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.     

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.     

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.     

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.     

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.     

Temporal resource partitioning mediates vertebrate coexistence at carcasses: The role of competitive and facilitative interactions

Unravelling how biodiversity is maintained despite species competition for shared resources has been a central question in community ecology, and is gaining relevance amidst the current biodiversity crisis. Yet, we have still a poor understanding of the mechanisms that regulate species coexistence and shape the structure of assemblages in highly competitive environments such as carrion pulsed resources. Here, we study how large vertebrates coexist in scavenger assemblages by adapting their diel activity at large ungulate carcasses in NW Spain. We used camera traps to record vertebrate scavengers consuming 34 carcasses of livestock and hunted wild ungulates, which allowed us to assess also differences regarding carcass origin. To evaluate temporal resource partition among species, we estimated the overlap of diel activity patterns and the mean times of each scavenger at carcasses. We recorded 16 species of scavengers, 7 mammals and 9 birds, and found similar richness at both types of carcasses. Birds and mammals showed contrasting diel activity patterns, with birds using carcasses during daytime (mean= 11:38 h) and mammals mostly at night (23:09 h). The unimodal activity patterns of scavengers showed asynchronous peaks among species. Subordinate species modified their activity patterns at carcasses used by apex species to reduce temporal overlap. Also, diel activity patterns of vultures closely followed those of corvids, suggesting facilitation processes in which corvids would enhance carcass detection by vultures. Two mammal species (12.5%) increased nocturnality at carcasses of hunted ungulates, which could be a response to human disturbance. Our results suggest that both temporal segregation and coupling mediate the coexistence of large vertebrates at carcasses. These mechanisms might lead to richer scavenger assemblages and thereby more efficient ones in driving critical ecosystem functions related to carrion consumption, such as energy and nutrient recycling and biodiversity maintenance.     

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.     

Species and individual replacements contribute more than nestedness to shape vertebrate scavenger metacommunities

Understanding the mechanisms that organize biodiversity is central in ecology and conservation. Beta diversity links local (alfa) and regional (gamma) diversity, giving insight into how communities organize spatially. Metacommunity ecology provides the framework to interpret regional and local processes interacting to shape communities. However, the lack of metacommunity studies for large vertebrates may limit the understanding and compromise the preservation of ecosystem functions and services. We aim to understand the mechanisms underlying differences in species composition among vertebrate scavenger communities ? which provide key ecosystem functions, e.g. carrion consumption ? within a metacommunity context. We obtained species richness and abundances at scavenger communities consuming ungulate carcasses monitored through motion‐triggered remote cameras in seven terrestrial ecosystems in Spain. We partitioned beta diversity to decompose incidence‐based (species presence/absence) and abundance‐based dissimilarities into their components (turnover/balanced variation and nestedness/abundance gradient, respectively). We identified the environmental factors explaining the observed patterns. The vertebrate scavenger metacommunity consisted of 3101 individuals from 30 species. Changes in composition among ecosystems were mostly (> 84%) due to species or individual replacement (i.e. turnover or balanced variation). Species or individual loss/gain (i.e. nestedness or abundance gradient) accounted for 13–16% of these changes. Mean carcass weight, elevation and habitat diversity were the main factors explaining species/individual replacement. Our findings suggest that local processes such as species‐sorting through habitat heterogeneity would dominate scavenger metacommunity dynamics together with stochastic forces (i.e. related to carrion unpredictability and scavenging being a widespread strategy among vertebrates). The presence of structured patterns (i.e. nestedness) in beta diversity could reflect a role of deterministic processes: mass‐effects through dispersal and defaunation. Vultures are long‐distance foragers and functionally dominant species, which would connect local assemblages within the metacommunity, supporting scavenger diversity and functions across space. These results highlight the importance of managing vertebrate scavenger assemblages within a metacommunity context.     

Scavenging affects persistence of avian carcasses resulting from window collisions in an urban landscape

ABSTRACT Collisions with windows remain an important human‐related threat to bird survival in urban landscapes. Accurately estimating the magnitude of avian mortality at windows is difficult and may be influenced by many sources of error, such as scavenging of carcasses. Failure to account for removal of carcasses by scavengers can bias estimates of window mortality. We tested the hypothesis that carcass survival depends on local habitat factors known to influence scavenger behavior. Scavenger activity on bird carcasses was documented at 20 buildings in an urban landscape in northwestern Illinois for 1 week during each season of a year. Known‐fate models were used to relate carcass survival to local habitat composition and to evaluate temporal variation in survival. We also documented species of scavengers and the timing of scavenging using motion‐triggered cameras. Daily carcass survival was greater in winter than during spring, summer, and fall. Survival was related negatively to canopy cover (trees and shrubs within a 50‐m buffer) and window area, and positively to pavement cover. Using an exponential model of survival time, estimated mean time of survival of carcasses (t± SE) was 82.9 ± 11.7 d for winter and 11.8 ± 7.2 d for other seasons. Raccoons (Procyon lotor) scavenged more carcasses than other species. Our results suggest that (1) carcass survival times may be short at locations with preferred habitats of known scavengers and predictable sources of food, and (2) knowledge of scavenger distribution and activity can inform predictive models of persistence. In studies of bird‐window collisions, the influence of scavenger bias can be minimized by maintaining short time intervals between carcass searches. Search intervals can be inferred by estimating the number of days that a carcass should persist at a site, which can be calculated using predicted daily survival probabilities of carcasses at study buildings.     

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.     

Scavenger guild and consumption patterns of an invasive alien fish species in a Mediterranean wetland

Invasive Alien Species (IAS) alter ecosystems, disrupting ecological processes and driving the loss of ecosystem services. The common carp Cyprinus carpio is a hazardous and widespread IAS, becoming the most abundant species in many aquatic ecosystems. This species transforms ecosystems by accumulating biomass to the detriment of other species, thus altering food webs. However, some terrestrial species, such as vertebrate scavengers, may benefit from dead carps, by incorporating part of the carp biomass into the terrestrial environment. This study describes the terrestrial vertebrate scavenger assemblage that benefits from carp carcasses in a Mediterranean wetland. We also evaluate the seasonal differences in the scavenger assemblage composition and carrion consumption patterns. Eighty carp carcasses (20 per season) were placed in El Hondo Natural Park, a seminatural mesohaline wetland in south‐eastern Spain, and we monitored their consumption using camera traps. We recorded 14 scavenger species (10 birds and four mammals) consuming carp carcasses, including globally threatened species. Vertebrates consumed 73% of the carrion biomass and appeared consuming at 82% of the carcasses. Of these carcasses consumed, 75% were completely consumed and the mean consumption time of carcasses completely consumed by vertebrates was 44.4 h (SD = 42.1 h). We recorded differences in species richness, abundance, and assemblage composition among seasons, but we did not find seasonal differences in consumption patterns throughout the year. Our study recorded a rich and efficient terrestrial vertebrate scavenger assemblage benefitting from carp carcasses. We detected a seasonal replacement on the scavenger species, but a maintenance of the ecological function of carrion removal, as the most efficient carrion consumers were present throughout the year. The results highlight the importance of vertebrate scavengers in wetlands, removing possible infectious focus, and moving nutrients between aquatic and terrestrial environments.     

Landscape heterogeneity metrics as indicators of bird diversity: Determining the optimal spatial scales in different landscapes

Species distribution models are often used to study the biodiversity of ecosystems. The modelling process uses a number of parameters to predict others, such as the occurrence of determinate species, population size, habitat suitability or biodiversity. It is well known that the heterogeneity of landscapes can lead to changes in species’ abundance and biodiversity. However, landscape metrics depend on maps and spatial scales when it comes to undertaking a GIS analysis.We explored the goodness of fit of several models using the metrics of landscape heterogeneity and altitude as predictors of bird diversity in different landscapes and spatial scales. Two variables were used to describe biodiversity: bird richness and trophic level diversity, both of which were obtained from a breeding bird survey by means of point counts. The relationships between biodiversity and landscape metrics were compared using multiple linear regressions. All of the analyses were repeated for 14 different spatial scales and for cultivated, forest and grassland environments to determine the optimal spatial scale for each landscape typology.Our results revealed that the relationships between species’ richness and landscape heterogeneity using 1:10,000 land cover maps were strongest when working on a spatial scale up to a radius of 125–250 m around the sampled point (circa 4.9–19.6 ha). Furthermore, the correlation between measures of landscape heterogeneity and bird diversity was greater in grasslands than in cultivated or forested areas. The multi-spatial scale approach is useful for (a) assessing the accuracy of surrogates of bird diversity in different landscapes and (b) optimizing spatial model procedures for biodiversity mapping, mainly over extensive areas.     

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.     

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.     

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.     

The underestimated role of carrion in vertebrates' diet studies

Aim

Despite the increasing scientific evidence on the importance of carrion in the ecology and evolution of many vertebrates, scavenging is still barely considered in diet studies. Here, we draw attention to how scientific literature has underestimated the role of vertebrates as scavengers, identifying the ecological traits that characterize those species whose role as scavengers could have gone especially unnoticed.

Location

Global.

Time Period

1938–2022.

Major Taxa Studied

Terrestrial vertebrate scavengers.

Methods

We analysed and compared (a) the largest database available on scavenging patterns by carrion-consuming vertebrates, (b) 908 diet studies about 156 scavenger species and (c) one of the most complete databases on bird and mammal diets (Elton Traits database). For each of these 156 species, we calculated their scavenging degree (i.e. proportion of carcases where the species is detected consuming carrion) as a proxy for carrion consumption, and related their ecological traits with the probability of being identified as scavengers in diet studies and in the Elton Traits database.

Results

More than half of the species identified as scavengers at monitored carcasses were not assigned carrion as food source in their diet studies nor in the Elton Traits database. Using a subset of study sites, we found a direct relationship between a species' scavenging degree and its rate of carrion biomass removal. In addition, scavenger species, which were classified as non-predators and mammals had a lower probability of being identified as scavengers in diet studies and in the Elton Traits database, respectively.

Main Conclusions

Our results clearly indicate an underestimation of the role of scavenging in vertebrate food webs. Given that detritus recycling is fundamental to ecosystem functioning, we encourage further recognition and investigation of the role of carrion as a food resource for vertebrates, especially for non-predator species and mammals with higher scavenging degree.     

Trophic Facilitation or Limitation? Comparative Effects of Pumas and Black Bears on the Scavenger Community

Scavenging is a widespread behaviour and an important process influencing food webs and ecological communities. Large carnivores facilitate the movement of energy across trophic levels through the scavenging and decomposition of their killed prey, but competition with large carnivores is also likely to constrain acquisition of carrion by scavengers. We used an experimental approach based on motion-triggered video cameras at black-tailed deer (Odocoileus hemionus columbianus) carcasses to measure the comparative influences of two large carnivores in the facilitation and limitation of carrion acquisition by scavengers. We found that pumas (Puma concolor) and black bears (Ursus americanus) had different effects on their ecological communities. Pumas, as a top-level predator, facilitated the consumption of carrion by scavengers, despite significantly reducing their observed sum feeding times (165.7 min±21.2 SE at puma kills 264.3 min±30.1 SE at control carcasses). In contrast, black bears, as the dominant scavenger in the system, limited consumption of carrion by scavengers as evidenced by the observed reduction of scavenger species richness recorded at carcasses where they were present (mean = 2.33±0.28 SE), compared to where they were absent (mean = 3.28±0.23 SE). Black bears also had large negative effects on scavenger sum feeding times (88.5 min±19.8 SE at carcasses where bears were present, 372.3 min±50.0 SE at carcasses where bears were absent). In addition, we found that pumas and black bears both increased the nestedness (a higher level of order among species present) of the scavenger community. Our results suggest that scavengers have species-specific adaptions to exploit carrion despite large carnivores, and that large carnivores influence the structure and composition of scavenger communities. The interactions between large carnivores and scavengers should be considered in future studies of food webs and 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.     

Scavenging by vertebrates: behavioral, ecological, and evolutionary perspectives on an important energy transfer pathway in terrestrial ecosystems

Carrion use by terrestrial vertebrates is much more prevalent than conventional theory implies, and, rather than a curiosity of animal behavior, is a key ecological process that must be accounted for. Human aversion to rotted substances and difficulties associated with identifying scavenged material in studies of food habits have contributed to the relative lack of information concerning scavenging behavior in vertebrates. Several lines of evidence, however, suggest that carrion resources are more extensively used by vertebrates than has been widely assumed: 1) a substantial number of animals die from causes other than predation and become available to scavengers, 2) a wide variety of vertebrate scavengers, rather than microbes or arthropods, consume most available carcasses, and 3) intense competition exists between vertebrate scavengers and decomposers, especially in warm climates. Although vultures are best adapted to use carrion, nearly all vertebrate predators are also scavengers to some extent. The costs and benefits associated with carrion use influences the evolution of scavenging behavior in vertebrates, resulting in a continuum of facultative scavengers that use carrion to varying degrees. The realized usage of carrion by a vertebrate species is influenced by the speed and efficiency with which it forages, its visual and olfactory abilities, and its capacity for detoxifying products of decomposition. A deeper understanding of carrion use by facultative scavengers will improve our knowledge of community and ecosystem processes, especially the flow of energy through food webs.     

Towards indicators of butterfly biodiversity based on a multiscale landscape description

The conservation of biodiversity has gained prominence in ecological research for the last decades. Conservation actions require a measure of biodiversity such as species richness, but its assessment is very difficult, even for small areas and therefore the search for surrogates (i.e. indicators) of biodiversity has emerged as an active research topic. We investigated the relationships between butterfly species richness and landscape structure and composition in two pine plantation sites in Southwest France. We assessed the correlation between butterfly species richness and a set of 15 landscape metrics computed for 18 land-uses at 10 different spatial scales. Spatial scales were accounted for by computing landscape metrics for circular buffers with radius ranging from 100 to 1000 m. The joint use of the Partial Least Squares Regression (PLSR) and a stepwise regression procedure revealed strong correlations between butterfly species richness and various landscape metrics in both study sites. The selected landscape metrics differed from one site to another and mostly involved measures of landscape fragmentation. We found a very strong effect of the spatial scale of investigation upon the perception of the landscape–butterfly richness relationship. Our main conclusions are that (i) certain landscape attributes can potentially serve as indicators for butterfly species richness at the landscape scale; (ii) future indicators of biodiversity based on landscape features should consider various spatial scales.