Modeling the impact of feeding stations on vulture scavenging service efficiency

Vultures, the only vertebrate obligate scavengers, are currently facing a dramatic worldwide decline with over half of vulture species now classified as threatened. To address this widespread decline, the use of feeding stations has been widely advocated in recovery programs. However, providing food that is more predictable in time and space than natural resources could disrupt the ecological scavenging service provided by vultures. In this study, we build a multi-agent system (MAS), which describes the social foraging behavior of Gyps vultures in order to investigate how resource predictability in space and resource management affect scavenging service efficiency. We study the possibility that vultures take into account feeding station location in addition of using local enhancement. Our results show that the efficiency of the scavenging service is heavily affected by the way resources are distributed between feeding stations and natural areas. Nevertheless, it appears possible to minimize a loss in service efficiency by increasing the number of feeding stations while keeping the total amount of resources available constant, thus reducing the predictability of the resources located on the stations. We illustrate our work in the case of European feeding stations that provide supplementary resources in areas where natural ones are scarce. Moreover, we discuss the implications worldwide, in particular in the case of India, where feeding stations for vultures are intended to lure individuals away from potentially harmful natural carrion.     

Effects of vulture declines on facultative scavengers and potential implications for mammalian disease transmission

Vultures (Accipitridae and Cathartidae) are the only known obligate scavengers. They feed on rotting carcasses and are the most threatened avian functional group in the world. Possible effects of vulture declines include longer persistence of carcasses and increasing abundance of and contact between facultative scavengers at these carcasses. These changes could increase rates of transmission of infectious diseases, with carcasses serving as hubs of infection. To evaluate these possibilities, we conducted a series of observations and experimental tests of the effects of vulture extirpation on decomposition rates of livestock carcasses and mammalian scavengers in Kenya. We examined whether the absence of vultures changed carcass decomposition time, number of mammalian scavengers visiting carcasses, time spent by mammals at carcasses, and potential for disease transmission at carcasses (measured by changes in intraspecific contact rates). In the absence of vultures, mean carcass decomposition rates nearly tripled. Furthermore, the mean number of mammals at carcasses increased 3-fold (from 1.5 to 4.4 individuals/carcass), and the average time spent by mammals at carcasses increased almost 3-fold (from 55 min to 143 min). There was a nearly 3-fold increase in the mean number of contacts between mammalian scavengers at carcasses without vultures. These results highlight the role of vultures in carcass decomposition and level of contact among mammalian scavengers. In combination, our findings lead us to hypothesize that changes in vulture abundance may affect patterns of disease transmission among mammalian carnivores.     

Bearded vultures (<Emphasis Type="Italic">Gypaetus barbatus</Emphasis>) prefer fatty bones

In animal species, prey processing and the provisioning of nutrients are subject to several constraints related with finding, ingesting and processing food. In most bird species, these constraints are obvious as a consequence of food morphology. In the case of the bearded vulture (Gypaetus barbatus), in comparison with other species, its behavioural and physiological adaptations apparently allow this vulture to ingest bone remains irrespective of their morphology. Here, by comparing bones delivered to the nest to be consumed (selected) and remains found at an experimental feeding station and at bone-breaking sites or ossuaries (rejected), I tested whether bearded vultures are capable of choosing from among the various anatomical parts of an animal carcass in relation to their fatty acid content (nutrient concentration hypothesis), their size (width-reduction hypothesis) or both. The results suggest that bearded vultures prefer the fatty anatomical parts (with a high percentage of oleic acid) of an animal carcass regardless of bone length, although bone morphology as a consequence of handling efficiency or the ingestion process may also play a secondary role in food selection. The close association between the bones selected and their high fat value implies an optimisation of foraging time and of the increased energy gained from the food. This is in line with selective foraging to redress specific nutritional imbalances (nutrient concentration hypothesis) and, secondarily, the width-reduction hypothesis.     

Priority areas for conservation of Old World vultures

The prosperity and well-being of human societies relies on healthy ecosystems and the services they provide. However, the biodiversity crisis is undermining ecosystems services and functions. Vultures are among the most imperiled taxonomic groups on Earth, yet they have a fundamental ecosystem function. These obligate scavengers rapidly consume large amounts of carrion and human waste, a service that may aid in both disease prevention and control of mammalian scavengers, including feral dogs, which in turn threaten humans. We combined information about the distribution of all 15 vulture species found in Europe, Asia, and Africa with their threats and used detailed expert knowledge on threat intensity to prioritize critical areas for conserving vultures in Africa and Eurasia. Threats we identified included poisoning, mortality due to collision with wind energy infrastructures, and other anthropogenic activities related to human land use and influence. Areas important for vulture conservation were concentrated in southern and eastern Africa, South Asia, and the Iberian Peninsula, and over 80% of these areas were unprotected. Some vulture species required larger areas for protection than others. Finally, countries that had the largest share of all identified important priority areas for vulture conservation were those with the largest expenditures related to rabies burden (e.g., India, China, and Myanmar). Vulture populations have declined markedly in most of these countries. Restoring healthy vulture populations through targeted actions in the priority areas we identified may help restore the ecosystem services vultures provide, including sanitation and potentially prevention of diseases, such as rabies, a heavy burden afflicting fragile societies. Our findings may guide stakeholders to prioritize actions where they are needed most in order to achieve international goals for biodiversity conservation and sustainable development.     

Suspected flunixin poisoning of a wild Eurasian Griffon Vulture from Spain

Exposure to residues of the nonsteroidal anti‐inflammatory drug (NSAID) diclofenac present in livestock carcasses has caused extensive declines in 3 Gyps vulture species across Asia. The carcass of a wild Eurasian Griffon Vulture (Gyps fulvus) was found in 2012 on an Andalucian (Spain) game hunting reserve and examined forensically. The bird had severe visceral gout, a finding consistent with Gyps vultures from Asia that have been poisoned by diclofenac. Liver and kidney samples from this Eurasian Griffon Vulture contained elevated flunixin (an NSAID) levels (median = 2.70 and 6.50 mg/kg, respectively). This is the first reported case of a wild vulture being exposed to and apparently killed by an NSAID outside Asia. It is also the first reported instance of mortality in the wild resulting from environmental exposure to an NSAID other than diclofenac. Caso de Sospecha de Envenenamiento por Flunixin de un Buitre Leonado en España     

Social dominance in migrant and resident turkey vultures at carcasses: evidence for a despotic distribution?

We placed carcasses in three different vegetation types in the heterogeneous savannas of central Venezuela to investigate the role of social dominance in habitat use by flocking migrant and resident turkey vultures (Cathartes aura meridionalis and C. a. ruficollis). Migrants foraged primarily in savanna habitats while residents foraged almost exclusively in gallery forest. In the gallery forest residents discovered carrion first significantly more often than migrants, despite there being equal densities of residents and migrants foraging over this habitat. Because residents fed in smaller groups than migrants at carcasses they had higher feeding rates. There was also a negative relationship between group sizes of residents and migrants. The feeding rate of residents declined in response to increased group size of migrants, but group size of residents had no effect. Migrant group size also had a greater effect on resident feeding rates than king vulture presence or absence. When the effect of migrant and resident group size on feeding rates in migrants was compared, the most significant factor was migrant group size. A second analysis showed that both resident group size and presence or absence of king vultures had a significant effect on feeding rates in migrants. Rates of agonistic encounters in migrant and resident turkey vultures increased weakly in relation to group size. However, there was an increase in residents' encounter rate with migrants in relation to increased migrant group size; there was no difference in resident encounter rates with other residents in relation to resident group size. Migrants dominated residents in almost all agonistic interactions over carcasses. We suggest that savanna habitats were less attractive to residents for foraging because they held larger groups of migrants.     

The causes and consequences of partial prey consumption by wolves preying on moose

For a wide range of taxa, partial prey consumption (PPC) is a frequent occurrence. PPC may arise from physiological constraints to gut capacity or digestive rate. Alternatively, PPC may represent an optimal foraging strategy. Assessments that clearly distinguish between these causes are rare and have been conducted only for invertebrate species that are ambush predators with extra-intestinal digestion (e.g., wolf spiders). We present the first strong test for the cause of PPC in a cursorial vertebrate predator with intestinal digestion: wolves (Canis lupus) feeding on moose (Alces alces). Previous theoretical assessments indicate that if PPC represents an optimal foraging strategy and is not caused by physiological limitations, then mean carcass utilization is negatively correlated with mean kill rate and the utilization of individual carcasses is uncorrelated with time between kills. Wolves exhibit exactly this pattern. We explore how the typical portrayal of PPC by wolves has been not only misleading but also detrimental to conservation by promoting negative attitudes toward wolves.     

Livestock carrying capacity evaluation in an integrated farming system: A case study from the mid-hills of Nepal

The crop livestock integrated farming system practiced in hilly regions of developing countries largely depends on the ecosystem as a whole. More especifically, the livestock component of the farming system relies heavily on natural resources such as forest, grazing and agricultural land for the supply of feed and fodder. The importance of animals as agents of nutrient recycling, sources of rural energy in terms of draft power and fuel, as well as being major contributors to the farm economy, has resulted in an increased population of ruminants in these regions, creating a threat to the sustainability and productivity of these land resources. This paper is an attempt to evaluate the livestock carrying capacity of land resources and to formulate the optimum herd size compatible with the differently resourced farm categories within the sub-watershed region of the mid-hills in Nepal. Our analysis reveals that the livestock carrying capacity of the land resources at the watershed level is 11696 Livestock Units (LU), whereas current stocking is 12985 LU, suggesting an overstocking of 1289 LU at the watershed level. Total feed supply from different sources is 12668 mt/yr whereas current Total Digestive Nutrient (TDN) demand is 14060 mt/yr, a negative balance of 1393 mt/yr. The excess livestock at the sub-watershed level directly implies excess livestock holding, producing a poor nutritional status for the farm household. Linear programming analysis reveals that the farmers of large, medium and small category farms can optimize their livestock holding by a combination of 3 LU buffaloes and 4 LU goats, 2 LU buffaloes and 4 LU goats and 1 LU buffaloes and 4.4 LU goats, thus giving maximum return to the farm family without exerting pressure on fragile natural resources.     

Can we measure carrying capacity with foraging behavior?

Carrying capacity is one of the most important, yet least understood and rarely estimated, parameters in population management and modeling. A simple behavioral metric of carrying capacity would advance theory, conservation, and management of biological populations. Such a metric should be possible because behavior is finely attuned to variation in environment including population density. We connect optimal foraging theory with population dynamics and life history to develop a simple model that predicts this sort of adaptive density-dependent change in food consumption. We then confirm the model's unexpected and manifold predictions with field experiments. The theory predicts reproductive thresholds that alter the marginal value of energy as well as the value of time. Both effects cause a pronounced discontinuity in quitting-harvest rate that we revealed with foraging experiments. Red-backed voles maintained across a range of high densities foraged at a lower density-dependent rate than the same animals exposed to low-density treatments. The change in harvest rate is diagnostic of populations that exceed their carrying capacity. Ecologists, conservation biologists, and wildlife managers may thus be able to use simple and efficient foraging experiments to estimate carrying capacity and habitat quality.     

Intraspecific variation in a predator affects community structure and cascading trophic interactions

Intraspecific phenotypic variation in ecologically important traits is widespread and important for evolutionary processes, but its effects on community and ecosystem processes are poorly understood. We use life history differences among populations of alewives, Alosa pseudoharengus, to test the effects of intraspecific phenotypic variation in a predator on pelagic zooplankton community structure and the strength of cascading trophic interactions. We focus on the effects of differences in (1) the duration of residence in fresh water (either seasonal or year-round) and (2) differences in foraging morphology, both of which may strongly influence interactions between alewives and their prey. We measured zooplankton community structure, algal biomass, and spring total phosphorus in lakes that contained landlocked, anadromous, or no alewives. Both the duration of residence and the intraspecific variation in foraging morphology strongly influenced zooplankton community structure. Lakes with landlocked alewives had small-bodied zooplankton year-round, and lakes with no alewives had large-bodied zooplankton year-round. In contrast, zooplankton communities in lakes with anadromous alewives cycled between large-bodied zooplankton in the winter and spring and small-bodied zooplankton in the summer. In summer, differences in feeding morphology of alewives caused zooplankton biomass to be lower and body size to be smaller in lakes with anadromous alewives than in lakes with landlocked alewives. Furthermore, intraspecific variation altered the strength of the trophic cascade caused by alewives. Our results demonstrate that intraspecific phenotypic variation of predators can regulate community structure and ecosystem processes by modifying the form and strength of complex trophic interactions.     

Agro-biodiversity in rice–wheat-based agroecosystems of eastern Uttar Pradesh,India: implications for conservation and sustainable management

In this study, we analysed the indigenous practices of agro-biodiversity, changing patterns of cropping systems and conservation of agriculturally important natural resources of rice–wheat-based agroecosystems. The research was carried out in three villages of Azamgarh district, eastern Uttar Pradesh, India. Participatory approaches and personal interviews were combined to collect the data from 180 farmers. The results indicated that resource-poor (RP) farmers, by applying traditional practices of natural resource conservation, outperform their rich (RR) counterparts in conserving agro-biodiversity. Increase in irrigation facilities, adoption of improved crop varieties, commercialisation of farming and socio-economic changes have threatened the agro-biodiversity and related natural resources of these agroecosystems. For most of the crops, the RP farmers had better production stability as compared to the RR ones because the former tend to apply more sustainable and ecologically sound indigenous practices. A negative growth rate was observed in the area/numbers and production of legumes, livestock, pasture lands and community forestry. Women, who are the local custodians of conserving indigenous biodiversity (uncultivated species) and crop varieties through localised knowledge and sociocultural institutions, had significantly higher knowledge on agro-biodiversity than men. We concluded that integration of farmers’ eco-friendly practices with formal science and policy-making in a participatory mode may enhance sustainability of functions of agroecosystem leading to the strengthening of adaptive practices in changing climate and socio-economic scenario.     

Chihuahuan Desert kangaroo rats: nonlinear effects of population dynamics, competition, and rainfall

Using long-term data on two kangaroo rats in the Chihuahuan Desert of North America, we fitted logistic models including the exogenous effects of seasonal rainfall patterns. Our aim was to test the effects of intraspecific interactions and seasonal rainfall in explaining and predicting the numerical fluctuations of these two kangaroo rats. We found that logistic models fit both data sets quite well; Dipodomys merriami showed lower maximum per capita growth rates than Dipodomys ordii, and in both cases logistic models were nonlinear. Summer rainfall appears to be the most important exogenous effect for both rodent populations; models including this variable were able to predict independent data better than models including winter rainfall. D. merriami was also negatively affected by another kangaroo rat (Dipodomys spectabilis), consistent with previous experimental evidence. We hypothesized that summer rainfall influences the carrying capacity of the environment by affecting seed availability and the intensity of intraspecific competition.     

Time stress,predation risk and diurnal–nocturnal foraging trade-offs in larval prey

Insect larvae increase in size with several orders of magnitude throughout development making them more conspicuous to visually hunting predators. This change in predation pressure is likely to impose selection on larval anti-predator behaviour and since the risk of detection is likely to decrease in darkness, the night may offer safer foraging opportunities to large individuals. However, forsaking day foraging reduces development rate and could be extra costly if prey are subjected to seasonal time stress. Here we test if size-dependent risk and time constraints on feeding affect the foraging–predation risk trade-off expressed by the use of the diurnal–nocturnal period. We exposed larvae of one seasonal and one non-seasonal butterfly to different levels of seasonal time stress and time for diurnal–nocturnal feeding by rearing them in two photoperiods. In both species, diurnal foraging ceased at large sizes while nocturnal foraging remained constant or increased, thus larvae showed ontogenetic shifts in behaviour. Short night lengths forced small individuals to take higher risks and forage more during daytime, postponing the shift to strict night foraging to later on in development. In the non-seasonal species, seasonal time stress had a small effect on development and the diurnal–nocturnal foraging mode. In contrast, in the seasonal species, time for pupation and the timing of the foraging shift were strongly affected. We argue that a large part of the observed variation in larval diurnal–nocturnal activity and resulting growth rates is explained by changes in the cost/benefit ratio of foraging mediated by size-dependent predation and time stress.     

Interacting Effects of Landownership, Land Use, and Endangered Species on Conservation of Southwestern U.S. Rangelands

Abstract:  The contemporary southwestern United States is characterized by fire-adapted ecosystems; large numbers of federally listed threatened and endangered species; a patchwork of federal, state, and private landownership; and a long history of livestock grazing as the predominant land use. I compared eight sites in southern Arizona and New Mexico to assess the interacting effects of these characteristics on conservation practices and outcomes. There was widespread interest and private-sector leadership in restoring fire to southwestern rangelands, and there is a shortage of predictive scientific knowledge about the effects of fire and livestock grazing on threatened and endangered species. It was easier to restore fire to lands that were either privately owned or not grazed, in part because of obstacles created by threatened and endangered species on grazed public lands. Collaborative management facilitated conservation practices and outcomes, and periodic removal of livestock may be necessary for conservation, but permanent livestock exclusion may be counterproductive because of interactions with land-use and landownership patterns.     

Metabolism of aceclofenac in cattle to vulture‐killing diclofenac

The nonsteroidal anti‐inflammatory drug (NSAID) diclofenac is highly toxic to Gyps vultures, and its recent widespread use in South Asia caused catastrophic declines in at least 3 scavenging raptors. The manufacture of veterinary formulations of diclofenac has since been banned across the region with mixed success. However, at least 12 other NSAIDs are available for veterinary use in South Asia. Aceclofenac is one of these compounds, and it is known to metabolize into diclofenac in some mammal species. The metabolic pathway of aceclofenac in cattle, the primary food of vultures in South Asia, is unknown. We gave 6 cattle the recommended dose of aceclofenac (2 mg/kg), collected blood thereafter at intervals for up to 12 h, and used liquid chromatography with mass spectrometry in a pharmacokinetic analysis of aceclofenac and diclofenac in the plasma. Nearly all the aceclofenac administered to the cattle was very rapidly metabolized into diclofenac. At 2 h, half the aceclofenac had been converted into diclofenac, and at 12 h four‐fifths of the aceclofenac had been converted into diclofenac. Therefore, administering aceclofenac to livestock poses the same risk to vultures as administering diclofenac to livestock. This, coupled with the risk that aceclofenac may replace diclofenac in the veterinary market, points to the need for an immediate ban on all aceclofenac formulations that can be used to treat livestock. Without such a ban, the recovery of vultures across South Asia will not be successful.     

Turkey vultures foraging at experimental food patches: a test of information transfer at communal roosts

Summary Experimental food patches were used to assess the importance of food information transfer in communally roosting turkey vultures (Cathartes aura) in southern Ontario. Feeding trials failed to provide evidence of recruitment due to information transfer. Overall, fewer birds arrived at novel food patches on the days following discovery than was expected, had information transfer been operating. Earlier arrivals on second days were more likely due to local enhancement rather than information transfer since the size of groups arriving at food patches did not differ between the 2 days. These results indicate that turkey vulture roosts in Ontario do not operate as centers for food information transfer. Intraspecific competition, preferential selection of small-sized carrion, and a low degree of kin association make the proposed benefits of information transfer much less applicable to turkey vultures than has been found for other scavenging species.Offprint requests to: P.J. Weatherhead     

Functional Equivalency between Rice Fields and Seminatural Wetland Habitats

Abstract: Evaluating the potential for anthropogenic habitats to act as surrogates for the natural habitats they replace is a key issue in conservation biology. In California, flooded rice fields are used by numerous aquatic birds during winter. If this habitat functions similarly to more natural wetlands, increased flooding may help replace the extensive wetlands that occurred in the region prior to agricultural development. I tested whether food abundance, perceived predation threat, foraging performance, and the way in which birds allocate their time to different behaviors differed between flooded rice fields and seminatural wetlands for several species of aquatic bird. When appropriate, I also compared flooded and unflooded fields. Invertebrate densities did not differ among habitats. Seminatural wetlands had less rice grain but more seeds from other plants than the two rice habitats. The frequency with which predators passed over a feeding area was lower in flooded fields than in unflooded fields or seminatural wetlands. Most differences in feeding performance and time allocation among habitats were small and statistically insignificant. For some species, feeding efficiency was greater in seminatural wetlands than in flooded fields. Increasing attack rates and the amount of time spent feeding when in flooded fields, however, may allow birds to compensate for reduced efficiency. Multivariate analyses showed that group size, predation threat, time of day, date, and water depth often were associated with behaviors, but that these variables rarely accounted for habitat differences. Flooded fields apparently provide equivalent foraging habitat to seminatural wetlands and, because of reduced predation threat, may be a safer habitat for waterbirds. Thus, if managed appropriately, one of the world's dominant forms of agriculture can provide valuable waterbird habitat.     

Males are faster foragers than females: intersexual differences of foraging behaviour in the Apennine chamois

Availability of food resources and individual characteristics can influence foraging behaviour, which can differ between males and females, leading to different patterns of food/habitat selection. In dimorphic species, females are usually more selective in food choice, show greater bite rates and spend more time foraging than males. We evaluated sexual differences in foraging behaviour in Apennine chamois Rupicapra pyrenaica ornata, during the warm season, before the rut. Both sexes selected nutritious vegetation patches and spent a comparable amount of time feeding. However, males had a significantly greater feeding intensity (bite rate) and a lower search effort for feeding (step rate), as well as they spent more time lying down than females. Females selected foraging sites closer to refuge areas than males. In chamois, sexual size dimorphism is seasonal, being negligible in winter–spring, but increasing to 30–40 % in autumn. Our results suggest that males enhance their energy and mass gain by increasing their food intake rate during the warm season, to face the costs of the mating season (November). Conversely, females seem to prioritize a fine-scale selection of vegetation and the protection of offspring. A great food intake rate of males in the warm season could have developed as a behavioural adaptation leading herbivores to the evolutionary transition from year-round monomorphism to permanent dimorphism, through seasonal dimorphism.     

Nutritional constraints on mountain baboons (Papio ursinus): Implications for baboon socioecology

Summary Populations of baboon (Papio sp.) at geographic and climatic extremes for the genus show a tendency to one-male organization, whereas most baboons live in multimale social groups; this effect has been attributed largely to limitation of food supply, but baboons' complex diet has hindered proper nutritional analyses. To test these optimal-diet explanations of social variation, we quantified intake and used phytochemical analysis of foods to compare the nutrition, during seasonal changes, of two groups of mountain baboons (P. ursinus) living at different altitudes of a continuous grassland habitat. The majority of plant foods were eaten uniquely by one or other group, though their altitudinal separation was only 400 m, and the time budget of feeding choices varied with age-sex class as well as season. Converting to a common currency of nutrients reveals that baboons gained the same yield from a unit time spent foraging (whether this is measured in edible dry weight, or simply protein) in both groups, despite their differing mean altitude, whereas seasonal variation was large and statistically significant. Increased feeding time at the winter bottleneck made no effective compensation for the poorer food yields: in late winter there was a minimum for daily nutrient gain at both altitudes. Apparently this population is already at an extreme for the time animals devote to foraging in winter, when they rely on inconspicuous and slow-to-harvest swollen shoot bases and underground plant storage organs. Since an individual's nutrient yield does not vary with altitude, we conclude that socioecological parameters are effectively optimized for feeding. Since contest competition is absent, this adjustment of foraging efficiency is largely through the effect of differential density on scramble competition. Differences in social structure are considered to be a secondray consequence of optimal foraging, mediated through altitudinal variation in either population density or in day range limits.     

Dominance in feeding territories relates to foraging success and offspring growth in brown skuas <Emphasis Type="Italic">Catharacta antarctica lonnbergi</Emphasis>

Territoriality should lead to strict dominance, as territory holders typically control access to resources and exclude others from their use. In feeding territories, dominance should be reflected in foraging success and ultimately in reproduction differences; however, these successive links have rarely been made explicit. Therefore, we investigated a population of brown skuas Catharacta antarctica lonnbergi, in which only part of the breeding population occupied feeding territories within penguin colonies. We identified the dominance hierarchy and determined the foraging success of the participants in fights for access to penguin carcasses within the territories. Furthermore, we monitored offspring growth from parents with and without feeding territories. Our results indicated a clear dominance hierarchy with territorial birds in their own territory dominating over territorial breeders from other territories, non-territorial breeders and non-breeding birds. However, territory owners could not completely exclude others from access to food. Foraging success was positively related to dominance scores: The dominant territory owners received 63% of a carcass, whereas non-territorial pairs could get less than 10%. The link between foraging success and offspring development was less clear: Although male chicks of non-territorial parents suffered from lower growth rates and, thus, delayed fledging, there were no such differences in female chicks. Territoriality in skuas did not imply a complete occupation of food, but guaranteed optimal growth conditions for offspring. Non-territorial individuals were forced to search for alternative resources, and the restricted access to the preferred food resulted in inferior conditions for offspring development, making this foraging strategy less rewarding.