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.     

Willow on Yellowstone's northern range: evidence for a trophic cascade?

Reintroduction of wolves (Canis lupus) to Yellowstone National Park in 1995-1996 has been argued to promote a trophic cascade by altering elk (Cervus elaphus) density, habitat-selection patterns, and behavior that, in turn, could lead to changes within the plant communities used by elk. We sampled two species of willow (Salix boothii and S. geyeriana) on the northern winter range to determine whether (1) there was quantitative evidence of increased willow growth following wolf reintroduction, (2) browsing by elk affected willow growth, and (3) any increase in growth observed was greater than that expected by climatic and hydrological factors alone, thereby indicating a trophic cascade caused by wolves. Using stem sectioning techniques to quantify historical growth patterns we found an approximately twofold increase in stem growth-ring area following wolf reintroduction for both species of willow. This increase could not be explained by climate and hydrological factors alone; the presence of wolves on the landscape was a significant predictor of stem growth above and beyond these abiotic factors. Growth-ring area was positively correlated with the previous year's ring area and negatively correlated with the percentage of twigs browsed from the stem during the winter preceding growth, indicating that elk browse impeded stem growth. Our results are consistent with the hypothesis of a behaviorally mediated trophic cascade on Yellowstone's northern winter range following wolf reintroduction. We suggest that the community-altering effects of wolf restoration are an endorsement of ecological-process management in Yellowstone National Park.     

A dynamic explanation of size-density scaling in carnivores

Population abundance is negatively related to body size for many types of organisms. Despite the ubiquity of size-density scaling relationships, we lack a general understanding of the underlying mechanisms. Although dynamic models suggest that it is possible to predict the intercept and slope of the scaling relationship from prior observations, this has never been empirically attempted. Here we fully parameterize a set of consumer-resource models for mammalian carnivores and successfully predict the size-density scaling relationship for this group without the use of free parameters. All models produced similar predictions, but comparison of nested models indicated that the primary factors generating size-density scaling in mammalian carnivores are prey productivity, predator-prey size ratios, and consumer area of capture.     

A simple plant mutation abets a predator-diversity cascade

Resource consumption often increases with greater consumer biodiversity. This could result either from complementarity among consumers or the inclusion of particular key species, and it is often difficult to differentiate between these two mechanisms. We exploited a simple plant mutation (reduced production of surface waxes) to alter foraging within a community of aphid predators, and thus perhaps shift the nature of resulting predator diversity effects. We found that greater predator species richness dramatically increased prey suppression and plant biomass only on mutant, reduced-wax pea plants (Pisum sativum). On pea plants from a sister line with wild type, waxier plant surfaces, predator species richness did not influence predators' impacts on herbivores or plants. Thus, a change in plant surface structure acted to turn on, or off, the cascading effects of predator diversity. Greater predator richness encouraged higher densities of true predators but did not lead to greater reproduction by a parasitoid, Aphidius ervi; fecundity of each natural enemy species was similar for the two plant types. Behavioral observations indicated that although A. ervi was less likely to forage within species-rich predator communities, low-wax plants mitigated this interference by encouraging generally greater A. ervi foraging and thus high rates of aphid dislodgement (aphids dropped from plants to escape A. ervi, but not the other predators). Thus, only species-rich, low-wax plants simultaneously encouraged strong species-specific effects of A. ervi, and strong complementarity among the other predator species. In summary, our study provides evidence that diversity effects in predator assemblages are sensitive to habitat characteristics. Further, we show that a simple plant morphological trait, controlled by a single gene mutation, can dramatically alter the cascading effects of predator species richness on herbivores and plants.     

Energetics,reproductive suppression and obligate communal breeding in carnivores

Summary Leptogenys processionalis Jerdon forages on termites and other arthropods by raiding in branched trails. Growth and topology of these search trails were studied using Horton's (1945) technique orginally developed to analyze the branching pattern of river systems. Branching was always a bifurcating process and branches emerged symmetrically on either side of the main trail. Branching coefficients (R _b ) were similar to those of a few biological branching systems, such as lungs, that are considered to be non-random in their branching pattern. The R _b values indicated that the rate of branching and growth of trails remained constant within each foraging bout. The length of trails became shorter as they grew out and branched. The branching process was a function of the spatial separation of food patches in the terminal search field. Ants in the terminal search field send signals on encountering prey. The recruits cannot discriminate between these signals if they arise from two food patches situated <40 cm from each other, and hence converge on them in a single trail. However, discrimination is possible when food patches are >40 cm apart and hence recruits congregate on them separately in two trails, resulting in branching. Thus, the branching process is a result of independent decision by the ants conforming to certain simple rules and not a collective decision of the whole colony. We argue that mass recruiting ants selected to forage by branching pattern of trails because of its efficiency over other topologies (Stevens 1973) in minimizing the cost of travel, both from the nest to the food patches and between food patches. Further, the branch angles appear to be a trade-off to minimize travel cost and the resistance to the flow of ants comprising the column.Offprint requests to: T. Veena     

Bite forces and evolutionary adaptations to feeding ecology in carnivores

The Carnivora spans the largest ecological and body size diversity of any mammalian order, making it an ideal basis for studies of evolutionary ecology and functional morphology. For animals with different feeding ecologies, it may be expected that bite force represents an important evolutionary adaptation, but studies have been constrained by a lack of bite force data. In this study we present predictions of bite forces for 151 species of extant carnivores, comprising representatives from all eight families and the entire size and ecological spectrum within the order. We show that, when normalized for body size, bite forces differ significantly between the various feeding categories. At opposing extremes and independent of genealogy, consumers of tough fibrous plant material and carnivores preying on large prey both have high bite forces for their size, while bite force adjusted for body mass is low among specialized insectivores. Omnivores and carnivores preying on small prey have more moderate bite forces for their size. These findings indicate that differences in bite force represent important adaptations to and indicators of differing feeding ecologies throughout carnivoran evolution. Our results suggest that the incorporation of bite force data may assist in the construction of more robust evolutionary and palaeontological analyses of feeding ecology.     

The land-cover cascade: relationships coupling land and water

We introduce the land-cover cascade (LCC) as a conceptual framework to quantify the transfer of land-cover-disturbance effects to stream biota. We hypothesize that disturbance is propagated through multivariate systems through key variables that transform a disturbance and pass a reorganized disturbance effect to the next hierarchical level where the process repeats until ultimately affecting biota. We measured 31 hydrologic, geomorphic, erosional, and substrate variables and 26 biotic responses that have been associated with land-use disturbance in third- and fourth-order streams in the Blue Ridge physiographic province in western North Carolina (USA). Regression analyses reduced this set of variables to include only those that responded to land cover and/or affected biota. From this reduced variable set, hypotheses were generated that predicted the disturbance pathways affecting each biotic response following the land-cover-cascade design. Cascade pathways began with land cover and ended with biotic responses, passing through at least one intermediate ecosystem abiotic component. Cascade models were tested for predictive ability and goodness-of-fit using path analysis. Biota were influenced by near-stream urban, agricultural, and forest land cover as propagated by hydrologic (e.g., discharge), geomorphic (e.g., stream bank height), erosional (e.g., suspended sediments), and depositional streambed (e.g., substrate size) features occurring along LCC pathways, reflecting abiotic mechanisms mediating land-cover disturbance. Our results suggest that communities are influenced by land-cover change indirectly through a hierarchy of associated abiotic components that propagate disturbance to biota. More generally, the land-cover cascade concept and experimental framework demonstrate an organized approach to the generic study of cascades and the complex relationships between landscapes and streams.     

Induced defenses in herbivores and plants differentially modulate a trophic cascade

Inducible defenses are dynamic traits that modulate the strength of both plant-herbivore and herbivore-carnivore interactions. Surprisingly few studies have considered the relative contributions of induced plant and herbivore defenses to the overall balance of bottom-up and top-down control. Here we compare trophic cascade strengths using replicated two-level and three-level plankton communities in which we systematically varied the presence or absence of induced defenses at the plant and/or herbivore levels. Our results show that a trophic cascade, i.e., significantly higher plant biomass in three-level than in two-level food chains, occurred whenever herbivores were undefended against carnivores. Trophic cascades did not occur when herbivores exhibited an induced defense. This pattern was obtained irrespective of the presence or absence of induced defenses at the plant level. We thus found that herbivore defenses, not plant defenses, had an overriding effect on cascade strength. We discuss these results in relation to variation in cascade strengths in natural communities.     

Defended versus undefended home range size of carnivores,ungulates and primates

Summary We tested the hypothesis that undefended home ranges are larger than defended home ranges using data collected from the literature for three groups of mammals. A matched-pairs analysis of populations within species or species within genera showed that undefended home ranges were larger than defended home ranges for carnivores and male ungulates, but not for primates. Primates may have been an exception because they violated a key assumption of the hypothesis, that defence costs increase with the size of the defended area. Undefended home ranges were 5.4 and 15.2 times larger than defended home ranges for carnivores and male ungulates, respectively. Whether or not a home range is defended is an important source of variation that should be included in future studies of home range size. Correspondence to: J.W.A. Grant     

Constraints on group size in primates and carnivores: population density and day-range as assays of exploitation competition

Summary We hypothesise that foraging group size (FGS) and population group size (PGS) in primates and carnivores are related to quantifiable variables indexing the intensity of exploitative competition. Group size is predicted to increase with both food density and travel capabilities, as estimated by the constraint-free day-range (DR_s), i.e. the day-range of a solitary individual uninfluenced by competition from conspecifics. We test this exploitation competition hypothesis among primates and carnivores, using data on populations, species and genera. Food density was indexed by population density. Where DR_s could be estimated by regression it was found to be correlated with observed day-range (DR_m). DR_s was therefore indexed by DR_m in all species. Population density and DR_m were associated with each other, but in a multiple regression each variable contributed independently to explaining variation in group size. PGS was predicted better than FGS, but regressions involving either measure of group size were significant in all analyses. The multiple regression analyses were validated by the method of linear contrasts, which accounts for possible lack of independence among taxa. We conclude that species differences in group size are influenced by both food density and DR_s. This suggests that variation in the intensity of exploitation competition is partly responsible for differences in group size.Correspondence to: R.W. Wrangham     

Primmorphs formation dynamics: a screening among Mediterranean sponges

We performed a screening on 21 Mediterranean sponge species in order to know from which ones primmorphs can be obtained, with a comparison of their formation among the outputs. Thirteen species produced primmorphs, evidencing a high variability among species concerning the number and size of primmorphs, without evidencing any taxonomical specificity. Additionally, the formation process and trend of growth during 1 month of monitoring were studied from a quantitative point of view. The dynamics of primmorphs formation follow different patterns: slow and quick formation, short and long-time survival and in the case of long-time survival, the fission and fusion models are proposed. Besides these patterns, each analysed species showed characteristic trends in the time required for the aggregation of cells, the time required for the production of small primmorphs, growth fluctuation and the acquirement of a final stable size.     

A trophic cascade triggers collapse of a salt-marsh ecosystem with intensive recreational fishing

Overexploitation of predators has been linked to the collapse of a growing number of shallow-water marine ecosystems. However, salt-marsh ecosystems are often viewed and managed as systems controlled by physical processes, despite recent evidence for herbivore-driven die-off of marsh vegetation. Here we use field observations, experiments, and historical records at 14 sites to examine whether the recently reported die-off of northwestern Atlantic salt marshes is associated with the cascading effects of predator dynamics and intensive recreational fishing activity. We found that the localized depletion of top predators at sites accessible to recreational anglers has triggered the proliferation of herbivorous crabs, which in turn results in runaway consumption of marsh vegetation. This suggests that overfishing may be a general mechanism underlying the consumer-driven die-off of salt marshes spreading throughout the western Atlantic. Our findings support the emerging realization that consumers play a dominant role in regulating marine plant communities and can lead to ecosystem collapse when their impacts are amplified by human activities, including recreational fishing.     

Effects of combining an intraguild predator with a cannibalistic intermediate predator on a species-level trophic cascade

A greater diversity of natural enemies can in some cases disrupt prey suppression, particularly when natural enemies engage in intraguild predation, where natural enemies compete with and prey upon each other. However, empirical studies have often demonstrated enhanced prey suppression despite intraguild predation. A recent theoretical study proposed the hypothesis that, when the intermediate predator is cannibalistic, intraguild predation can reduce cannibalism within the intermediate predator population, leading to little change in intermediate predator mortality and thus enhanced prey suppression. The goal of this study was to examine this hypothesis empirically. Two summer-long field enclosure experiments were conducted in cotton fields. We investigated the effects of adding an intraguild predator, Zelus renardii, on (1) the abundance of a cannibalistic intermediate predator, Geocoris pallens, (2) the abundance of a herbivore, Lygus hesperus, and (3) cotton plant performance. G. pallens adult abundance did not increase, even when food availability was high and natural enemies were absent, suggesting that density-dependent cannibalism imposes an upper limit on its densities. Furthermore, although Z. renardii is an intraguild predator of G. pallens, G. pallens long-term densities were unaffected by Z. renardii. In the presence of the intermediate predator, the addition of the intraguild predator Z. renardii enhanced suppression of L. hesperus, and there were suggestions that Z. renardii and G. pallens partitioned the L. hesperus population. Effects of herbivore suppression cascaded to the plant level, improving plant performance. In conclusion, we provide empirical support for the hypothesis that the addition of an intraguild predator may enhance prey suppression if the intermediate predator expresses density-dependent cannibalism. Intraguild predation and cannibalism co-occur in many communities; thus their joint effects may be broadly important in shaping predator effects on herbivores and plant performance.     

Interactions among plant defense compounds: a method for analysis

Summary. Plants contain an enormous diversity and quantity of secondary metabolites, some of which are toxic and deterrent to herbivores and pathogens. This impressive diversity of plant compounds suggests a high probability of interactions among them. Synergistic interactions are those in which the combined activity of two or more chemicals is greater than that expected given their individual activities. On the other hand, antagonistic interactions are those in which the combined activity of two or more chemicals is less than that expected given their individual activities. Synergistic interactions could increase plant fitness whereas antagonistic interactions could decrease plant fitness. Interactions are thus potentially very important, not only in explaining the diversity of defense compounds within individual plants, but also in providing insight into plant defense strategies. Although synergistic interactions have received increased attention in the ecological literature in the last decade, the number of documented cases of synergy remains small and antagonistic interactions are rarely considered. The primary reason for this scarcity may be the difficulty of detecting, analyzing and displaying such interactions. Analysis by ANOVA, though sometimes used, often is not appropriate. We introduce a simple technique, isobolographic analysis, that is used in pharmacology for detecting and rigorously quantifying synergy and antagonism and provide an example using the brine shrimp toxicity assay. More statistically sophisticated approaches, such as isobolographic analysis, will allow ecologists to effectively document the role of chemical synergy and antagonism in interactions between species. Such chemical interactions may ultimately provide insight into longstanding, ecological questions. Received 26 October 1998; accepted 24 March 1999.     

Suppression of soybean aphid by generalist predators results in a trophic cascade in soybeans.

Top-down regulation of herbivores in terrestrial ecosystems is pervasive and can lead to trophic cascades that release plants from herbivory. Due to their relatively simplified food webs, agroecosystems may be particularly prone to trophic cascades, a rationale that underlies biological control. However, theoretical and empirical studies show that, within multiple enemy assemblages, intraguild predation (IGP) may lead to a disruption of top-down control by predators. We conducted a factorial field study to test the separate and combined effects of predators and parasitoids in a system with asymmetric IGP. Specifically we combined ambient levels of generalist predators (mainly Coccinellidae) of the soybean aphid, Aphis glycines Matsumura, with controlled releases of the native parasitoid Lysiphlebus testaceipes (Cresson) and measured their impact on aphid population growth and soybean biomass and yield. We found that generalist predators provided strong, season-long aphid suppression, which resulted in a trophic cascade that doubled soybean biomass and yield. However, contrary to our expectations, L. testaceipes provided minor aphid suppression and only when predators were excluded, which resulted in nonadditive effects when both groups were combined. We found direct and indirect evidence of IGP, but because percentage parasitism did not differ between predator exclusion and ambient predator treatments, we concluded that IGP did not disrupt parasitism during this study. Our results support theoretical predictions that intraguild predators which also provide strong herbivore suppression do not disrupt top-down control of herbivores.     

Optimal Allocation of Resources among Threatened Species: a Project Prioritization Protocol

Abstract: Conservation funds are grossly inadequate to address the plight of threatened species. Government and conservation organizations faced with the task of conserving threatened species desperately need simple strategies for allocating limited resources. The academic literature dedicated to systematic priority setting usually recommends ranking species on several criteria, including level of endangerment and metrics of species value such as evolutionary distinctiveness, ecological importance, and social significance. These approaches ignore 2 crucial factors: the cost of management and the likelihood that the management will succeed. These oversights will result in misallocation of scarce conservation resources and possibly unnecessary losses. We devised a project prioritization protocol (PPP) to optimize resource allocation among New Zealand's threatened‐species projects, where costs, benefits (including species values), and the likelihood of management success were considered simultaneously. We compared the number of species managed and the expected benefits gained with 5 prioritization criteria: PPP with weightings based on species value; PPP with species weighted equally; management costs; species value; and threat status. We found that the rational use of cost and success information substantially increased the number of species managed, and prioritizing management projects according to species value or threat status in isolation was inefficient and resulted in fewer species managed. In addition, we found a clear trade‐off between funding management of a greater number of the most cost‐efficient and least risky projects and funding fewer projects to manage the species of higher value. Specifically, 11 of 32 species projects could be funded if projects were weighted by species value compared with 16 projects if projects were not weighted. This highlights the value of a transparent decision‐making process, which enables a careful consideration of trade‐offs. The use of PPP can substantially improve conservation outcomes for threatened species by increasing efficiency and ensuring transparency of management decisions.     

Chronic renal failure among farm families in cascade irrigation systems in Sri Lanka associated with elevated dietary cadmium levels in rice and freshwater fish (Tilapia)

Chronic renal failure (CRF), in the main agricultural region under reservoir based cascade irrigation in Sri Lanka has reached crisis proportion. Over 5,000 patients in the region are under treatment for CRF. The objective of this study is to establish the etiology of the CRF. Concentrations of nine heavy metals were determined in sediments, soils of reservoir peripheries, water and Nelumbo nucifera (lotus) grown in five major reservoirs that supply irrigation water. All five reservoirs carried higher levels of dissolved cadmium (Cd), iron (Fe) and lead (Pb). Dissolved Cd in reservoir water ranged from 0.03 to 0.06 mg/l. Sediment Cd concentration was 1.78-2.45 mg/kg. No arsenic (As) was detected. Cd content in lotus rhizomes was 253.82 mg/kg. The Provisional Tolerable Weekly Intake (PTWI) of Cd based on extreme exposure of rice is 8.702-15.927 mug/kg body weight (BW) for different age groups, 5-50 years. The PTWI of Cd due to extreme exposure of fish is 6.773-12.469 mug/kg BW. The PTWI on a rice staple with fish is 15.475-28.396 mug/kg BW. The mean urinary cadmium (UCd) concentration in CRF patients of age group 40-60 years was 7.58 mug Cd/g creatinine and in asymptomatic persons UCd was 11.62 mug Cd/g creatinine, indicating a chronic exposure to Cd. The possible source of Cd in reservoir sediments and water is Cd-contaminated agrochemicals. The CRF prevalent in north central Sri Lanka is a result of chronic dietary intake of Cd, supported by high natural levels of fluoride in drinking water, coupled with neglecting of routine de-silting of reservoirs for the past 20 years.     

Philopatry, male presence and grooming reciprocation among female primates: a comparative perspective

Competition for food and a safe location in a group are considered to be the main determinants of variation in social relationships among female primates. The effect of the presence of males is usually overlooked, however. Here we put forward two hypotheses connecting the (relative) number of males in a group, a statistic measuring the strength of positive relationships among females (the degree of reciprocity of grooming) and female residence. Under the first hypothesis, we assumed that philopatric females suffer especially from competition for males. Because females of these species are restrained to their native group and thus do not move to groups where better conditions (i.e. more males) prevail, a higher socionomic sex ratio would imply reduced competition and this should be reflected in better relationships among females (i.e. in a higher degree of reciprocation). The second hypothesis is based on the contention of Wrangham (1987) that groups with more males are better able to defend large food sources: in these groups female relationships would suffer less from within-group competition for food. We therefore also expected a positive correlation between the absolute number of males and grooming reciprocation. To test the hypotheses, matrices of grooming interactions were collected by going through the primatological literature. For ten female-resident and four female-transfer primate species, the degree of grooming reciprocation was quantified using a specially adapted matrix statistic. The results favoured the first (competition for males) hypothesis. The degree of grooming reciprocation among females was positively correlated with socionomic sex ratio, but neither with group size nor with the absolute number of males. Correlations between the degree of reciprocity and sex ratio were confirmed at three levels: within the same group at different points in time, between groups of the same species, and between species. In addition, the degree of reciprocation increased with sex ratio more strongly in typical single/variable male than in multi-male species. This we interpreted as a direct consequence of the lower social availability of males for females in multi-male groups than single-male groups. In turn, this effect may be due to males competing for females in multi-male groups. Received: 28 May 1997 / Accepted after revision: 9 November 1997     

A model for leveling coalitions among primate males: toward a theory of egalitarianism

We present a simple model of within-group leveling coalitions among male primates. The model assumes that the value of the coalition is the sum of the payoffs of its members, that the individuals payoff is monotonically decreasing with its rank and that coalitions do not cause rank changes. It predicts that mainly mid- to low rankers engage in leveling coalitions, and that most coalition partners are of adjacent ranks. These predictions agree reasonably well with observations in nature. The model also makes the novel predictions that leveling coalitions are found where male mating competition has only a moderate contest component, and that male dominance ranks will become poorly differentiated where leveling coalitions are frequent. Both these predictions are consistent with observations on groups of macaques and baboons. The model also may account for leveling coalitions among egalitarian human foragers, without making additional assumptions about special human capabilities.