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.     

Rapid scavenging of jellyfish carcasses reveals the importance of gelatinous material to deep-sea food webs

Jellyfish blooms are common in many oceans, and anthropogenic changes appear to have increased their magnitude in some regions. Although mass falls of jellyfish carcasses have been observed recently at the deep seafloor, the dense necrophage aggregations and rapid consumption rates typical for vertebrate carrion have not been documented. This has led to a paradigm of limited energy transfer to higher trophic levels at jelly falls relative to vertebrate organic falls. We show from baited camera deployments in the Norwegian deep sea that dense aggregations of deep-sea scavengers (more than 1000 animals at peak densities) can rapidly form at jellyfish baits and consume entire jellyfish carcasses in 2.5 h. We also show that scavenging rates on jellyfish are not significantly different from fish carrion of similar mass, and reveal that scavenging communities typical for the NE Atlantic bathyal zone, including the Atlantic hagfish, galatheid crabs, decapod shrimp and lyssianasid amphipods, consume both types of carcasses. These rapid jellyfish carrion consumption rates suggest that the contribution of gelatinous material to organic fluxes may be seriously underestimated in some regions, because jelly falls may disappear much more rapidly than previously thought. Our results also demonstrate that the energy contained in gelatinous carrion can be efficiently incorporated into large numbers of deep-sea scavengers and food webs, lessening the expected impacts (e.g. smothering of the seafloor) of enhanced jellyfish production on deep-sea ecosystems and pelagic–benthic coupling.     

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.     

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.     

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.     

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.     

Moose (<Emphasis Type="Italic">Alces alces</Emphasis>) hunters subsidize the scavenger community in Alaska

In many temperate ecosystems animal carcasses resultant from wildlife harvest can provide a high-quality food source for myriad facultative scavengers. We investigated scavenger use of human-provisioned ungulate carrion from a fall moose (Alces alces) hunt during 2010 and 2011 on the Gustavus Forelands, Alaska, USA. Using data from remote cameras, we (1) identified the scavenger species that used these resources and (2) evaluated their spatial and temporal responses to this seasonal resource event by indexing their activity patterns and relative order of arrival at carrion sites. We also quantified the length of time carrion persisted and estimated the amount of moose biomass provisioned to vertebrate scavengers by human hunters. Our results indicated that 11 vertebrate species (five birds and six mammals) scavenged moose carrion. We found that the common raven was the only species documented at all carrion sites and the most abundant species at moose carrion sites. As a species group, corvids [black-billed magpie (Pica hudsonia), common raven (Corvus corax); 0.1 ± 2.3 days] were the first to arrive at human-provisioned moose carrion sites, whereas ursids [brown bear (Ursus arctos), black bear (U. americanus); 1.3 ± 1.0 days] arrived after corvids but sooner than expected and canids [gray wolf (Canis lupus), coyote (C. latrans); 3.9 ± 3.0] arrived later than expected compared to our null model. On average, carrion persisted >20 days and hunters provided scavengers with a minimum of 2720 kg (82.7 kg/km²) and 1815 kg (64.8 kg/km²) of moose carrion during 2010 and 2011, respectively. Understanding how scavengers, particularly large carnivores, interact with human-provisioned moose carrion at the rural–wildland interface is essential for mitigating potential human–wildlife conflicts associated with humans subsidizing predators with a high-quality food resource.     

Declines in scavenging by endangered vultures in the Horn of Africa

Scavenging is an important ecological process. By quickly locating and consuming carrion, vertebrate scavengers cycle nutrients, stabilize food webs, and may help mitigate disease transmission to humans. Across Africa, many scavengers feed at abattoirs (i.e. slaughterhouses), thereby aiding in waste removal. Little information exists on the scavenger community composition and dynamics at abattoirs, and, to our knowledge, the carrion removal that scavengers provide at these sites has never been quantified. We studied vertebrate scavenger ecology at 6 abattoirs in Ethiopia with time-lapse photography and in-person surveys from 2014–2019. Specifically, we investigated daily, seasonal, and inter-annual patterns in use of abattoirs by vertebrate scavengers and estimated carrion consumption rates. We demonstrated the importance of abattoirs for supporting a large number and diversity of scavenger species, including 3 critically endangered, 2 endangered, 1 vulnerable, and 2 regionally endemic bird species. At the start of the study, vultures contributed 57% of carrion removal provided by vertebrate scavengers. Detections of critically endangered Rüppell's (Gyps rueppelli) and white-backed (G. africanus) vultures declined by 73% and critically endangered hooded vultures (Necrosyrtes monachus) declined by 15% over the study period. Simultaneously, the detections of dogs more than doubled. Using estimates of species-specific carrion consumption rates from the literature, coupled with changes in scavenger detections in our study, we estimated a 12% (54 kg/day) reduction in carrion consumption, or nearly 20,000 kg carrion less consumed per year by the end of the study at these 6 abattoirs. Our results indicate that ongoing vulture declines across Africa could significantly reduce carrion removal. We recommend that improving fencing around abattoir facilities could help restrict access by feral dogs, increase foraging by vultures, and, therefore, increase overall carrion removal rates.     

Comparing scavenging in marine and terrestrial ecosystems: a case study with fish and gull carcasses in a small Mediterranean island

Carrion consumption by scavengers is a key component of both terrestrial and aquatic food webs. However, there are few direct comparisons of the structure and functioning of scavenging communities in different ecosystems. Here, we monitored the consumption of 23 fish (seabream Sparus aurata) and 34 bird (yellow-legged gull Larus michahellis) carcasses on a small Mediterranean island (Isla Grosa, southeastern Spain) and surrounding waters in summer to compare the structure of the scavenger assemblages and their carrion consumption efficiencies in terrestrial and shallow water habitats. Scavenging was highly efficient both in marine and terrestrial environments, especially in the presence of a highly abundant vertebrate scavenger species, the yellow-legged gull. The vertebrate scavenger community was richer in the marine environment, whereas the invertebrate community was richer on land. The scavenger network was usually well-structured (i.e., nested), with the exception of the community associated with fish terrestrial carcasses, which were almost monopolized by yellow-legged gulls. In contrast, gulls left conspecific carcasses untouched, thus allowing longer persistence of gull carcasses on land and their exploitation by a diverse insect community. Our study shows important differences in the scavenging process associated with environment and carcass type. Promising avenues for further eco-evolutionary and applied research arise from the comparison of scavenging processes in terrestrial and marine ecosystems, from small islands to continents.     

The mesoscavenger release hypothesis and implications for ecosystem and human well‐being

Many apex scavenger species, including nearly all obligate scavengers, are in a state of rapid decline and there is growing evidence these declines can drastically alter ecological food webs. Our understanding of how apex scavengers regulate populations of mesoscavengers, those less‐efficient scavengers occupying mid‐trophic levels, is improving; yet, there has been no comprehensive evaluation of the evidence around the competitive release of these species by the loss of apex scavengers. Here we present current evidence that supports the mesoscavenger release hypothesis, the increase in mesoscavengers and increase in carrion in the face of declining apex scavengers. We provide two models of scavenger dynamics to demonstrate that the mesoscavenger release hypothesis is consistent with ecological theory. We further examine the ecological and human well‐being implications of apex scavenger decline, including carrion removal and disease regulation services.     

Seal carrion is a predictable resource for coastal ecosystems

The timing, magnitude, and spatial distribution of resource inputs can have large effects on dependent organisms. Few studies have examined the predictability of such resources and no standard ecological measure of predictability exists. We examined the potential predictability of carrion resources provided by one of the UK's largest grey seal (Halichoerus grypus) colonies, on the Isle of May, Scotland. We used aerial (11 years) and ground surveys (3 years) to quantify the variability in time, space, quantity (kg), and quality (MJ) of seal carrion during the seal pupping season. We then compared the potential predictability of seal carrion to other periodic changes in food availability in nature. An average of 6893 kg of carrion ∙yr⁻¹ corresponding to 110.5 × 10³ MJ yr⁻¹ was released for potential scavengers as placentae and dead animals. A fifth of the total biomass from dead seals was consumed by the end of the pupping season, mostly by avian scavengers. The spatial distribution of carcasses was similar across years, and 28% of the area containing >10 carcasses ha⁻¹ was shared among all years. Relative standard errors (RSE) in space, time, quantity, and quality of carrion were all below 34%. This is similar to other allochthonous-dependent ecosystems, such as those affected by migratory salmon, and indicates high predictability of seal carrion as a resource. Our study illustrates how to quantify predictability in carrion, which is of general relevance to ecosystems that are dependent on this resource. We also highlight the importance of carrion to marine coastal ecosystems, where it sustains avian scavengers thus affecting ecosystem structure and function.     

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.     

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.     

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.     

Use of salmonid carcasses by vertebrate scavengers

The use of salmon Salmo salar carrion by otters Lutra lutra and other scavengers along the River Dee in north-east Scotland was studied by radio-tagging and individual marking of fish carcasses. More carcasses were available on the Dee than on tributary streams used for spawning, indicating that salmon returned to the river after spawning and died there. The amount of salmon carrion available to terrestrial and avian scavengers along the Dee varied from 6.7 kg. km^-1 on an upstream study area to 36 kg. km^-1 downstream. Fish carcasses in the Dee were moved by spates up to 20 km but in streams used for spawning less than 1 km. Of 86 carcasses examined in 1990/91, 64 were available to terrestrial and avian scavengers on the bank or awash and of these 45 had been fed upon by otters and 16 by birds. In 1991/92, 23 of 30 carcasses were available to terrestrial and avian scavengers. All had been fed upon, 19 by otters, four by birds. Other carcasses, in shallow water, were not available to terrestrial and avian scavengers. Subsequent scavenging was mainly by otters and continued for up to three weeks after the carcasses were found. Heron Ardea cinerea , great black-backed gull Larus marinus and crow Corvus corone also scavenged salmon carcasses along the Dee. Great black-backed gulls were the most frequent scavengers, but heron (dominant to black-backed gull) was a major scavenger in 1990/91. Crows, subordinate to other scavengers, waited, often in pairs, upon dominant scavengers. There were more scavenging birds downstream and numbers did not change between years. Of 20 salmon carcasses placed in spawning areas eight were probably, two possibly, removed by otters. Otters continued to scavenge carcasses for up to a month. Scavenging by foxes Vulpes vulpes and birds followed the removal of fish carcasses from the water by otters. Radio transmitters were removed by otters and left lying alongside carcasses.     

Carrion cycling in food webs: comparisons among terrestrial and marine ecosystems

In light of current global changes to ecosystem function (e.g. climate change, trophic downgrading, and invasive species), there has been a recent surge of interest in exploring differences in nutrient cycling among ecosystem types. In particular, a growing awareness has emerged concerning the importance of scavenging in food web dynamics, although no studies have focused specifically on exploring differences in carrion consumption between aquatic and terrestrial ecosystems. In this forum we synthesize the scavenging literature to elucidate differences in scavenging dynamics between terrestrial and marine ecosystems, and identify areas where future research is needed to more clearly understand the role of carrion consumption in maintaining ecosystem function within each of these environments. Although scavenging plays a similar functional role in terrestrial and aquatic food webs, here we suggest that several fundamental differences exist in scavenging dynamics among these ecosystem types due to the unique selection pressures imposed by the physical properties of water and air. In particular, the movement of carcasses in marine ecosystems (e.g. wave action, upwelling, and sinking) diffuses biological activity associated with scavenging and decomposition across large, three‐dimensional spatial scales, creating a unique spatial disconnect between the processes of production, scavenging, and decomposition, which in contrast are tightly linked in terrestrial ecosystems. Moreover, the limited role of bacteria and temporal stability of environmental conditions on the sea floor appears to have facilitated the evolution of a much more diverse community of macrofauna that relies on carrion for a higher portion of its nutrient consumption than is present in terrestrial ecosystems. Our observations are further discussed as they pertain to the potential impacts of climate change and trophic downgrading (i.e. removal of apex consumers from ecosystems) on scavenging dynamics within marine and terrestrial ecosystems.     

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.     

Effects of carrion resources on herbivore spatial distribution are mediated by facultative scavengers

Carcasses of large herbivores are pulsed resources whose impact on animal communities and ecological processes is poorly understood. In temperate forests, long-lasting ungulate carcasses are a prime resource for many species of birds and mammals during winter. Facultative carrion-eaters also consume live prey, thus potentially leading to unexpected secondary effects on populations of species not directly linked to carcass exploitation. By snow-tracking and direct observations we investigated in Bia?owie?a Forest (E. Poland) whether large ungulate carcasses elicit spatial responses in facultative scavengers and their prey. We found that in the vicinity of carcass sites the probability of the presence of common ravens Corvus corax, jays Garrulus glandarius and red foxes Vulpes vulpes increased significantly. Indeed, large groups of the two bird species were exclusively found in those places. Because of these aggregations, the probability of predator–prey encounters (red foxes and brown hares Lepus europaeus) was significantly higher near carcass sites. Accordingly, the abundance of hares and other live prey such as red squirrels Sciurus vulgaris decreased at their vicinities, probably as a consequence of direct killing and/or predator avoidance. This study provides the first evidence of carrion pulses permeating into apparently distant trophic levels, such as herbivores, via facultative scavengers, thus highlighting some unnoticed but relevant effects of carrion resources on community structure.     

Factors affecting the fate of Pacific lamprey carcasses and resource transport to riparian and stream macrohabitats

1. Anadromous fish transport marine-derived nutrients to freshwaters during spawning migrations with potential implications for stream food webs. While many studies have explored the role of marine-derived nutrients instream ecosystems (particularly via Pacific salmonids [Oncorhynchus spp.]), relatively few have examined the spatial distribution and patchiness of non-salmonid fish carcasses or rates of transport to the riparian zone.
2. We radio-tagged and released 144 mature Pacific lamprey (Entosphenus tridentatus) prior to spawning and tracked the fate of post-spawn carcasses in two inland Columbia River basin streams to characterise spatial distribution of carcasses and marine-derived nutrient deposition. We found that 27 and 40% of lamprey that could be assigned a fate were moved into the riparian zone adjacent to stream segments exhibiting higher velocity conditions with larger substrates. Conversely, lamprey with instream fates were associated with depositional microhabitats and woody debris dams. Estimated carcass loading rates varied by more than an order of magnitude among habitats. These patterns probably reflect a combination of processes influencing the likelihood of carcass removal (e.g. by predators or scavengers, or stranding) and factors affecting the distribution of carcasses remaining within the stream.
3. Our results demonstrate substantial transport of lamprey carcasses across the stream-riparian ecotone and a non-random distribution of carcasses within streams, patterns which probably influence how resources enter stream and riparian food webs. More broadly, the results suggest local and landscape-scale hydrogeomorphic factors, along with species-specific traits and phenology, affect the distribution and potential roles of fish carrion in stream food webs.

     

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.