data from the monitoring of the Eurasian lynx (Lynx lynx) in the European Alps. The addition of ambiguous detections increased the precision of parameter estimates. For the Eurasian lynx, incorporating ambiguous detections produced more precise estimates of the ecological parameters and revealed additional occupied sites in areas where the species is likely expanding. Overall, we found that ambiguous data should be considered when studying the distribution of large carnivores through the use of dynamic occupancy models that account for misidentification.     

Accounting for Uncertainty in Making Species Protection Decisions

Abstract:  Uncertainty gives rise to two decision errors in implementing the U.S. Endangered Species Act: listing species that are not in danger of extinction and delisting species that are in danger of extinction. I evaluated four methods (minimum standard, precautionary principle, minimax regret criterion, adaptive management) for deciding whether to list or delist a species when there is uncertainty about how those decisions are likely to influence survival of the species. A safe minimum standard criterion preserves some minimum amount or safe standard (population) of a species unless maintaining that amount generates unacceptable social cost. The precautionary principle favors not delisting a species when there is insufficient evidence on the efficacy of state management plans for protecting them. A minimax regret criterion selects the delisting decision that minimizes the maximum loss likely to occur under alternative ecosystem states. When the cost of making a correct decision is less than the cost of making an incorrect decision, the minimax regret criteria indicates that delisting is the optimal decision. Active adaptive management employs statistically valid experiments to test hypotheses about the likely impacts of delisting decisions. Safe minimum standard and minimax regret criterion are not compatible with the U.S. Endangered Species Act. The precautionary principle comes closest to describing how federal agencies make delisting decisions. Active adaptive management is scientifically superior to the other methods but is costly and time consuming and may not be compatible with the U.S. National Environmental Policy Act.     

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.