Wolves-coyotes-foxes: a cascade among carnivores

Due to the widespread eradication of large canids and felids, top predators in many terrestrial ecosystems are now medium-sized carnivores such as coyotes. Coyotes have been shown to increase songbird and rodent abundance and diversity by suppressing populations of small carnivores such as domestic cats and foxes. The restoration of gray wolves to many parts of North America, however, could alter this interaction chain. Here we use a 30-year time series of wolf, coyote, and fox relative abundance from the state of Minnesota, USA, to show that wolves suppress coyote populations, which in turn releases foxes from top-down control by coyotes. In contrast to mesopredator release theory, which has often considered the consequence of top predator removal in a three-species interaction chain (e.g., coyote-fox-prey), the presence of the top predator releases the smaller predator in a four-species interaction chain. Thus, heavy predation by abundant small predators might be more similar to the historical ecosystem before top-predator extirpation. The restructuring of predator communities due to the loss or restoration of top predators is likely to alter the size spectrum of heavily consumed prey with important implications for biodiversity and human health.     

The Role of Incentive Programs in Conserving the Snow Leopard

Abstract:   Pastoralists and their livestock share much of the habitat of the snow leopard ( Uncia uncia ) across south and central Asia. The levels of livestock predation by the snow leopard and other carnivores are high, and retaliatory killing by the herders is a direct threat to carnivore populations. Depletion of wild prey by poaching and competition from livestock also poses an indirect threat to the region's carnivores. Conservationists working in these underdeveloped areas that face serious economic damage from livestock losses have turned to incentive programs to motivate local communities to protect carnivores. We describe a pilot incentive program in India that aims to offset losses due to livestock predation and to enhance wild prey density by creating livestock-free areas on common land. We also describe how income generation from handicrafts in Mongolia is helping curtail poaching and retaliatory killing of snow leopards. However, initiatives to offset the costs of living with carnivores and to make conservation beneficial to affected people have thus far been small, isolated, and heavily subsidized. Making these initiatives more comprehensive, expanding their coverage, and internalizing their costs are future challenges for the conservation of large carnivores such as the snow leopard.     

Landscape ecology of eastern coyotes based on large-scale estimates of abundance.

Since their range expansion into eastern North America in the mid-1900s, coyotes (Canis latrans) have become the region's top predator. Although widespread across the region, coyote adaptation to eastern forests and use of the broader landscape are not well understood. We studied the distribution and abundance of coyotes by collecting coyote feces from 54 sites across a diversity of landscapes in and around the Adirondacks of northern New York. We then genotyped feces with microsatellites and found a close correlation between the number of detected individuals and the total number of scats at a site. We created habitat models predicting coyote abundance using multi-scale vegetation and landscape data and ranked them with an information-theoretic model selection approach. These models allow us to reject the hypothesis that eastern forests are unsuitable habitat for coyotes as their abundance was positively correlated with forest cover and negatively correlated with measures of rural non-forest landscapes. However, measures of vegetation structure turned out to be better predictors of coyote abundance than generalized "forest vs. open" classification. The best supported models included those measures indicative of disturbed forest, especially more open canopies found in logged forests, and included natural edge habitats along water courses. These forest types are more productive than mature forests and presumably host more prey for coyotes. A second model with only variables that could be mapped across the region highlighted the lower density of coyotes in areas with high human settlement, as well as positive relationships with variables such as snowfall and lakes that may relate to increased numbers and vulnerability of deer. The resulting map predicts coyote density to be highest along the southwestern edge of the Adirondack State Park, including Tug Hill, and lowest in the mature forests and more rural areas of the central and eastern Adirondacks. Together, these results support the need for a nuanced view of how eastern coyotes use forested habitats.     

Managing conflict between large carnivores and livestock

Large carnivores are persecuted globally because they threaten human industries and livelihoods. How this conflict is managed has consequences for the conservation of large carnivores and biodiversity more broadly. Mitigating human–predator conflict should be evidence‐based and accommodate people's values while protecting carnivores. Despite much research into human and large‐carnivore coexistence strategies, there have been few attempts to document the success of conflict‐mitigation strategies on a global scale. We conducted a meta‐analysis of global research on conflict mitigation related to large carnivores and humans. We focused on conflicts that arise from the threat large carnivores pose to livestock. We first used structured and unstructured searching to identify replicated studies that used before–after or control–impact design to measure change in livestock loss as a result of implementing a management intervention. We then extracted relevant data from these studies to calculate an overall effect size for each intervention type. Research effort and focus varied among continents and aligned with the histories and cultures that shaped livestock production and attitudes toward carnivores. Livestock guardian animals most effectively reduced livestock losses. Lethal control was the second most effective control, although its success varied the most, and guardian animals and lethal control did not differ significantly. Financial incentives have promoted tolerance of large carnivores in some settings and reduced retaliatory killings. We suggest coexistence strategies be location‐specific, incorporate cultural values and environmental conditions, and be designed such that return on financial investment can be evaluated. Improved monitoring of mitigation measures is urgently required to promote effective evidence‐based policy.     

Wild Dog Demography in Hluhluwe-Umfolozi Park, South Africa

Abstract: Wild dogs (   Lycaon pictus ) are highly threatened carnivores, and conservation of their dwindling numbers is needed. Isolated populations contribute little to these conservation efforts, so linking populations of dogs is a necessary goal to prevent extinction. I gathered demographic information from a small population of wild dogs in Hluhluwe-Umfolozi Park, KwaZulu-Natal, South Africa. Photographs and archive records from 1981 to 1996 were used to compile information on this population. Demographic parameters from the released population were compared with information on wild dogs elsewhere. Most demographic parameters of the Hluhluwe-Umfolozi Park population were similar to published information from other populations. The main difference was the effect of a single pack on population performance: pack formation was unlikely, unrelated animals were rare, and major losses (emigration and mortalities) occurred. Increasing the number of packs by introducing more dogs would be useful but would be only a short-term solution. Increasing the local population size and artificially linking populations in southern Africa appears to be the only longer-term solution to ensure the viability of wild dogs on the subcontinent.     

Limiting Depredation by African Carnivores: the Role of Livestock Husbandry

Abstract:   Most large carnivore species are in global decline. Conflict with local people, particularly over depredation on livestock, is a major cause of this decline, affecting both nominally protected populations and those outside protected areas. For this reason, techniques that can resolve conflicts between large carnivores and livestock farmers may make important contributions to conservation. We monitored rates of livestock depredation by lions (  Panthera leo ), leopards ( Panthera pardus ), cheetahs ( Acinonyx jubatus ), and spotted hyenas ( Crocuta crocuta ), and retributive killing of these species by farmers in livestock-producing areas of Laikipia District, Kenya. Farmers killed more lions, leopards, and spotted hyenas where these predators killed more livestock. Livestock husbandry had a clear effect on rates of depredation and hence on the numbers of predators killed. Cattle, sheep, and goats experienced the lowest predation rates when attentively herded by day and enclosed in traditional corrals (bomas) by night. Construction of the boma, the presence of watchdogs, and high levels of human activity around the boma were all associated with lower losses to predators. Although most of this work was carried out on commercial ranches, local Maasai and Samburu pastoralists have practiced nearly identical forms of husbandry for generations. Our study shows that traditional, low-tech husbandry approaches can make an important contribution to the conservation of large carnivores.     

Recolonizing wolves and mesopredator suppression of coyotes: impacts on pronghorn population dynamics.

Food web theory predicts that the loss of large carnivores may contribute to elevated predation rates and, hence, declining prey populations, through the process of mesopredator release. However, opportunities to test predictions of the mesopredator release hypothesis are rare, and the extent to which changes in predation rates influence prey population dynamics may not be clear due to a lack of demographic information on the prey population of interest. We utilized spatial and seasonal heterogeneity in wolf distribution and abundance to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus) throughout much of the United States, contributes to high rates of neonatal mortality in ungulates. To test this hypothesis, we contrasted causes of mortality and survival rates of pronghorn (Antilocapra americana) neonates captured at wolf-free and wolf-abundant sites in western Wyoming, USA, between 2002 and 2004. We then used these data to parameterize stochastic population models to heuristically assess the impact of wolves on pronghorn population dynamics due to changes in neonatal survival. Coyote predation was the primary cause of mortality at all sites, but mortality due to coyotes was 34% lower in areas utilized by wolves (P < 0.001). Based on simulation modeling, the realized population growth rate was 0.92 based on fawn survival in the absence of wolves, and 1.06 at sites utilized by wolves. Thus, wolf restoration is predicted to shift the trajectory of the pronghorn population from a declining to an increasing trend. Our results suggest that reintroductions of large carnivores may influence biodiversity through effects on prey populations mediated by mesopredator suppression. In addition, our approach, which combines empirical data on the population of interest with information from other data sources, demonstrates the utility of using simulation modeling to more fully evaluate ecological theories by moving beyond estimating changes in vital rates to analyses of population-level impacts.     

Food webs and intraguild predation: community interactions of a native mesocarnivore

Trophic level interactions between predators create complex relationships such as intraguild predation. Theoretical research has predicted two possible paths to stability in intraguild systems: intermediate predators either outcompete higher-order predators for shared resources or select habitat based on security. The effects of intraguild predation on intermediate mammalian predators such as swift foxes (Vulpes velox) are not well understood. We examined the relationships between swift foxes and both their predators and prey, as well the effect of vegetation structure on swift fox-coyote (Canis latrans) interactions, between August 2001 and August 2004. In a natural experiment created by the Pinon Canyon Maneuver Site in southeastern Colorado, USA, we documented swift fox survival and density in a variety of landscapes and compared these parameters in relation to prey availability, coyote abundance, and vegetation structure. Swift fox density varied significantly between study sites, while survival did not. Coyote abundance was positively related to the basal prey species and vegetation structure, while swift fox density was negatively related to coyote abundance, basal prey species, and vegetation structure. Our results support the prediction that, under intraguild predation in terrestrial systems, top predator distribution matches resource availability (resource match), while intermediate predator distribution inversely matches predation risk (safety match). While predation by coyotes may be the specific cause of swift fox mortality in this system, the more general mechanism appears to be exposure to predation moderated by shrub density.     

Birds as predators in tropical agroforestry systems

Insectivorous birds reduce arthropod abundances and their damage to plants in some, but not all, studies where predation by birds has been assessed. The variation in bird effects may be due to characteristics such as plant productivity or quality, habitat complexity, and/or species diversity of predator and prey assemblages. Since agroforestry systems vary in such characteristics, these systems provide a good starting point for understanding when and where we can expect predation by birds to be important. We analyze data from bird exclosure studies in forests and agroforestry systems to ask whether birds consistently reduce their arthropod prey base and whether bird predation differs between forests and agroforestry systems. Further, we focus on agroforestry systems to ask whether the magnitude of bird predation (1) differs between canopy trees and understory plants, (2) differs when migratory birds are present or absent, and (3) correlates with bird abundance and diversity. We found that, across all studies, birds reduce all arthropods, herbivores, carnivores, and plant damage. We observed no difference in the magnitude of bird effects between agroforestry systems and forests despite simplified habitat structure and plant diversity in agroforests. Within agroforestry systems, bird reduction of arthropods was greater in the canopy than the crop layer. Top-down effects of bird predation were especially strong during censuses when migratory birds were present in agroforestry systems. Importantly, the diversity of the predator assemblage correlated with the magnitude of predator effects; where the diversity of birds, especially migratory birds, was greater, birds reduced arthropod densities to a greater extent. We outline potential mechanisms for relationships between bird predator, insect prey, and habitat characteristics, and we suggest future studies using tropical agroforests as a model system to further test these areas of ecological theory.     

The Paradox of the Long‐Term Positive Effects of a North American Crayfish on a European Community of Predators

Abstract: Invasions of non‐native species are one of the major causes of losses of native species. In some cases, however, non‐natives may also have positive effects on native species. We investigated the potential facilitative effects of the North American red swamp crayfish (Procambarus clarkii) on the community of predators in southwestern Spain. To do so, we examined the diets of predators in the area and their population trends since introduction of the crayfish. Most predator species consumed red swamp crayfish, which sometimes occurred in over 50% of their diet samples. Moreover, the abundance of species preying on crayfish increased significantly in the area as opposed to the abundance of herbivores and to predator populations in other areas of Europe, where those predators are even considered threatened. Thus, we report the first case in which one non‐native species is both beneficial because it provides prey for threatened species and detrimental because it can drive species at lower trophic levels to extinction. Increases in predator numbers that are associated with non‐native species of prey, especially when some of these predators are also invasive non‐natives, may increase levels of predation on other species and produce cascading effects that threaten native biota at longer temporal and larger spatial scales. Future management plans should include the complexity of interactions between invasive non‐natives and the entire native community, the feasibility of successful removal of non‐native species, and the potential social and economic interests in the area.     

Laboratory studies of the effect of predation on production and the production:biomass ratio of the opportunistic polychaete Capitella capitata (Type I)

The effect of simulated predation upon the secondary production and P:B (production:biomass) ratio of the polychaete Capitella capitata (Type I) was estimated from laboratory studies. The first method (the maximum sustainable yield, MSY, method) summed net increases in biomass of each population over time with biomass exploited by predation. In the specific growth-rate method, experimentally determined specific-growth rates were applied to changes in size classes and standing stock over time, providing another estimate of production for comparison to the MSY method. Predation had a pronounced effect on the magnitude of production, standing stocks, and hence the P:B ratio causing a fourfold difference in P:B ratios between the controls (P:B=4.9) and the 23% wk^-1 predation rate (P:B=19.6). Production reached a high of 87 g ash-free dry wt m^-2 yr^-1 in the highest predation treatment (23% wk^-1). An estimate of the number of individuals recruited in each population showed that predation caused an increase in population turnover rate. Gross ecological efficiency (calories of food ingested by the predator/calories of food consumed by the prey) and food-chain efficiency (calories of prey ingested by the predator/calories of food supplied to the prey) were 7.4 and 5.8% respectively, for the 23% wk^-1 predation treatment.     

Behavioral resource depression and decaying perceived risk of predation in two species of coexisting gerbils

Summary Behavioral resource depression occurs when the behavior of prey individuals changes in response to the presence of a predator, resulting in a reduction of the encounter rate of the predator with its prey. Here I present experimental evidence on the response of two species of gerbils (Gerbillus allenbyi and G. pyramidum) to the presence of barn owls. I conducted the experiments in a large aviary. Both gerbils responded to the presence of barn owl predators by foraging in fewer resource patches (seed trays) and by quitting foraged resource patches at a higher resource harvest rate (giving-up density of resource; GUD). This reduced the amount of time gerbils were exposed to owl predation, and hence the encounter rate of owls with gerbils, i.e., behavioral resource depression. Thus, the presence of owls imposes a foraging cost on gerbils due to risk of predation, and also on the owls themselves due to resource depression. I then examined how resource depression relaxed over time following exposure to owls. In the days following an encounter with the predator, the reduction in foraging activity for both gerbil species eased. Increasing numbers of trays were foraged each day, and GUDs in seed trays declined. The two gerbils differed in their rate of recovery, with G. pyramidum returning to prepredator levels of foraging after 1 or 2 nights and G. allenbyi taking 5 nights or longer. Interspecific differences in recovery rates may be based on differences between the species in vulnerability to predation and/or ability to detect the presence of predators. The differences in recovery rates may be due to optimal memory windows or decay rates, where differences between species are based on risk of predation or on how perceived risk changes with time since a predator was last encountered. Finally, differences between or among competitors in recovery from resource depression may provide foraging opportunities in time for the species which recover most quickly and may have implications for species coexistence.     

Indirect effects and traditional trophic cascades: a test involving wolves, coyotes, and pronghorn

The traditional trophic cascades model is based on consumer resource interactions at each link in a food chain. However, trophic-level interactions, such as mesocarnivore release resulting from intraguild predation, may also be important mediators of cascades. From September 2001 to August 2004, we used spatial and seasonal heterogeneity in wolf distribution and abundance in the southern Greater Yellowstone Ecosystem to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus), accounts for high rates of coyote predation on pronghorn (Antilocapra americana) fawns observed in some areas. Results of this ecological perturbation in wolf densities, coyote densities, and pronghorn neonatal survival at wolf-free and wolf-abundant sites support the existence of a species-level trophic cascade. That wolves precipitated a trophic cascade was evidenced by fawn survival rates that were four-fold higher at sites used by wolves. A negative correlation between coyote and wolf densities supports the hypothesis that interspecific interactions between the two species facilitated the difference in fawn survival. Whereas densities of resident coyotes were similar between wolf-free and wolf-abundant sites, the abundance of transient coyotes was significantly lower in areas used by wolves. Thus, differential effects of wolves on solitary coyotes may be an important mechanism by which wolves limit coyote densities. Our results support the hypothesis that mesopredator release of coyotes contributes to high rates of coyote predation on pronghorn fawns, and demonstrate the importance of alternative food web pathways in structuring the dynamics of terrestrial systems.     

Effects of Management of Domestic Dogs and Recreation on Carnivores in Protected Areas in Northern California

Abstract: In developed countries dogs (Canis lupus familiaris) are permitted to accompany human visitors to many protected areas (e.g., >96% of protected lands in California, U.S.A.), and protected‐area management often focuses on regulating dogs due to concerns about predation, competition, or transmission of disease and conflicts with human visitors. In 2004 and 2005, we investigated whether carnivore species richness and abundance were associated with management of domestic dogs and recreational visitation in protected areas in northern California. We surveyed for mammalian carnivores and human visitors in 21 recreation areas in which dogs were allowed offleash or onleash or were excluded, and we compared our observations in the recreation areas with observations in seven reference sites that were not open to the public. Carnivore abundance and species richness did not differ among the three types of recreation areas, but native carnivore species richness was 1.7 times greater (p < 0.01) and the relative abundances of native coyotes (Canis latrans) and bobcats (Lynx rufus) were over four times greater (p < 0.01) in the reference sites. Abundances of bobcats and all carnivores declined as the number of visitors increased. The policy on domestic dogs did not appear to affect species richness and abundance of mammalian carnivores. But the number of dogs we observed was strongly associated with human visitation (R²= 0.54), so the key factors associated with recreational effects on carnivores appear to be the presence and number of human visitors to protected areas.     

Impact of cannibalism on predator-prey dynamics: size-structured interactions and apparent mutualism

Direct and indirect interactions between two prey species can strongly alter the dynamics of predator-prey systems. Most predators are cannibalistic, and as a consequence, even systems with only one predator and one prey include two prey types: conspecifics and heterospecifics. The effects of the complex direct and indirect interactions that emerge in such cannibalistic systems are still poorly understood. This study examined how the indirect interaction between conspecific and heterospecific prey affects cannibalism and predation rates and how the direct interactions between both species indirectly alter the effect of the cannibalistic predator. I tested for these effects using larvae of the stream salamanders Eurycea cirrigera (prey) and Pseudotriton ruber (cannibalistic predator) by manipulating the relative densities of the conspecific and heterospecific prey in the presence and absence of the predator in experimental streams. The rates of cannibalism and heterospecific predation were proportional to the respective densities and negatively correlated, indicating a positive indirect interaction between conspecific and heterospecific prey, similar to "apparent mutualism." Direct interactions between prey species did not alter the effect of the predator. Although both types of prey showed a similar 30% reduction in night activity and switch in microhabitat use in response to the presence of the predator, cannibalism rates were three times higher than heterospecific predation rates irrespective of the relative densities of the two types of prey. Cumulative predation risks differed even more due to the 48% lower growth rate of conspecific prey. Detailed laboratory experiments suggest that the 3:1 difference in cannibalism and predation rate was due to the higher efficiency of heterospecific prey in escaping immediate attacks. However, no difference was observed when the predator was a closely related salamander species, Gyrinophilus porphyriticus, indicating that this difference is species specific. This demonstrates that cannibalism can result in the coupling of predator and prey mortality rates that strongly determines the dynamics of predator-prey systems.     

The growth-predation risk trade-off under a growing gape-limited predation threat

Growth is a critical ecological trait because it can determine population demography, evolution, and community interactions. Predation risk frequently induces decreased foraging and slow growth in prey. However, such strategies may not always be favored when prey can outgrow a predator's hunting ability. At the same time, a growing gape-limited predator broadens its hunting ability through time by expanding its gape and thereby creates a moving size refuge for susceptible prey. Here, I explore the ramifications of growing gape-limited predators for adaptive prey growth. A discrete demographic model for optimal foraging/growth strategies was derived under the realistic scenario of gape-limited and gape-unconstrained predation threats. Analytic and numerical results demonstrate a novel fitness minimum just above the growth rate of the gape-limited predator. This local fitness minimum separates a slow growth strategy that forages infrequently and accumulates low but constant predation risk from a fast growth strategy that forages frequently and experiences a high early predation risk in return for lower future predation risk and enhanced fecundity. Slow strategies generally were advantageous in communities dominated by gape-unconstrained predators whereas fast strategies were advantageous in gape-limited predator communities. Results were sensitive to the assumed relationships between prey size and fecundity and between prey growth and predation risk. Predator growth increased the parameter space favoring fast prey strategies. The model makes the testable predictions that prey should not grow at the same rate as their gape-limited predator and generally should grow faster than the fastest growing gape-limited predator. By focusing on predator constraints on prey capture, these results integrate the ecological and evolutionary implications of prey growth in diverse predator communities and offer an explanation for empirical growth patterns previously viewed to be anomalies.     

Food and predators affect egg production in song sparrows

Although the possibility that food and predators may interact in limiting avian populations has long been recognized, there have been few attempts to test this experimentally in the field. We conducted a manipulative food addition experiment on the demography of Song Sparrows (Melospiza melodia) across sites that varied in predator abundance, near Victoria, British Columbia, Canada, over three consecutive breeding seasons. We previously showed that food and predators had interactive effects on annual reproductive success (young fledged per female). Here, we report the effects on egg production. Our results show that food limits the total number of eggs laid over the breeding season ("total egg production") and that interactive food and predator effects, including food effects on nest predation, determine how those eggs are "parceled out" into different nests. Food addition alone significantly affected total egg production, and there was no significant interannual variability in this result. At the same time, both food and predators affected the two determinants of total egg production: "clutch number" (total number of clutches laid) and average clutch size. Both clutch number and size were affected by a food x predator x year interaction. Clutch number was lower at low-predator locations because there was less nest predation and thus less renesting. Food addition also significantly reduced nest predation, but there was significant interannual variation in this effect. This interannual variation was responsible for the food x predator x year interactions because the larger the effect of food on nest predation in a given year, the smaller was the effect of food on clutch number; and the smaller the effect of food on clutch number, the larger was the effect of food on clutch size. Potential predator and year effects on total egg production were thus cancelled out by an inverse relationship between clutch number and clutch size. We suggest that combined food and predator effects on demography could be the norm in both birds and mammals.     

Effects of climate and exurban development on nest predation and predator presence in the southern Appalachian Mountains (USA)

In the eastern United States, land-use and climate change have likely contributed to declines in the abundance of Neotropical migrant birds that occupy forest interiors, but the mechanisms are not well understood. We conducted a nest-predation experiment in southern Appalachian Mountain forests (North Carolina, U.S.A.) during the 2009 and 2010 breeding seasons to determine the effects of exurban development and temperature on predator presence and the average number of days until eggs in an artificial nest were disturbed by predators. We baited artificial nests with quail (Excalfactoria chinensi) eggs and monitored them for 18 days. We used clay eggs, track plates, and motion-triggered cameras to detect and identify nest predators. The average number of days a nest was undisturbed decreased as mean temperature increased and, to a lesser extent, as the density of buildings increased. Nests on the ground were more often depredated than those in trees, likely due to increased predation by opossum (Didelphis virginiana) and other carnivores. Raccoons (Procyon lotor), opossums, corvids (Corvus brachyrhynchos and Cyanocitta cristata), chipmunks (Tamias striatus), black bears (Ursus americanus), and domestic cats (Felis catus) were the most commonly detected predators. Presence of these predators did not vary as a function of mean temperature. Domestic cats and corvids were detected more frequently in plots with high rather than low densities of buildings. Forest-interior specialists and Neotropical migrants often nest in cool, high-elevation areas with low housing density. These bird species, especially those that nest on the ground, may be most vulnerable to increased nest predation if temperature and exurban development increase at higher elevations as anticipated.     

Can rare positive interactions become common when large carnivores consume livestock?

Livestock populations in protected areas are viewed negatively because of their interaction with native ungulates through direct competition for food resources. However, livestock and native prey can also interact indirectly through their shared predator. Indirect interactions between two prey species occur when one prey modifies either the functional or numerical responses of a shared predator. This interaction is often manifested as negative effects (apparent competition) on one or both prey species through increased predation risk. But indirect interactions can also yield positive effects on a focal prey if the shared predator modifies its functional response toward increased consumption of an abundant and higher-quality alternative prey. Such a phenomenon between two prey species is underappreciated and overlooked in nature. Positive indirect effects can be expected to occur in livestock-dominated wildlife reserves containing large carnivores. We searched for such positive effects in Acacia-Zizhypus forests of India's Gir sanctuary where livestock (Bubalus bubalis and Bos indicus) and a coexisting native prey (chital deer, Axis axis) are consumed by Asiatic lions (Panthera leo persica). Chital vigilance was higher in areas with low livestock density than in areas with high livestock density. This positive indirect effect occurred because lion predation rates on livestock were twice as great where livestock were abundant than where livestock density was low. Positive indirect interactions mediated by shared predators may be more common than generally thought with rather major consequences for ecological understanding and conservation. We encourage further studies to understand outcomes of indirect interactions on long-term predator-prey dynamics in livestock-dominated protected areas.     

Novel features of an inducible defense system in larval tree frogs (Hyla chrysoscelis)

Organisms in aquatic ecosystems must often tolerate variable environmental conditions, including an uncertain risk of predation. Individuals that can maintain plastic defenses against predation will increase their survival when predators are present, but will not incur the costs of these defenses when the risk of predation is low and the defense is not induced. Larvae of the pond-breeding anuran Hyla chrysoscelis develop a conspicuous phenotype in the presence of predators consisting of a brightly colored tail and a deeper tail fin. In this study, I attempted to identify the source of the chemical signal that induces this defensive morphology in this species. I tested whether metabolites alone, originating from the prey but passing through the predator, were able to induce the same morphological response as the combination of alarm signals released directly by attacked conspecifics, and metabolites. I used morphometric and tail conspicuousness data to assess tadpole response to the perceived risk of predation by larval odonate predators (Anax junius). I also tested whether this inducing cue could be recognized across species by measuring the morphological response of H. chrysoscelis tadpoles exposed to cues emitted when tadpoles of a closely related genus (Pseudacris crucifer) were consumed. Tadpoles exhibited a clean graded response of both overall shape and tail morphology in response to all cues, corresponding to their relative reliability as indicators of a risk of predation. H. chrysoscelis tadpoles were also able to respond to cues emitted when tadpoles of a closely related genus were consumed by predators. These results illustrate that tadpoles of this species are able to respond to metabolites alone without alarm signals, and that interspecific chemical communication is a primary mechanism for predator avoidance in this inducible defense system.