Selfish-herd behaviour of sheep under threat

Flocking is a striking example of collective behaviour that is found in insect swarms, fish schools and mammal herds [1]. A major factor in the evolution of flocking behaviour is thought to be predation, whereby larger and/or more cohesive groups are better at detecting predators (as, for example, in the 'many eyes theory'), and diluting the effects of predators (as in the 'selfish-herd theory') than are individuals in smaller and/or dispersed groups [2]. The former theory assumes that information (passively or actively transferred) can be disseminated more effectively in larger/cohesive groups, while the latter assumes that there are spatial benefits to individuals in a large group, since individuals can alter their spatial position relative to their group-mates and any potential predator, thus reducing their predation risk [3]. We used global positioning system (GPS) data to characterise the response of a group of 'prey' animals (a flock of sheep) to an approaching 'predator' (a herding dog). Analyses of relative sheep movement trajectories showed that sheep exhibit a strong attraction towards the centre of the flock under threat, a pattern that we could re-create using a simple model. These results support the long-standing assertion that individuals can respond to potential danger by moving towards the centre of a fleeing group [2].     

Assessing the roles of wolves and dogs in livestock predation with suggestions for mitigating human–wildlife conflict and conservation of wolves

Predation on livestock is a cause of serious and long-lasting conflict between farmers and wildlife, promoting negative public attitudes and endangering conservation of large carnivores. However, while large carnivores, especially the grey wolf (Canis lupus), are often blamed for killing sheep and other farm animals, free-ranging dogs may also act as predators. To develop appropriate measures for livestock protection, reliable methods for identifying predator species are critical. Identification of predators from visual examination of livestock wounds can be ambiguous and genetic analysis is strongly preferable for accurate species determination. To estimate the proportion of wolves and dogs implicated in sheep predation, we developed a sensitive genetic assay to distinguish between wolves and domestic dogs. A total of 183 predator saliva samples collected from killed sheep in Estonia were analysed. The assay identified the predator species in 143 cases (78%). Sheep were most often killed by wolves (81%); however, predation by dogs was substantial (15%). We compared the molecular results with field observations conducted by local environmental officials and recorded some disagreement, with the latter underestimating the role of dogs. As predator saliva samples collected from prey are often of poor quality, we suggest using mitochondrial DNA as a primary tool to maximise the number of successfully analysed samples. We also suggest adopting forensic DNA analysis more widely in livestock predation assessments as a legislative measure since misidentification that is biased against wolves can be counterproductive for conservation by enhancing conflict with society and leading to increased culling and poaching.     

Differential vulnerability of lambs to coyote predation

Coyote-lamb interactions in 65-ha pastures were observed during a study of Komondor dogs guarding sheep. Vulnerability of lambs to coyote predation was related to total time lambs were exposed to coyotes in the enclosures and total time present in the flock. Lambs were most vulnerable during the first two weeks of exposure (P < 0.001, X² = 49.3), and were somewhat less vulnerable in the next 4 weeks of exposure. If the lambs survived 6 weeks of exposure, their chances of succumbing to predation were small. Lambs were most vulnerable if they were added to an established flock. Orphan lambs were more vulnerable than ewe-reared lambs. Even though the coyotes in this study were persistent in attacking lambs, their success was limited for at least 3 reasons. First, the lambs sometimes refused to run from the coyotes. In addition, they occasionally actively defended themselves by foot-stomping, butting and approaching the coyote. Finally, no lambs were killed while in close proximity to a guardian dog. Based on this study, predation might best be reduced by minimizing introductions of new lambs, and especially orphan lambs, to established flocks, and by using a guardian dog, if possible.     

Local prey vulnerability increases with multiple attacks by a predator

Theory and empirical evidence suggest that predator activity makes prey more wary and less vulnerable to predation. However if at least some prey in the population are energetically or spatially constrained, then predators may eventually increase local prey vulnerability because of the cumulative costs of anti‐predation behaviour. We tested whether repeated attacks by a predator might increase prey vulnerability in a system where redshanks on a saltmarsh are attacked regularly by sparrowhawks from adjacent woodland. Cumulative attack number led to a reduction in redshank numbers and flock size (but had no effect on how close redshanks fed to predator‐concealing cover) because some redshanks moved to safer but less profitable habitats, leaving smaller flocks on the saltmarsh. This effect held even though numbers of redshank on the saltmarsh increased with time of day. As a result of the change in flock size, predicted attack‐success increased up to 1.6‐fold for the sparrowhawk, while individual risk of capture for the redshank increased up to 4.5‐fold among those individuals remaining on the saltmarsh. The effect did not arise simply because hawks were more likely to attack smaller flocks because attack rate was not dependent on flock size or abundance. Our data demonstrate that when some individual prey are constrained in their ability to feed on alternative, safer foraging sites, their vulnerability to predation increases as predator attacks accumulate, although those, presumably better quality individuals that leave the immediate risky area will have lower vulnerability, so that the mean vulnerability across the entire population may not have changed substantially. This suggests that the selective benefits of multiple low‐cost attacks by predators on prey could potentially lead to 1) locally heightened trait‐mediated interactions, 2) locally reduced interference among competing predators, and 3) the evolution of active prey manipulation by predators.     

Domestic Dogs in Rural Communities around Protected Areas: Conservation Problem or Conflict Solution?

Although domestic dogs play many important roles in rural households, they can also be an important threat to the conservation of wild vertebrates due to predation, competition and transmission of infectious diseases. An increasing number of studies have addressed the impact of dogs on wildlife but have tended to ignore the motivations and attitudes of the humans who keep these dogs and how the function of dogs might influence dog-wildlife interactions. To determine whether the function of domestic dogs in rural communities influences their interactions with wildlife, we conducted surveys in rural areas surrounding protected lands in the Valdivian Temperate Forests of Chile. Sixty percent of farm animal owners reported the use of dogs as one of the primary means of protecting livestock from predators. The probability of dog–wild carnivore interactions was significantly associated with the raising of poultry. In contrast, dog–wild prey interactions were not associated with livestock presence but had a significant association with poor quality diet as observed in previous studies. Dog owners reported that they actively encouraged the dogs to chase off predators, accounting for 25–75% of the dog–wild carnivore interactions observed, depending on the predator species. Humans controlled the dog population by killing pups and unwanted individuals resulting in few additions to the dog population through breeding; the importation of predominantly male dogs from urban areas resulted in a sex ratios highly dominated by males. These results indicate that dog interactions with wildlife are related to the role of the dog in the household and are directly influenced by their owners. To avoid conflict with local communities in conservation areas, it is important to develop strategies for managing dogs that balance conservation needs with the roles that dogs play in these rural households.     

Nest predators of woodland open-nesting songbirds in central Europe

I used time-lapse videotaping to identify predators of open songbird nests in fragmented deciduous woodland (nine plots, 2–10 ha each) in the Czech Republic from 2002 to 2006. I documented 22 species of predators at 171 nests of 13 species (mainly Blackcap Sylvia atricapilla , Song Thrush Turdus philomelos , Common Blackbird Turdus merula , Yellowhammer Emberiza citrinella and Chaffinch Fringilla coelebs ). The main predators were Pine Marten Martes martes (37% of 178 predation events), Jay Garrulus glandarius (29%), Buzzard Buteo buteo (7%) and Great Spotted Woodpecker Dendrocopos major (7%); mammals accounted for 48% of total predation. At least 3% of nests were depredated by multiple predators. In spite of their local abundance, Hooded Crows Corvus cornix did not present a serious threat for shrub nesting songbirds (< 1% of total predation). No predation by mice was recorded, suggesting that their importance has been overestimated in artificial nest studies. The proportional species composition of predators depended on which species occupied the monitored nest and location (study plot), but not on the year or the time of season. Corvids and raptors accounted for a relatively larger percentage of total predation of small ('warblers') and large ('thrushes') prey species, respectively, whereas carnivores were important predators of all prey species. Active nests of thrushes were only rarely robbed by Jays (< 4% of 52 events), presumably due to parental nest defence. Predation by woodpeckers was spatially clumped, probably due to individual foraging specialization. Predation by the other major predators was documented on most/all study plots.     

Behavioural changes and aversive conditioning in hunting dogs by the second-year confrontation with domestic sheep

Domesticated dogs occasionally exhibit predatory behaviour towards domestic sheep when running loose in pasture. Both young and old dogs of either sex may chase sheep. Electronic dog collars applying electric shocks are utilised as one method of training dogs to refrain from attacking sheep. This device is used for a number of other training purposes which have raised concern for the welfare of the dogs being trained. This study aims at testing long-term learning effects of previous sheep tests on sheep chasing in hunting dog breeds (Norwegian elkhounds (grey), English setters, and hare hunting dogs), in particular with use of electronic dog collars, in addition to uncovering potential secondary negative effects on dogs' behaviour and mental stability. The dogs (N=114) were subjected to three tests for two subsequent years, the second year being reported here. Dogs were tested for reactions to different stimuli, including a sheep, in a path test. In a sheep confrontation test, dogs were fenced in with a sheep group and given el. shocks when approaching 1-2m from sheep. A questionnaire to the dog owners reported differences in dogs' behaviour between the years.Dogs showed weaker or delayed behavioural responses in both tests in the second year. No dogs showed interest in or attacked a lone sheep in the path test in the second year, while almost two thirds of them did so the first year. In the sheep confrontation test, the dogs exhibited comparatively hesitant initial hunting motivation the second year, being more evident in dogs which received el. shocks the first year. No dogs chased or attacked sheep as their first response in this test, while half of them did so the first year. The proportion of dogs attacking sheep during the entire test was reduced to almost one fourth. The number of el. shocks administered reduced by the second year, and only one of the dogs that received el. shocks the first year received el. shocks the second year. The owners reported no negative effect on the dogs' behaviour during the year ensuing el. shock treatment. Eighteen of the 24 dogs reported by owners to exhibit behavioural changes lost their previous interest in sheep.The second-year tests indicate that aversive conditioning with the use of electronic dog collar may be an efficient method for reducing the probability of a dog chasing or attacking grazing sheep. No adverse effects were observed with our test procedure.     

Behavioural differences between three breed groups of hunting dogs confronted with domestic sheep

When running free in open fields, domestic dogs occasionally display predatory behaviour towards domestic sheep. This has not yet been studied scientifically. The aim of the present study was to investigate the inclination to chase sheep in three breed groups of hunting dogs that are most frequently used in areas with grazing sheep. We studied 41 elkhounds, 29 hare hunting dogs and 68 English setters. Behaviours indicative of motivation for chasing or attacking sheep were examined in three different ways. A path test examined functional traits such as hunting ability, contact willingness, reactivity to sudden noise, and response towards a lone sheep. In a sheep confrontation test, loose-leashed dogs were observed in a fenced enclosure with sheep and given electric shocks through an electronic dog collar if within 1-2m from the sheep. A questionnaire to the dog owners supplied information on their dog's previous experience with sheep and behavioural responses to various types of novel stimuli. No significant sex differences were found. The elkhounds showed the highest interest in a lone sheep in the path test, and displayed the highest initial hunting motivation, the highest percentage of dogs starting a sheep attack, the highest attack severity, and were most frequently given el. shocks. The hare hunting dogs were intermediate, while setters showed the lowest values for these variables. Dogs reported as showing low fearfulness more frequently acted as potential sheep chasers in the tests. Dogs up to 3 years of age showed a more pronounced initial hunting motivation and more frequent attacks than older dogs, although there were no age differences in the number of el. shocks given in the test. The latter may be related to the more frequent abruption of attacks among younger dogs. The main factors predicting a high hunting motivation and attack severity were lack of previous opportunity to chase sheep, low fearfulness towards gunshots and unfamiliar people, and general interest in sheep shown when encountering them. Probability of sheep chasing differed between dog breeds and age groups. Previous experience and certain character traits were indicative of a high predatory motivation towards sheep.     

Predator‐avoidance behaviour in a nocturnal petrel exposed to a novel predator

Many species of bird recognize acoustic and visual cues given by their predators and have complex defence adaptations to reduce predation risk. Recognition of threats posed by specific predators and specialized anti‐predation behaviours are common. In this study we investigated predator recognition and anti‐predation behaviours in a pelagic seabird, Leach's Storm‐petrel Oceanodroma leucorhoa, at a site where predation risk from Great Skuas Stercorarius skua is exceptionally high. Leach's Storm‐petrels breed in burrows and come on land only at night. Counter‐predator adaptations were investigated correlatively in relation to changing natural light levels at night, and experimentally in relation to nocturnal visual and acoustic signals from Great Skuas. Colony attendance by Leach's Storm‐petrels was attuned to changes in light conditions at night and was highest when nights were darkest. This behaviour is likely to reduce predation risk on land; however, specific recognition of Great Skuas and specialized defence behaviours were not found. Leach's Storm‐petrels, in particular apparently non‐breeding individuals, were entirely naïve to the threat posed by Great Skuas and were captured easily in a variety of different ways, on the ground and in the air. Lack of specialized behavioural adaptations in Leach's Storm‐petrels against Great Skuas may be because spatial overlap of breeding distributions of these species appears to be a rare and recent phenomenon.     

Livestock predation in the Bale Mountains,Ethiopia

In the Web Valley of the Bale Mountains National Park, the pastoral people suffer from livestock predation by wild carnivores. A total of 704 livestock were reported to be killed by wild carnivores over a 3‐year period, causing a loss of potential revenue of 12 USD per year per household. Reported annual predation rates equated to 1.4% of the livestock population of the study area. Spotted hyaenas were responsible for most livestock predation (57%), followed by leopards (18%), common jackals (16%) and servals (9%). Hyaenas killed all livestock types (horses, donkeys, mules, cattle, goats and sheep) whilst leopards, common jackals and servals killed mostly goats and sheep. A survey of 362 households revealed that the pastoral people keep dogs to protect livestock from carnivores. During 250 nights of observation in the ten settlements, pastoralists were alerted to the presence of hyaenas on 80 occasions by the barking of their dogs. Such tradition of keeping dogs presents a threat to the persistence of the endangered Ethiopian wolf through diseases transmission. Given the frequency of carnivore attacks on livestock, it is desirable to develop alternative livestock protection methods that both minimize livestock losses and reduce the risk of disease transmission to Ethiopian wolves.     

Prevalence of scrapie infection in Great Britain: interpreting the results of the 1997-1998 abattoir survey

An accurate estimate of the prevalence of scrapie infection in the Great Britain (GB) sheep flock is essential when assessing any potential risk to human health through exposure to sheep transmissible spongiform encephalopathies (TSEs). One method for assessing the prevalence is to sample sheep intended for human consumption using a diagnostic test capable of detecting infected animals prior to the onset of clinical signs. An abattoir survey conducted in Great Britain in 1997-1998 tested brain samples from 2809 apparently healthy sheep of which none was found to be positive for scrapie by histopathology or immunohistochemistry (IHC) although 10 were positive for scrapie-associated fibrils (SAF). Subsequently, the tonsils from a subset of the animals sampled were examined using IHC, one of which tested positive. To interpret these results we use a likelihood-based approach, which accounts for the variation in the prevalence of infection with age and test sensitivity and specificity with stage of infection. Combining the results for all of the diagnostic tests yields an estimate of the prevalence of scrapie infection in the GB sheep flock of 0.22% (95% confidence interval: 0.01-0.97%). Moreover, our analysis suggests that all of the diagnostic tests used are very specific (greater than 99%). Indeed, only SAF detection yields a specificity estimate of less than 100%, which helps to account for the high number of samples found to be positive for SAF.     

Intraflock variation in the speed of escape-flight response on attack by an avian predator

The benefits of flocking to prey species, whether through collective vigilance,dilution of risk, or predator confusion, depend on flock members respondingin a coordinated way to attack. We videotaped sparrowhawks attackingredshank flocks to determine if there were differences in thetiming of escape flights between flock members and the factorsthat might affect any differences. Sparrowhawks are surpriseshort-chase predators, so variation in the time taken to takeflight on attack is likely to be a good index of predation risk.Most birds in a flock flew within 0.25 s of the first bird flying,and all birds were flying within 0.7 s. Redshanks that werevigilant, that were closest to the approaching raptor, and thatwere close to their neighbors took flight earliest within aflock. Birds in larger flocks took longer, on average, to takeflight, measured from the time that the first bird in the flockflew. Most birds took flight immediately after near neighbors tookoff, but later flying birds were more likely to fly immediatelyafter more distant neighbors took flight. This result, alongwith the result that increased nearest neighbor distance increasedflight delay, suggests that most redshanks flew in responseto conspecifics flying. The results strongly suggest that thereis significant individual variation in predation risk withinflocks so that individuals within a flock will vary in benefitsthat they gain from flocking.     

A communication-based spatial model of antipredator vigilance

Many animals spend their lives performing two often mutually exclusive tasks: feeding and watching out for predators (anti-predator vigilance). There have been many theoretical and empirical studies investigating this trade-off, especially for birds. An important characteristic of a flock of real birds is the area occupied by the flock. Individuals feeding close together experience increased competition so that the feeding rate decreases. Widely spaced individuals may suffer a loss in vigilance efficiency, since communication between individuals is more difficult, such that the predation risk increases. A vigilance model is developed which allows birds to control their spacing (and so the area of the flock) as well as their vigilance rate. The best strategy for the birds is found under a variety of environmental conditions, under the assumption that each individual acts selfishly to maximize its own fitness.     

Naïve,bold, or just hungry? An invasive exotic prey species recognises but does not respond to its predators

Alien species experience both costs and benefits in invaded environments, through naiveté of potential prey species, but also predation pressure from native predators. The question of whether alien prey recognise and respond to native predators has been relatively understudied, despite the hypothesised potential for native predators to provide biotic resistance to invasion. There are two main hypotheses about whether exotic prey should recognise native and exotic predators in their new ranges: (1) naiveté—predicting recognition of evolutionarily familiar predators only, and (2) pre-adaptation—predicting recognition of all predators through a generalist recognition template. With regards to antipredator responses, (3) naïveté theory presumes that exotic prey will respond to the predators they recognise, but we suggest that (4) a bold behavioural syndrome, and/or a high marginal value of food in invaded environments might result in weak or absent responses, even to recognised predators. Here we combine the giving-up density framework with behavioural analysis of remote camera footage to experimentally test these ideas in a disturbed, peri-urban, Australian ecosystem, where alien black rats are predated on by alien dogs, foxes, cats, and native quolls. Black rats recognised dogs and foxes, but appear naïve towards quolls. However, they showed no antipredator responses at all, consistent with a bold behavioural syndrome, elevated predation risk, and/or a high marginal value of food in invaded environments.     

The effect of group size on vigilance in Ruddy Turnstones Arenaria interpres varies with foraging habitat

Foraging birds can manage time spent vigilant for predators by forming groups of various sizes. However, group size alone will not always reliably determine the optimal level of vigilance. For example, variation in predation risk or food quality between patches may also be influential. In a field setting, we assessed how simultaneous variation in predation risk and intake rate affects the relationship between vigilance and group size in foraging Ruddy Turnstones Arenaria interpres. We compared vigilance, measured as the number of ‘head‐ups’ per unit time, in habitat types that differed greatly in prey energy content and proximity to cover from which predators could launch surprise attacks. Habitats closer to predator cover provided foragers with much higher potential net energy intake rates than habitats further from cover. Foragers formed larger and denser flocks on habitats closer to cover. Individual vigilance of foragers in all habitats declined with increasing flock size and increased with flock density. However, vigilance by foragers on habitats closer to cover was always higher for a given flock size than vigilance by foragers on habitats further from cover, and habitat remained an important predictor of vigilance in models including a range of potential confounding variables. Our results suggest that foraging Ruddy Turnstones can simultaneously assess information on group size and the general likelihood of predator attack when determining their vigilance contribution.     

Putting (Big) Black Dog Syndrome to the Test: Evidence from a Large Metropolitan Shelter

Most support for Black Dog Syndrome or Big Black Dog Syndrome is anecdotal or theoretical. Yet some animal shelters/organizations have implemented strategies to address what they believe are lower rates of adoption and/or higher rates of euthanasia for “Big Black Dogs.” This study examines the persistent anecdotes and theories on humans’ preferences and aversions to dogs of various shades, using hierarchical multinomial logistic regression to predict outcomes for an analytic sample of 7,440 dogs from 2010–2011 in an urban, public animal shelter serving Louisville, Kentucky, USA. The relationship between coat shade and dog outcomes was not straightforward; while no relationship existed at the bivariate level, after controls were added, entirely black dogs showed somewhat lower odds of adoption—and higher euthanasia risk—than those characterized as secondarily black or sans black. Breed category, breed size, and purebred status were stronger predictors of dog outcomes than coat shade. Big Black Dog Syndrome was not supported by these data; smaller dogs were more likely to be euthanized if they were partly or wholly black. These findings may offer nuances to adoption strategies employed by shelters/organizations, help make better use of resources, and, perhaps, improve the likelihood of homing or rehoming shelter animals.     

Evaluating apparent competition in limiting the recovery of an endangered ungulate

Predation can disproportionately affect endangered prey populations when generalist predators are numerically linked to more abundant primary prey. Apparent competition, the term for this phenomenon, has been increasingly implicated in the declines of endangered prey populations. We examined the potential for apparent competition to limit the recovery of Sierra Nevada bighorn sheep (Ovis canadensis sierrae), an endangered subspecies under the US Endangered Species Act. Using a combination of location, demographic, and habitat data, we assessed whether cougar (Puma concolor) predation on endangered bighorn sheep was a consequence of their winter range overlap with abundant mule deer (Odocoileus hemionus). Consistent with the apparent competition hypothesis, bighorn sheep populations with higher spatial overlap with deer exhibited higher rates of cougar predation which had additive effects on adult survival. Bighorn sheep killed by cougars were primarily located within deer winter ranges, even though those areas constituted only a portion of the bighorn sheep winter ranges. We suspect that variation in sympatry between bighorn sheep and deer populations was largely driven by differences in habitat selection among bighorn sheep herds. Indeed, bighorn sheep herds that experienced the highest rates of predation and the greatest spatial overlap with deer also exhibited the strongest selection for low elevation habitat. Although predator-mediated apparent competition may limit some populations of bighorn sheep, it is not the primary factor limiting all populations, suggesting that the dynamics of different herds are highly idiosyncratic. Management plans for endangered species should consider the spatial distributions of key competitors and predators to reduce the potential for apparent competition to hijack conservation success.     

Development, growth, and egg production of Ageneotettix deorum (Orthoptera: Acrididae) in response to spider predation risk and elevated resource quality

Abstract.  1. Predation risk to insects is often size- or stage-selective and usually decreases as prey grow. Any factor, such as food quality, that accelerates developmental and growth rates is likely to reduce the period over which prey are susceptible to size-dependent predation.
2. Using field experiments, several hypotheses that assess growth, development, and egg production rates of the rangeland grasshopper Ageneotettix deorum (Scudder) were tested in response to combinations of food quality and predation risk from wolf spiders to investigate performance variation manifested through a behaviourally mediated path affecting food ingestion rates.
3. Grasshoppers with nutritionally superior food completed development ≈ 8–18% faster and grew 15–45% larger in the absence of spiders, in comparison with those subjected to low quality food exposed to spider predators. Growth and development did not differ for grasshoppers feeding on high quality food when predators were present in comparison with lower quality food unimpeded by predators. Responses indicated a compensatory relationship between resource quality and predation risk.
4. Surviving grasshoppers produced fewer eggs compared with individuals not exposed to spiders. Because no differences were found in daily egg production rate regardless of predation treatment, lower egg production was attributed to delayed age of first reproduction. Results compare favourably with responses observed in natural populations.
5. Risk of predation from spiders greatly reduced growth, development, and ultimately egg production. Increased food quality counteracts the impact of predation risk on grasshoppers through compensatory responses, suggesting that bottom-up factors mediate effects of spiders.     

Optimal predator management for mountain sheep conservation depends on the strength of mesopredator release

Large predators often suppress ungulate population growth, but they may also suppress the abundance of smaller predators that prey on neonatal ungulates. Antagonistic interactions among predators may therefore need to be integrated into predator–prey models to effectively manage ungulate–predator systems. We present a modeling framework that examines the net impact of interacting predators on the population growth rate of shared prey, using interactions among wolves Canis lupus, coyotes Canis latrans and Dall sheep Ovis dalli dalli as a case study. Wolf control is currently employed on approximately 16 million ha in Alaska to increase the abundance of ungulates for human harvest. We hypothesized that the positive effects of wolf control on Dall sheep population growth could be counteracted by increased levels of predation by coyotes. Coyotes and Dall sheep adult females (ewes) and lambs were radiocollared in the Alaska Range from 1999–2005 to estimate fecundity, age‐specific survival rates, and causes of mortality in an area without wolf control. We used stage‐structured population models to simulate the net effect of wolf control on Dall sheep population growth (λ). Our models accounted for stage‐specific predation rates by wolves and coyotes, compensatory mortality, and the potential release of coyote populations due to wolf control. Wolves were the main predators of ewes, coyotes were the main predators of lambs, and wolves were the main source of mortality for coyotes. Population models predicted that wolf control could increase sheep λ by 4% per year in the absence of mesopredator release. However, if wolf control released coyote populations, our models predicted that sheep λ could decrease by up to 3% per year. These results highlight the importance of integrating antagonistic interactions among predators into predator–prey models, because the net effect of predator management on shared prey can depend critically on the strength of mesopredator release.     

Coyote Abundance,Sheep Predation,and Wild Prey Correlates Illuminate Mediterranean Trophic Dynamics

ABSTRACT Sheep predation by coyotes (Canis latrans) is a major problem for sheep producers in North America. Solutions are facilitated by a basic understanding of the trophic dynamic context of this problem, one that likely varies geographically in important qualitative ways. Little is known about vertebrate trophic dynamics in Mediterranean ecosystems, where prey are diverse and their biomass is strongly influenced multi-annually by variable rainfall. We used long-term data sets from north-coastal California, USA, to investigate whether wild prey fluctuations caused immediate negative effects on sheep predation via a reduction in the coyote functional response or delayed positive effects on sheep predation via a numerical response by coyote predators. Because we could not measure prey biomass directly, we used variables associated with lower trophic levels (e.g., annual plant productivity, vole abundance, rainfall) as proxies for wild prey biomass. Coyote population growth rate was positively correlated with lower-trophic-level variables of the previous year, suggesting a numerical response, and sheep (ad F + lambs) predation was positively correlated with coyote abundance in the current year. Sheep predation also was negatively correlated with lower-trophic-level variables of the current year, suggesting an immediate buffering effect of wild prey on sheep predation. Together, coyote abundance and lower-trophic-level variables explained 47% of the multi-annual variation in sheep kills. The negative pathway between lower-trophic-level variables and sheep predation was stronger than the positive pathway, possibly due to the erratic nature of multi-annual fluctuations in lower-trophic-level variables, which could prevent the numerical response from reaching its full potential. Monthly analyses revealed a type III functional response of coyotes to lambs, which is expected to enhance buffering effects of wild prey on sheep predation. Our findings suggest the dominant effect of wild prey biomass on sheep predation by coyotes in this Mediterranean-type community is as a buffer.