Habitat characteristics from local to landscape scales combine to shape vertebrate scavenging communities

Abiotic and biotic factors modulate carcass consumption by scavengers, affecting ecosystem functioning. Habitat structure is arguably a key factor influencing scavenging, but its role remains poorly understood, particularly at small spatial scales. We examine how habitat characteristics at landscape (50–1000 m radius) and local (≤10 m radius) scales around carrion affect the structure of vertebrate scavenging communities. We used remote cameras to monitor the consumption of 151 ungulate carcasses in one temperate (55 carcasses) and two Mediterranean (56 and 40 carcasses) study areas in Spain in 2011–2013. Our results showed complex habitat–scavenger relationships that mainly relied upon the spatial scale, the type of carcass and the study area. While the response of scavenger richness to habitat characteristics was consistent across study areas, the effects of diversity varied regionally at the landscape scale. Large and medium-sized carcasses in open landscapes had lower scavenger richness, likely because open habitats promote vulture dominance. At the local scale, shrub cover lowered scavenger richness and diversity, hindering carrion location by avian scavengers. Our results suggest that the structure of vertebrate scavenging assemblages at carcasses is driven by carcass and habitat characteristics operating as ecological filters at different scales (i.e. local, landscape, and biogeographical), which affect a species’ ability to locate, access and dominate carrion. Understanding the factors underlying the complex habitat–community relationships shown here has implications for managing key ecosystem functions and services. We propose a multi-scale conceptual framework to disentangle scavenger–carcass relationships.     

Novel Scavenger Removal Trials Increase Wind Turbine—Caused Avian Fatality Estimates

ABSTRACT For comparing impacts of bird and bat collisions with wind turbines, investigators estimate fatalities/megawatt (MW) of rated capacity/year, based on periodic carcass searches and trials used to estimate carcasses not found due to scavenger removal and searcher error. However, scavenger trials typically place ≥10 carcasses at once within small areas already supplying scavengers with carcasses deposited by wind turbines, so scavengers may be unable to process and remove all placed carcasses. To avoid scavenger swamping, which might bias fatality estimates low, we placed only 1–5 bird carcasses at a time amongst 52 wind turbines in our 249.7-ha study area, each carcass monitored by a motion-activated camera. Scavengers removed 50 of 63 carcasses, averaging 4.45 days to the first scavenging event. By 15 days, which corresponded with most of our search intervals, scavengers removed 0% and 67% of large-bodied raptors placed in winter and summer, respectively, and 15% and 71% of small birds placed in winter and summer, respectively. By 15 days, scavengers removed 42% of large raptors as compared to 15% removed in conventional trials, and scavengers removed 62% of small birds as compared to 52% removed in conventional trials. Based on our methodology, we estimated mean annual fatalities caused by 21.9 MW of wind turbines in Vasco Caves Regional Preserve (within Altamont Pass Wind Resource Area, California, USA) were 13 red-tailed hawks (Buteo jamaicensis), 12 barn owls (Tyto alba), 18 burrowing owls (Athene cunicularia), 48 total raptors, and 99 total birds. Compared to fatality rates estimated from conventional scavenger trials, our estimates were nearly 3 times higher for red-tailed hawk and barn owl, 68% higher for all raptors, and 67% higher for all birds. We also found that deaths/gigawatt-hour of power generation declined quickly with increasing capacity factor among wind turbines, indicating collision hazard increased with greater intermittency in turbine operations. Fatality monitoring at wind turbines might improve by using scavenger removal trials free of scavenger swamping and by relating fatality rates to power output data in addition to rated capacity (i.e., turbine size). The resulting greater precision in mortality estimates will assist wildlife managers to assess wind farm impacts and to more accurately measure the effects of mitigation measures implemented to lessen those impacts.     

Carcass size shapes the structure and functioning of an African scavenging assemblage

The particle size of the food resource strongly determines the structure and dynamics of food webs. However, the ecological implications of carcass size variation for scavenging networks structure and functioning have been largely overlooked. Here we investigate differences in scavenging patterns due to carcass size in a complex vertebrate scavenger community, Hluhluwe‐iMfolozi Park, South Africa, while taking into account seasonality. We monitored the consumption of three types of experimental carcasses: ‘small’ (< 10 kg), ‘medium’ (10–100 kg) and ‘large’ (> 100 kg). We employed general lineal models to explore the influence of carcass size on 1) scavenging network structure (scavenger species richness per carcass) and 2) functioning (carcass detection time, consumption time, consumption rate and percentage of carrion consumed). We also tested whether the structure of the scavenging network of each carcass size was nested, i.e. whether the scavenging assemblage in species‐poor carcasses was a subset of the assemblage consuming species‐rich carcasses. We found strong evidence indicating that carcass size is a major factor governing the associated scavenger assemblage. Scavenger species richness per carcass and carcass consumption time and rate increased with carcass size, while carcass detection time and percentage of carrion biomass consumed were negatively related to carcass size. Strikingly, most of the carrion biomass was consumed by facultative scavengers, represented by large mammalian carnivores, rather than by obligate scavengers (i.e. vultures). Scavenging network nestedness tended to be higher at larger carcasses, and nestedness was sensitive to the removal of the most connected species in the network (spotted hyena) rather than vultures. When comparing scavenging and predation assemblages, crucial size‐dependent differences emerge. Also, we identified a traditionally ignored mechanism by which hunting large prey could be relatively less profitable for predators, namely the costs associated with competition from scavengers and decomposers.     

Ecological role of vertebrate scavengers in urban ecosystems in the UK

Recent research has demonstrated how scavenging, the act of consuming dead animals, plays a key role in ecosystem structure, functioning, and stability. A growing number of studies suggest that vertebrate scavengers also provide key ecosystem services, the benefits humans gain from the natural world, particularly in the removal of carcasses from the environment. An increasing proportion of the human population is now residing in cities and towns, many of which, despite being highly altered environments, contain significant wildlife populations, and so animal carcasses. Indeed, non‐predation fatalities may be higher within urban than natural environments. Despite this, the fate of carcasses in urban environments and the role vertebrate scavengers play in their removal have not been determined. In this study, we quantify the role of vertebrate scavengers in urban environments in three towns in the UK. Using experimentally deployed rat carcasses and rapid fire motion‐triggered cameras, we determined which species were scavenging and how removal of carcass biomass was partitioned between them. Of the 63 experimental carcasses deployed, vertebrate scavenger activity was detected at 67%. There was a significantly greater depletion in carcass biomass in the presence (mean loss of 194 g) than absence (mean loss of 14 g) of scavengers. Scavenger activity was restricted to three species, Carrion crows Corvus corone, Eurasian magpies Pica pica, and European red foxes Vulpes vulpes. From behavioral analysis, we estimated that a maximum of 73% of the carcass biomass was removed by vertebrate scavengers. Despite having low species richness, the urban scavenger community in our urban study system removed a similar proportion of carcasses to those reported in more pristine environments. Vertebrate scavengers are providing a key urban ecosystem service in terms of carcass removal. This service is, however, often overlooked, and the species that provide it are among some of the most disliked and persecuted.     

Identification of vertebrate scavengers of small mammal carcasses in a forested landscape

We identified vertebrate scavengers of small mammal carcasses at the 780-km² Savannah River Site during the winter of 2000–2001. Rodent carcasses, differing in size and visual conspicuousness, were placed in upland pine forests and bottomland hardwood forests during six 2-week periods. Sixty-two of the 96 carcasses (65%) were removed by vertebrates. With the aid of remote photography, we identified 11 species of scavengers removing carcasses. RaccoonsProcyon lotor, gray foxesUrocyon cinereoargenteus, and feral pigsSus scrofa scavenged most frequently. The mean elapsed time for carcass removal was 5.6 days. The number of carcasses removed by vertebrates did not differ significantly with respect to carcass size, visual conspicuousness, or habitat type; however, air temperature was strongly correlated (positively) with carcass removal. Our study demonstrates that many mammal species are capable of utilizing small carrion items as a food resource, and suggests that scavenging may account for a higher proportion of the diet of some facultative scavengers than is now widely assumed.     

Deer Carcass Decomposition and Potential Scavenger Exposure to Chronic Wasting Disease

ABSTRACT Chronic wasting disease (CWD) is a transmissible spongiform encephalopathy afflicting the Cervidae family in North America, causing neurodegeneration and ultimately death. Although there are no reports of natural cross-species transmission of CWD to noncervids, infected deer carcasses pose a potential risk of CWD exposure for other animals. We placed 40 disease-free white-tailed deer (Odocoileus virginianus) carcasses and 10 gut piles in the CWD-affected area of Wisconsin (USA) from September to April in 2003 through 2005. We used photos from remotely operated cameras to characterize scavenger visitation and relative activity. To evaluate factors driving the rate of carcass removal (decomposition), we used Kaplan-Meier survival analysis and a generalized linear mixed model. We recorded 14 species of scavenging mammals (6 visiting species) and 14 species of scavenging birds (8 visiting species). Prominent scavengers included American crows (Corvus brachyrhynchos), raccoons (Procyon lotor), and Virginia opossums (Didelphis virginiana). We found no evidence that deer consumed conspecific remains, although they visited gut piles more often than carcasses relative to temporal availability in the environment. Domestic dogs, cats, and cows either scavenged or visited carcass sites, which could lead to human exposure to CWD. Deer carcasses persisted for 18 days to 101 days depending on the season and year, whereas gut piles lasted for 3 days. Habitat did not influence carcass decomposition, but mammalian and avian scavenger activity and higher temperatures were positively associated with faster removal. Infected deer carcasses or gut piles can serve as potential sources of CWD prions to a variety of scavengers. In areas where surveillance for CWD exposure is practical, management agencies should consider strategies for testing primary scavengers of deer carcass material.     

Functional differences in scavenger communities and the speed of carcass decomposition

Carcass decomposition largely depends on vertebrate scavengers. However, how behavioral differences between vertebrate scavenger species, the dominance of certain species, and the diversity of the vertebrate scavenger community affect the speed of carcass decomposition is poorly understood. As scavenging is an overlooked trophic interaction, studying the different functional roles of vertebrate species in the scavenging process increases our understanding about the effect of the vertebrate scavenger community on carcass decomposition. We used motion‐triggered infrared camera trap footages to profile the behavior and activity of vertebrate scavengers visiting carcasses in Dutch nature areas. We grouped vertebrate scavengers with similar functional roles. We found a clear distinction between occasional scavengers and more specialized scavengers, and we found wild boar (Sus scrofa) to be the dominant scavenger species in our study system. We showed that these groups are functionally different within the scavenger community. We found that overall vertebrate scavenger diversity was positively correlated with carcass decomposition speed. With these findings, our study contributes to the understanding about the different functional roles scavengers can have in ecological communities.     

Vertebrate scavenger guild composition and utilization of carrion in an East Asian temperate forest

Scavenging is a common feeding behavior by many species that plays an important role in ecosystem stability and function while also providing ecosystem services. Despite its importance, facultative scavenging on large animal carcasses has generally been overlooked in Asian temperate forest ecosystems. The aim of this study was to determine the composition and feeding behavior of the facultative scavenger guild as it relates to sika deer (Cervus nippon) carcasses in Japanese forests. There are no obligate scavengers or large predators that kill adult ungulates, but humans fill the role of large predators by culling deer for population management. We documented nine vertebrate species scavenging on deer carcasses and found that mammals were more frequent scavengers than birds and also fed for longer durations. This result suggests that there is a facultative scavenger guild composed mainly of mammals in our forest ecosystem and that carcass utilization by birds was restricted to only forest species. Raccoon dogs (Nyctereutes procyonoides) and Asian black bears (Ursus thibetanus) were the most frequent scavenger species and also fed for longer durations than other scavengers. There were significant seasonal differences in scavenging by Asian black bear, Japanese marten (Martes melampus), and mountain hawk‐eagle (Nisaetus nipalensis), suggesting the availability of other food resources may alter facultative scavenging by each species. Our results support that scavenging is widespread in this system and likely has important functions including building links in the food web.     

Ecological links between salmon,large carnivore predation,and scavenging birds

We compared scavenging bird abundance and diversity across 17 estuaries on the Central Coast of British Columbia, Canada in relation to landscape characteristics and biomass of spawning salmon and senescent and depredated salmon carcasses. We discovered that all metrics for spawning salmon and carcass biomasses were strong predictors of scavenger abundance and diversity. Specifically, Shannon’s diversity, which emphasizes rare species richness, and total abundances of scavengers, corvids (Corvus spp.), and small and large gulls (Larus spp.) were most strongly predicted by total biomass of carcasses. In contrast, the abundance of bald eagles Haliaeetus leucocephalus was most strongly predicted by biomass of carcasses that had been killed or scavenged by other predators (mostly bears and wolves). Simpson’s diversity, which emphasizes evenness of common species, was best predicted by total spawning salmon biomass. Estuary area also featured prominently among top predictors of most scavenger metrics. Our results suggest a link between terrestrial salmon predators and bald eagles, and that available salmon biomass is important for maintaining the abundance and diversity of scavenging birds that congregate at estuaries throughout the spawning season.     

Estimating carcass persistence and scavenging bias in a human‐influenced landscape in western Alaska

ABSTRACT We examined variation in persistence rates of waterfowl carcasses placed along a series of transects in tundra habitats in western Alaska. This study was designed to assess the effects of existing tower structures and was replicated with separate trials in winter, summer and fall as both the resident avian population and the suite of potential scavengers varied seasonally. Carcass persistence rates were uniformly low, with <50% of carcasses persisting for more than a day on average. Persistence rate varied by carcass age, carcass size, among transects and was lowest in the fall and highest in the summer. We found little support for models where persistence varied in relation to the presence of tower structures. We interpret this as evidence that scavengers were not habituated to searching for carcasses near these structures. Our data demonstrate that only a small fraction of bird carcasses are likely to persist between searches, and if not appropriately accounted for, scavenging bias could significantly influence bird mortality estimates. The variation that we documented suggests that persistence rates should not be extrapolated among tower locations or across time periods as the variation in carcass persistence will result in biased estimates of total bird strike mortality.     

Foraging risk in scavenging ecology: A study of scavenger behavior and patterns of bacterial growth

Scavenging constitutes an understudied energy pathway in terrestrial ecosystems, with important connections to disease ecology. A prevailing null hypothesis in scavenging ecology is that carcasses serve as a risk-free meal for whatever animal first encounters them on the landscape. We tested this hypothesis by focusing on a suspected risk that scavengers would face at carcasses: the risk of pathogen exposure. We conducted field trials with mouse carcasses in which we manipulated potential cues to pathogen risk and then monitored scavenger foraging decisions. Separately, we studied pathogen and commensal bacteria dynamics within mouse carcasses through time in the laboratory to better understand how carcass age might impact pathogen risk to scavengers. A visual cue to pathogen risk in the field (carcasses deployed in groups of 7 in 1 m²) caused facultative vertebrate scavengers to pass on the opportunity to feed at a rate six-times higher than for carcasses deployed singly (46.9% vs. 7.7%), suggesting an ability to perceive cues to pathogen risk at carcasses. The cues to carcass age, however, produced no effect on facultative scavenger behavior in our field trials. Laboratory trials demonstrated that both commensal enteric bacteria and a known pathogen (Listeria monocytogenes) increased at least through bloat and active decay stages in carcasses, suggesting that cues to carcass age may not help scavengers reduce pathogen risk early in decomposition. In providing evidence counter to the free meal hypothesis, our results support the continued formation of an alternative risk-based framework to understand scavenger behavior at carcasses.     

Scavenger community response to the removal of a dominant scavenger

The alteration of scavenging communities can reduce basic ecosystem services and increase risks to human and wildlife health. Recent work demonstrated that scavenging communities in agricultural landscapes are extremely efficient: superabundant mesopredators sequestered system energy by dominating scavenging activity. To explore how the disturbance of these communities affects the stability of carrion removal as an ecosystem function, we experimentally manipulated a scavenging community within an agricultural landscape by reducing the abundance of the dominant scavenger, raccoons Procyon lotor. We then monitored the fates of 676 mouse Mus musculus carcasses placed in 13 control and 13 removal woodlots from June 2007–May 2008. The diversity of vertebrate scavengers did not change between control and removal woodlots and scavenging by invertebrates was unaffected by our experiment. Although Virginia opossums Didelphis virginiana and other scavengers exhibited a functional response when raccoons were reduced in abundance, the increases did not change the proportional allocation of carcasses among scavengers. Finally, the reduced abundance of a major scavenger affected system efficiency. More carcasses remained un‐scavenged at the end of trials in removal woodlots than in control woodlots. This experiment demonstrates the vulnerability of a critical ecosystem service, carrion removal, to perturbations of the scavenging community and serves to highlight the method by which scavenger communities may respond to perturbations.     

Interactive effects of obligate scavengers and scavenger community richness on lagomorph carcass consumption patterns

Carrion consumption patterns in vertebrate scavenger communities may be influenced by several interacting factors. We assessed the effects of the number of scavenger species and the presence of obligate scavengers (vultures) on carcass detection and consumption times, and the structure (nestedness) of the scavenger assemblage by exploring consumption patterns of lagomorph carcasses provided experimentally. Carcass detection and consumption times were strongly inversely related to vulture presence, whereas scavenger richness had a low contribution, except when interacting with vulture presence. However, none of the scavenger communities presented a nested pattern, perhaps because of the small size of lagomorphs, which prevents large numbers of scavengers and interspecific interactions occurring at one carcass. Our results suggest that scavenger species richness, especially the presence of vultures, increases scavenging efficiency.     

Carcass Type Affects Local Scavenger Guilds More than Habitat Connectivity

Scavengers and decomposers provide an important ecosystem service by removing carrion from the environment. Scavenging and decomposition are known to be temperature-dependent, but less is known about other factors that might affect carrion removal. We conducted an experiment in which we manipulated combinations of patch connectivity and carcass type, and measured responses by local scavenger guilds along with aspects of carcass depletion. We conducted twelve, 1-month trials in which five raccoon (Procyon lotor), Virginia opossum (Didelphis virginiana), and domestic rabbit (Oryctolagus spp.) carcasses (180 trials total) were monitored using remote cameras in 21 forest patches in north-central Indiana, USA. Of 143 trials with complete data, we identified fifteen species of vertebrate scavengers divided evenly among mammalian (N = 8) and avian species (N = 7). Fourteen carcasses (9.8%) were completely consumed by invertebrates, vertebrates exhibited scavenging behavior at 125 carcasses (87.4%), and four carcasses (2.8%) remained unexploited. Among vertebrates, mammals scavenged 106 carcasses, birds scavenged 88 carcasses, and mammals and birds scavenged 69 carcasses. Contrary to our expectations, carcass type affected the assemblage of local scavenger guilds more than patch connectivity. However, neither carcass type nor connectivity explained variation in temporal measures of carcass removal. Interestingly, increasing richness of local vertebrate scavenger guilds contributed moderately to rates of carrion removal (≈6% per species increase in richness). We conclude that scavenger-specific differences in carrion utilization exist among carcass types and that reliable delivery of carrion removal as an ecosystem service may depend on robust vertebrate and invertebrate communities acting synergistically.     

Effects of vulture exclusion on carrion consumption by facultative scavengers

Vultures provide an essential ecosystem service through removal of carrion, but globally, many populations are collapsing and several species are threatened with extinction. Widespread declines in vulture populations could increase the availability of carrion to other organisms, but the ways facultative scavengers might respond to this increase have not been thoroughly explored. We aimed to determine whether facultative scavengers increase carrion consumption in the absence of vulture competition and whether they are capable of functionally replacing vultures in the removal of carrion biomass from the landscape. We experimentally excluded 65 rabbit carcasses from vultures during daylight hours and placed an additional 65 carcasses that were accessible to vultures in forested habitat in South Carolina, USA during summer (June–August). We used motion‐activated cameras to compare carrion use by facultative scavenging species between the experimental and control carcasses. Scavenging by facultative scavengers did not increase in the absence of competition with vultures. We found no difference in scavenger presence between control carcasses and those from which vultures were excluded. Eighty percent of carcasses from which vultures were excluded were not scavenged by vertebrates, compared to 5% of carcasses that were accessible to vultures. At the end of the 7‐day trials, there was a 10.1‐fold increase in the number of experimental carcasses that were not fully scavenged compared to controls. Facultative scavengers did not functionally replace vultures during summer in our study. This finding may have been influenced by the time of the year in which the study took place, the duration of the trials, and the spacing of carcass sites. Our results suggest that under the warm and humid conditions of our study, facultative scavengers would not compensate for loss of vultures. Carcasses would persist longer in the environment and consumption of carrion would likely shift from vertebrates to decomposers. Such changes could have substantial implications for disease transmission, nutrient cycling, and ecosystem functioning.     

Window Area and Development Drive Spatial Variation in Bird-Window Collisions in an Urban Landscape

Collisions with windows are an important human-related threat to birds in urban landscapes. However, the proximate drivers of collisions are not well understood, and no study has examined spatial variation in mortality in an urban setting. We hypothesized that the number of fatalities at buildings varies with window area and habitat features that influence avian community structure. In 2010 we documented bird-window collisions (BWCs) and characterized avian community structure at 20 buildings in an urban landscape in northwestern Illinois, USA. For each building and season, we conducted 21 daily surveys for carcasses and nine point count surveys to estimate relative abundance, richness, and diversity. Our sampling design was informed by experimentally estimated carcass persistence times and detection probabilities. We used linear and generalized linear mixed models to evaluate how habitat features influenced community structure and how mortality was affected by window area and factors that correlated with community structure. The most-supported model was consistent for all community indices and included effects of season, development, and distance to vegetated lots. BWCs were related positively to window area and negatively to development. We documented mortalities for 16/72 (22%) species (34 total carcasses) recorded at buildings, and BWCs were greater for juveniles than adults. Based on the most-supported model of BWCs, the median number of annual predicted fatalities at study buildings was 3 (range = 0–52). These results suggest that patchily distributed environmental resources and levels of window area in buildings create spatial variation in BWCs within and among urban areas. Current mortality estimates place little emphasis on spatial variation, which precludes a fundamental understanding of the issue. To focus conservation efforts, we illustrate how knowledge of the structural and environmental factors that influence bird-window collisions can be used to predict fatalities in the broader landscape.     

Rewilding traditional grazing areas affects scavenger assemblages and carcass consumption patterns

The abandonment of traditional livestock farming systems in Mediterranean countries is triggering a large-scale habitat transformation, which, in general, consists of the replacement of open grazing areas by woodlands through non-managed regeneration. As a consequence, wild ungulates are occupying rapidly the empty niche left by domestic ungulates. Both types of ungulates represent the main trophic resource for large vertebrate scavengers. However, a comparison of how vertebrate scavengers consume ungulate carcasses in different habitats with different ungulate species composition is lacking. This knowledge is essential to forecast the possible consequences of the current farmland abandonment on scavenger species. Here, we compared the scavenging patterns of 24 wild and 24 domestic ungulate carcasses in a mountainous region of southern Spain monitored through camera trapping. Our results show that carcasses of domestic ungulates, which concentrate in large numbers in open pasturelands, were detected and consumed earlier than those of wild ungulate carcasses, which frequently occur in much lower densities at more heterogenous habitats such as shrublands and forest. Richness and abundance of scavengers were also higher at domestic ungulate carcasses in open habitats. Vultures, mainly griffons (Gyps fulvus), consumed most of the carcasses, although mammalian facultative scavengers, mainly wild boar (Sus scrofa) and red fox (Vulpes vulpes), also contributed importantly to the consumption of wild ungulate carcasses in areas with higher vegetation cover. Our findings evidence that the abandonment of traditional grazing may entail consequences for the scavenger community, which should be considered by ecologists and wildlife managers.     

The nested structure of a scavenger community

Scavenging is a widespread phenomenon in vertebrate communities which has rarely been accounted for, in spite of playing an essential role in food webs by enhancing nutrient recycling and community stability. Most studies on scavenger assemblages have often presented an oversimplified view of carrion foraging. Here, we applied for the first time the concept of nestedness to the study of a species-rich scavenger community in a forest ecosystem (Białowieża Primeval Forest, Poland) following a network approach. By analysing one of the most complete datasets existing up to now in a pristine environment, we have shown that the community of facultative scavengers is not randomly assembled but highly nested. A nested pattern means that species-poor carcasses support a subset of the scavenger assemblage occurring at progressively species-rich carcasses. This result contradicts the conventional view of facultative scavenging as random and opportunistic and supports recent findings in scavenging ecology. It also suggests that factors other than competition play a major role in determining community structure. Nested patterns in scavenger communities appear to be promoted by the high diversity in carrion resources and consumers, the differential predictability of the ungulate carcass types and stressful environmental conditions.     

Factors affecting searcher efficiency and scavenger removal of bat carcasses in Neotropical wind facilities

Bat fatalities at wind facilities have been reported worldwide, and environmental impact assessments depend on searches for carcasses around wind turbines to quantify impacts. Some of the carcasses may go undetected by search teams or be removed by scavengers during search intervals, so these biases must be evaluated and taken into account in fatality estimation. We investigated the influence of different factors on searcher efficiency and scavenger removal in a dry forest area in northeastern Brazil, one of the regions with the highest density of wind turbines in the Neotropics. We conducted searcher efficiency and scavenger removal trials around 34 wind turbines from January 2017 to January 2018. Searcher efficiency was influenced by cover type, season, and carcass size, ranging between 12% for small bats in shrub vegetation during the rainy season and 96% for large bats in absent or sparse vegetation during the dry season. Carcass type and season affected scavenger removal; carcass persistence time was shorter for chicks (1.2 days) than for bats and mice (2.1 days), and the probability of a carcass persisting for a whole day was higher in the rainy season, while the probability of carcass persistence for 7, 14, and 28 days was higher in the dry season. The scavenger community was composed of canids, birds of prey, and insects, with systematic removal of carcasses by the crab-eating fox (Cerdocyon thous) throughout the year and by dung beetles in the rainy season. Based on our findings, impact assessments of wind facilities on bats should conduct searcher efficiency trials in all seasons and cover types around wind turbines, using bat carcasses or models of different sizes. Scavenger removal trials should cover all seasons as well, and use mouse carcasses (but not chick carcasses) as surrogates for bats.     

ENDURANCE RUNNING AND ITS RELEVANCE TO SCAVENGING BY EARLY HOMININS

It has been argued that endurance running ability may have been important in hominin evolution, giving hominins an enhanced ability to scavenge by allowing them to reach carcasses before other terrestrial vertebrate scavengers. This would have allowed them to exploit the carcass before eventually surrendering it on the arrival of potentially dangerous large terrestrial scavengers. Here, we use a simple spatial model to evaluate the ability of competitors to hominin scavengers to find carcasses. We argue that both hominin and nonhominin terrestrial scavengers would often first have been alerted to available carcasses by overflying aerial scavengers. Our model estimates that nonhominin scavengers will generally be able to reach the carcass within 30 min of detecting a plume of vultures above a nearby carcass. We argue that endurance running over periods greater than 30 min would not have provided a selective advantage to early hominins through increased scavenging opportunities. However, shorter distance running may have been selected, particularly if hominins could defend or usurp carcasses from other mammalian scavengers.