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.     

Functional compensation by insular scavengers: the relative contributions of vertebrates and invertebrates vary among islands

When the extinction of some species results in loss of ecosystem functions, other species may be able to compensate for this loss. Functional compensation has recently been observed on islands where species are likely to become extinct; however, few studies have analysed functional compensation by scavengers in insular environments. Here, we investigated the ecosystem functions of vertebrate and invertebrate scavengers on Honshu, the main island of Japan, and offshore islands (four of the Oki Islands). The Oki Islands lack several native vertebrate scavengers (raccoon dogs, boars, martens, and foxes) that are abundant on Honshu. We experimentally placed mouse carcasses on the forest floors of Honshu (12 sites) and the Oki Islands (7 sites on Dōgo, 4 sites on Nishinoshima, 4 sites on Nakanoshima, and 3 sites on Chiburijima) in September–October 2014 and 2015. Nearly all of the mouse carcasses (95–100%) placed on the forest floor disappeared within 1 week. However, the relative importance of vertebrate and invertebrate scavengers varied among islands. Vertebrate and invertebrate scavengers contributed equally on Honshu (carcass removal by vertebrates, 47%), whereas vertebrate scavengers rarely contributed to carcass removal on three of the Oki Islands (carcass removal by vertebrates: 17% on Dōgo, 30% on Nishinoshima, and 5% on Nakanoshima). Consequently, invertebrate scavengers (Nicrophorus burying beetles) functionally compensated for the low species diversity of vertebrate scavengers on the three islands (carcass burial by Nicrophorus beetles, 65–95%). However, on the small island of Chiburijima, introduced raccoon dogs contributed to nearly all of the removals (93%), suggesting that introduced scavengers can compensate for the functional reduction. This functional compensation by scavengers may help stabilise the functions and/or services of scavengers in island environments.     

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.     

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.     

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.     

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.     

Resource availability and population dynamics of Nicrophorus investigator, an obligate carrion breeder

1. Food resources for rearing young may influence insect populations. This is particularly true for insects that breed obligately on rare, ephemeral resources such as dung, fungi, or carrion. 2. Beetles in the genus Nicrophorus bury small vertebrate carcasses for rearing their young. Studies reviewed by Scott (1998) have found a positive relationship between carcass mass and total brood size. It is likely that access to carcasses suitable for breeding, and not food or mates, limits reproduction in both male and female Nicrophorus. Thus, small mammal densities could determine Nicrophorus population sizes. 3. The work reported here examined the relationship between Nicrophorus investigator (Coleoptera: Silphidae) population size and small mammal abundance at two sites over a 4‐year period. 4. Nicrophorus investigator buried and reared young on all the native small rodent species trapped at two sites in south‐western Colorado, U.S.A. (Peromyscus maniculatus, Microtus montanus, Zapus princeps, Tamias minimus, Thomomys talpoides). They preferred to bury and reproduce on rodent carcasses weighing between 16 and 48 g; rodents of this size represented 82% of captures. 5. Population sizes of N. investigator and small rodents were estimated simultaneously using mark‐recapture censuses over a 4‐year period. Considering only rodents within the size range used by N. investigator, the estimated small mammal biomass per hectare in one year and the beetle population size in the following year were correlated significantly.     

Carcass permanency time and its implications to the roadkill data

Roadkill studies are typically conducted without regard to the carcass permanency time on the highway, which may lead to underestimation of roadkill data, especially small animals that are quickly removed by scavengers. To evaluate the carcass duration time on the lane and its relation to the roadkill time event, we conducted an experiment comparing the removal rate of small carcasses between different stretches of a highway and stretches of dirt roads. The rates found in the study were considered high, with 89 % of carcasses being removed in the first 24 h and 66 % within 12 h. The removals were high for both road categories but reached their peaks at different periods of the day, being higher during the day on the highway and at night on dirt roads. We believe that removal on the highway is dictated by higher vehicular traffic and mainly by the action of scavenger birds during the day, while on dirt roads, it is due to the action of different opportunistic scavengers during the night.     

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.     

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.     

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.     

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.     

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.     

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.     

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.     

Soil and vegetation nutrient response to bison carcasses in Białowieża Primeval Forest,Poland

Ungulate carcasses can have important effects on the surrounding soil and vegetation. The impact of six carcasses of European bison (Bison bonasus) was investigated for the first time in a natural temperate forest (Białowieża, Poland) by measuring soil and plant nutrient concentrations along a gradient extending from the centre of each carcass. Calcium concentration and pH were found to be higher at the centre of the carcass, decreasing towards the periphery. This effect lasted up to 7 years after the death of the animal. The concentration of most nutrients in the soil and plants varied irregularly, suggesting an effect of the carcass at its centre but the absence of a clear pattern of variation along the gradient. Concentrations of NO₃⁻ in the soil differed only at the 1-year old carcass, suggesting a fast turnover of nitrate in temperate forests. Our results show that the effects of large herbivore carcasses on soil and plant nutrient concentrations are not easily detectable in a temperate forest as in more homogeneous habitats, such as tundra and prairie. This may be due to the high activity of scavengers and nutrient recycling in the study area, but it may also be a consequence of a more complex and patchy interaction between nutrient availability and other limiting factors in temperate forests.     

Fish Food in the Deep Sea: Revisiting the Role of Large Food-Falls

The carcasses of large pelagic vertebrates that sink to the seafloor represent a bounty of food to the deep-sea benthos, but natural food-falls have been rarely observed. Here were report on the first observations of three large ‘fish-falls’ on the deep-sea floor: a whale shark (Rhincodon typus) and three mobulid rays (genus Mobula). These observations come from industrial remotely operated vehicle video surveys of the seafloor on the Angola continental margin. The carcasses supported moderate communities of scavenging fish (up to 50 individuals per carcass), mostly from the family Zoarcidae, which appeared to be resident on or around the remains. Based on a global dataset of scavenging rates, we estimate that the elasmobranch carcasses provided food for mobile scavengers over extended time periods from weeks to months. No evidence of whale-fall type communities was observed on or around the carcasses, with the exception of putative sulphide-oxidising bacterial mats that outlined one of the mobulid carcasses. Using best estimates of carcass mass, we calculate that the carcasses reported here represent an average supply of carbon to the local seafloor of 0.4 mg m⁻²d⁻¹, equivalent to ∼4% of the normal particulate organic carbon flux. Rapid flux of high-quality labile organic carbon in fish carcasses increases the transfer efficiency of the biological pump of carbon from the surface oceans to the deep sea. We postulate that these food-falls are the result of a local concentration of large marine vertebrates, linked to the high surface primary productivity in the study area.     

Habitat patch and matrix effects on small-mammal persistence in Amazonian forest fragments

Tropical forest mammal assemblages are widely affected by the twin effects of habitat loss and habitat fragmentation. We evaluated the effects of forest patch metrics, habitat structure, age of patch isolation, and landscape metrics on the species richness, abundance and composition of small mammals at 23 forest fragments (ranging in size from 43 to 7,035 ha) in a highly deforested 3,609-km² landscape of southwestern Brazilian Amazonia. Using pitfall traps and both terrestrial and arboreal traplines of Sherman, Tomahawk and snap traps, we captured a total of 844 individuals over 34,900 trap-nights representing 26 species and 20 genera of small-mammals, including 13 rodent and 13 marsupial species. We also consider the effects of distance from forest edges on species occupancy and abundance. Overall small mammal abundance, species richness and species composition were primarily affected by the quality of the open-habitat matrix of cattle pastures, rather than by patch metrics such as fragment size. Ultimately, small mammal community structure was determined by a combination of both landscape- and patch-scale variables. Knowledge of the anthropogenic factors that govern small mammal community structure is of critical importance for managing the persistence of forest vertebrates in increasingly fragmented neotropical forest landscapes.     

The fate of cetacean carcasses in the deep sea: observations on consumption rates and succession of scavenging species in the abyssal north-east Atlantic Ocean

The fate of cetacean carcasses in the deep sea was investigated using autonomous deep-sea lander vehicles incorporating time-lapse camera systems, fish and amphipod traps. Three lander deployments placed cetacean carcasses at depths of 4000 to 4800 m in the north-east Atlantic for periods of 36 h, 152 h and 276 h before being recovered. The photographic sequences revealed that carcasses were rapidly consumed by fish and invertebrate scavengers with removal rates ranging from 0.05 to 0.4 kg h^-1. In the longest experiment the carcass was skeletonized within five days. In each deployment, approximately an hour after emplacement, the grenadier Coryphaenoides (Nematonurus) armatus and large numbers of lysianassid amphipods had arrived at the food-fall. The initially high numbers of grenadiers declined once the majority of the bait had been consumed and a variety of other fish and invertebrates were then observed, some taking up residence at the site. None of the fish species appeared to consume the carcass directly, but preyed upon amphipods instead. Funnel traps recovered with the carcass indicated a succession in the species composition of amphipods, with the specialist necrophages such as Paralicella spp. being replaced by more generalist feeders of the Orchomene species complex.