Biomass Flow and Scavengers Use of Carcasses after Re-Colonization of an Apex Predator

Background

Reestablishment of apex predators influences the availability and distribution of biomass for scavengers and can therefore be an important agent for structuring species communities. We studied how the re-colonization of the Scandinavian Peninsula by wolves (Canis lupus) affected the amount and temporal variation in use of moose (Alces alces) carcasses.

Methodology/Principal Findings

We compared the availability of biomass from remains at wolf kills with those killed by hunters, vehicle collisions and natural death. Movement-triggered cameras monitored patterns of use on wolf kills and remains from hunter harvest by scavengers (n = 15 276) in relation to time of year, available carcass biomass, time since the death of the moose and presence of wolves. Remains from hunter harvest were the largest food source for scavengers both within wolf territories (57%) and in areas without wolves (81%). The total annual biomass available were similar in areas with (25 648 kg) and without (24 289 kg) wolves. Presence of wolves lowered the peak biomass available from hunter harvest in October (20%) and increased biomass available during December to August (38–324% per month). The probability of scavengers being present decreased faster with time at remains from hunter harvest compared to wolf kills and both the probability of being present and the number of visits by scavengers to wolf kills increased as the amount of biomass available on the carcass increased.

Conclusions/Significance

Wolves reduced the seasonal variation of biomass from moose carcasses and most important increased it during spring. Scavengers also visited wolf kills most frequently during spring when most scavenging species have young, which may lead to an increase in survival and/or reproductive success of scavengers within wolf territories. This applies both for abundant scavenging species that were the most frequent visitors at wolf kills and threatened scavengers with lower visit frequency.     

Spatio-temporal segregation of facultative avian scavengers at ungulate carcasses

Spatial and/or temporal segregation of resource use are mechanisms that may allow coexistence between potential competitors. Spatial and temporal patterns of carrion use were studied in the main avian scavengers of Sierra Espuña Natural Park (SE, Spain). We monitored the use of ungulate carcasses provided by hunting during winter and summer of 2005–2006. Non-breeding ravens exploited aggregated carcasses and depleted the resource in a few days while golden eagles used scattered carcasses over a longer consumption period. Moreover, non-breeding ravens exploited carcasses when golden eagles were less active. Almost all available hunter kills were exploited by golden eagles and/or ravens during winter whereas a high number of carcasses were not used during the breeding season, suggesting strong competition in a period of low food availability. Thus carrion arrangement, seasonal behaviour and competitive interactions could be affecting the foraging patterns of these two species. Interestingly, intraspecific aggregation of the smaller species and dominance of the larger one may act on the use of shared resources and trigger segregation mechanisms. Our results support that differences in the spatial and temporal patterns of resource use may allow resource partitioning between two species, thus facilitating their coexistence in sympatric areas.     

Apex predators and the facilitation of resource partitioning among mesopredators

Apex predators may influence carnivore communities through the suppression of competitively dominant mesopredators, however they also provide carrion subsidies that could influence foraging and competition among sympatric mesopredators when small prey is scarce. We assessed coyote Canis latrans and red fox Vulpes vulpes winter diet overlap and composition from scats collected in two study areas with 3‐fold difference in grey wolf Canis lupus density due to a wolf control program. We hypothesized that differences in diet composition would be driven by the use of carrion, and tested whether 1) apex predators facilitate resource overlap, or 2) apex predators facilitate resource partitioning. We estimated the available biomass of snowshoe hares and voles based on pellet density and vole capture rates in each study area. We used molecular analysis to confirm species identification of predator scats, and used microscopic evaluation of prey remains to analyze diet composition of 471 coyote and fox scats. Ungulate carrion, voles and snowshoe hares comprised 73% of coyote and fox diet, and differences in use of carrion and microtines accounted for nearly 60% of the dissimilarity in diet among these canids. Carrion was the top‐ranked item in the coyote diet in both study areas, whereas carrion use by red foxes declined 3‐fold in the study area with higher wolf and small prey abundance. Diet overlap tended to be lower and diet diversity tended to be higher where wolves were more abundant, though these trends were not statistically significant. Taken together, our findings indicate that carrion provisions could facilitate resource partitioning in mesocarnivore communities by alleviating exploitation competition for small mammals.     

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.     

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.     

Gray wolves as climate change buffers in Yellowstone

Understanding the mechanisms by which climate and predation patterns by top predators co-vary to affect community structure accrues added importance as humans exert growing influence over both climate and regional predator assemblages. In Yellowstone National Park, winter conditions and reintroduced gray wolves (Canis lupus) together determine the availability of winter carrion on which numerous scavenger species depend for survival and reproduction. As climate changes in Yellowstone, therefore, scavenger species may experience a dramatic reshuffling of food resources. As such, we analyzed 55 y of weather data from Yellowstone in order to determine trends in winter conditions. We found that winters are getting shorter, as measured by the number of days with snow on the ground, due to decreased snowfall and increased number of days with temperatures above freezing. To investigate synergistic effects of human and climatic alterations of species interactions, we used an empirically derived model to show that in the absence of wolves, early snow thaw leads to a substantial reduction in late-winter carrion, causing potential food bottlenecks for scavengers. In addition, by narrowing the window of time over which carrion is available and thereby creating a resource pulse, climate change likely favors scavengers that can quickly track food sources over great distances. Wolves, however, largely mitigate late-winter reduction in carrion due to earlier snow thaws. By buffering the effects of climate change on carrion availability, wolves allow scavengers to adapt to a changing environment over a longer time scale more commensurate with natural processes. This study illustrates the importance of restoring and maintaining intact food chains in the face of large-scale environmental perturbations such as climate change.     

The relationship between wolverine and larger predators,lynx and wolf,in a historical ecosystem context

Apex predators play an important role in shaping ecosystem structure. They may suppress smaller predators (mesopredators) but also subsidize scavengers via carrion provisioning. However, the importance of these interactions can change with ecosystem context. The wolverine (Gulo gulo) is a cold-adapted carnivore and facultative scavenger. It has a circumboreal distribution, where it could be either suppressed or subsidized by larger predators. In Scandinavia, the wolverine might interact with two larger predators, wolf (Canis lupus) and lynx (Lynx lynx), but human persecution decimated the populations in the nineteenth and early twentieth century. We investigated potential relationships between wolverine and the larger predators using hunting bag statistics from 15 Norwegian and Swedish counties in 1846–1922. Our best models showed a positive association between wolverine and lynx trends, taking ecological and human factors into account. There was also a positive association between year-to-year fluctuations in wolverine and wolf in the latter part of the study period. We suggest these associations could result from positive lynx–wolverine interactions through carrion provisioning, while wolves might both suppress wolverine and provide carrion with the net effect becoming positive when wolf density drops below a threshold. Wolverines could thus benefit from lynx presence and low-to-intermediate wolf densities.     

Predicting the influence of wolf-provided carrion on scavenger community dynamics under climate change scenarios

Climate change poses an immediate threat to the persistence and distribution of many species, yet our ability to forecast changes in species composition is hindered by poor understanding of the extent to which higher trophic‐level interactions may buffer or exacerbate the adverse effects of warming. We incorporated species‐specific consumption data from 240 wolf‐killed elk carcasses from Yellowstone National Park into stochastic simulation models to link trends in the El Niño Southern Oscillation (ENSO) to food procurement by a guild of scavengers as a function of gray wolf reintroduction. We find that a shift in ENSO towards the El Niño (warming) phase of the cycle coincident with increasing global temperatures reduces carrion for scavengers, particularly those with strong seasonal patterns in resource use such as grizzly bears. Wolves alleviate these warming‐induced food shortages by rendering control over this crucial resource to biotic rather than abiotic factors. Ecosystems with intact top predators are likely to exhibit stronger biotic regulation and should be more resistant to climate change than ecosystems lacking them.     

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.     

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.     

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.     

Spatial variability in wolf diet and prey selection in Galicia (NW Spain)

We studied wolf (Canis lupus) diet for three different landscapes in the north-western Iberian Peninsula, differing in land uses and availability of food for wolves. We examined 2740 scats, collected over a period of 4 years, in order to describe wolf diet, its geographic variation, and trophic preferences. The most consumed species were wild pony, roe deer and cattle. We observed differences in wolf diet among the three study sites, related to the availability and accessibility of food resources in each habitat. For the two study sites in northern and central Galicia, wolves showed similar diet, with high occurrence of wild pony (37 vs. 34%) and cattle (20 vs. 23%), but differing in the consumption of wild ungulates (16 vs. 8%) and carrion (7 vs. 14%). Wolf diet in eastern Galicia’s mountain ranges was entirely different, due to the higher consumption of wild ungulates (70%). Wolves showed clear prey selection patterns. Between wild ponies and livestock, wolves positively selected ponies. Among wild ungulates, wolves positively selected roe deer. Wild pony and roe deer are key species for wolf feeding in Galicia. In the Galician wild pony range, ponies are the main food for wolves. Given that the availability of wild ponies may contribute to the decrease in wolf predation on cattle, it is essential to develop innovative administrative decisions in such areas to preserve this traditional equid population. In the same way, the population of roe deer should be strengthened in the livestock areas outside the range of wild pony.     

Evidence that vulture restaurants increase the local abundance of mammalian carnivores in South Africa

Vulture restaurants are used worldwide as a conservation tool to provide threatened vultures with a source of supplementary carrion free from anthropogenic contaminants such as poisons and veterinary drugs. While the impacts of supplementary feeding sites on ecosystem and scavenging community dynamics have been investigated in Europe, no information is currently available for southern Africa. This study presents evidence that providing supplementary carrion for vultures stimulated an increase in local abundance of two species of mammalian carnivores, the brown hyaena (Hyaena brunnea) and the black‐backed jackal (Canis mesomelas). These findings require that the wider impacts of providing supplementary carrion for conserving threatened species are fully investigated.     

Donor-Control of Scavenging Food Webs at the Land-Ocean Interface

Food webs near the interface of adjacent ecosystems are potentially subsidised by the flux of organic matter across system boundaries. Such subsidies, including carrion of marine provenance, are predicted to be instrumental on open-coast sandy shores where in situ productivity is low and boundaries are long and highly permeable to imports from the sea. We tested the effect of carrion supply on the structure of consumer dynamics in a beach-dune system using broad-scale, repeated additions of carcasses at the strandline of an exposed beach in eastern Australia. Carrion inputs increased the abundance of large invertebrate scavengers (ghost crabs, Ocypode spp.), a numerical response most strongly expressed by the largest size-class in the population, and likely due to aggregative behaviour in the short term. Consumption of carrion at the beach-dune interface was rapid and efficient, driven overwhelmingly by facultative avian scavengers. This guild of vertebrate scavengers comprises several species of birds of prey (sea eagles, kites), crows and gulls, which reacted strongly to concentrations of fish carrion, creating hotspots of intense scavenging activity along the shoreline. Detection of carrion effects at several trophic levels suggests that feeding links arising from carcasses shape the architecture and dynamics of food webs at the land-ocean interface.     

Mobility of moose—comparing the effects of wolf predation risk,reproductive status,and seasonality

In a predator–prey system, prey species may adapt to the presence of predators with behavioral changes such as increased vigilance, shifting habitats, or changes in their mobility. In North America, moose (Alces alces) have shown behavioral adaptations to presence of predators, but such antipredator behavioral responses have not yet been found in Scandinavian moose in response to the recolonization of wolves (Canis lupus). We studied travel speed and direction of movement of GPS‐collared female moose (n = 26) in relation to spatiotemporal differences in wolf predation risk, reproductive status, and time of year. Travel speed was highest during the calving (May–July) and postcalving (August–October) seasons and was lower for females with calves than females without calves. Similarly, time of year and reproductive status affected the direction of movement, as more concentrated movement was observed for females with calves at heel, during the calving season. We did not find support for that wolf predation risk was an important factor affecting moose travel speed or direction of movement. Likely causal factors for the weak effect of wolf predation risk on mobility of moose include high moose‐to‐wolf ratio and intensive hunter harvest of the moose population during the past century.     

Diet shift of a facultative scavenger, the wolverine, following recolonization of wolves

1. Wolves Canis lupus L. recolonized the boreal forests in the southern part of the Scandinavian peninsula during the late 1990s, but so far there has been little attention to its effect on ecosystem functioning. Wolf predation increases the availability of carcasses of large prey, especially moose Alces alces L., which may lead in turn to a diet switch in facultative scavengers such as the wolverine Gulo gulo L. 2. Using 459 wolverine scats collected during winter-spring 2001-04 for DNA identity and dietary contents, we compared diet inside and outside wolf territories while controlling for potential confounding factors, such as prey density. We tested the hypothesis that wolverine diet shifted towards moose in the presence of wolves, while taking into account possible sexual segregation between the sexes. Occurrence of reindeer, moose and small prey was modelled against explanatory covariates using logistic mixed-effects models. Furthermore, we compared diet composition and breadth among habitats and sexes. 3. Occurrence of reindeer, moose and small prey in the diet varied with prey availability and habitat. As expected, diet contained more moose and less reindeer and small prey in the presence of wolves. Their diet in tundra consisted of 40% reindeer Rangifer tarandus L., 39% moose and 9% rodents. In forest with wolf, their diet shifted to 76% moose, 18% reindeer and 5% rodents; compared to 42% moose, 32% reindeer and 15% rodents in forest without wolf. This diet switch could not be explained by higher moose density in wolf territories. Female diet consisted of more small prey than for males, but there was a tendency for females to use the highly available moose carrion opportunistically and to hunt less on small prey within wolf territories. 4. Our study highlights how wolves increase scavenging opportunities for wolverines, and how sexual differences in diet may also apply to large scavengers. Due to their more restricted home range, female wolverines are forced to rely more on hunting small prey. The relatively high occurrence of wolf kills, however, forms an important food source to wolverines in this area. The recolonization of wolves may therefore have contributed to the consequent recolonization of wolverines into the same area.     

The scavenger spyglass: how recruiting hunters to watch carrion boosts wildlife research

Hunters support scavengers with seasonal pulses of carrion. If those hunters also deploy remote cameras at kill sites, they could simultaneously contribute data to wildlife research while gaining first-hand knowledge of scavenger ecology. In 2018–2020, we recruited hunters to monitor carcasses and offal with remote cameras across western Montana, USA. We increased our sampling effort by also setting up cameras following successful elk (Cervus canadensis) hunts at a private ranch. Cameras recorded 19 scavenger species. Golden eagles (Aquila chrysaetos) appeared at 55% of sites, and 3 individuals wore auxiliary markers, demonstrating how hunters can augment efforts to detect tagged wildlife. Cameras also documented elusive predators (e.g., wolves [Canis lupus]) and a seasonality of scavenging among American black bears (Ursus americanus). At 42% of the sites, ≥1 cervid investigated the carrion within 1 m, a behavior that may transmit the prions associated with chronic wasting disease. Hunters are willing and competent citizen scientists that can help generate wildlife observations at a broad spatial scale.     

Restoration promotes ecological functioning through greater complementarity

Restoration projects are increasingly widespread and many promote habitat succession and the diversity and abundance of faunal communities. These positive effects on biodiversity and abundance may extend to enhancing the ecological functioning and resilience of previously degraded ecosystems, but this is rarely quantified. This study surveyed a 200-ha restoring coastal wetland and three control wetlands in the Maroochy River, eastern Australia to compare the effects of wetland restoration on the consumption of carrion and the biodiversity, abundance, and functional diversity of functionally important fish and crustaceans. Carrion consumption by fish and crustaceans was measured every 6 months from spring 2017 until spring 2021 for nine events using a combination of baited cameras and scavenging assays. We found restoration improved rates of carrion consumption and the biodiversity, functional diversity, and abundance of scavenger species. Despite positive effects on the diversity of scavengers and carrion consumption, the abundance of two species, longfin eels (Anguilla reinhardtii) and mud crabs (Scylla serrata), was the most important predictors of carrion consumption rates. The spatial distribution of carrion consumption was concentrated in areas with high saltmarsh extent, moderate to high mangrove extent, and high salinity, which also resembled the distribution of both longfin eels and mud crabs. We show that restoration can promote the rates of key ecological functions but that increases to functions are likely to be characterized by low functional redundancy and greater complementarity. Therefore, maintaining or increasing the abundance of functionally important species should become a key objective in future restoration projects.     

Diet,Prey Selection,and Predation Impact of Black-Backed Jackals in South Africa

ABSTRACT To investigate the role of black-backed jackals (Canis mesomelas) as predators, we studied diet, prey selection, and predation impact of jackals on 2 game ranches in South Africa that differed in ungulate diversity and biomass. Results showed that large (>15 kg) ungulate species dominated jackal diets throughout the year on both the less diverse (range of ingested biomass across seasons = 39–78%) and more diverse (26–69%) game ranch. Other important food items included medium-sized mammals (1–3 kg; 1–26%) and fruit (2–69%), whereas small mammals comprised 3–11% of ingested biomass across seasons on both sites. Jackals were not random in consumption of ungulates, and consumption patterns suggested jackals actively hunted certain species rather than consumed them as carrion. During ungulate birthing periods, jackals consumed almost exclusively those ungulate species that were hiders (i.e., fawns were hidden in tall vegetation away from herd) regardless of ungulate densities, suggesting that primarily fawns were preyed upon. Among hiders, there was a negative relationship (P = 0.01) between body size and percent of population consumed by jackals, indicating smaller species were more susceptible than larger species to jackal predation. Consequently, springbok (Antidorcas marsupialis) were always selected over other ungulate species on both sites, and this species was the most impacted by jackal predation. In contrast, ungulate species that were followers (i.e., fawns immediately followed mothers within protection of the herd) were scarcely or not at all consumed by jackals, regardless of body size or density. Medium-sized mammals were selectively consumed over ungulates, and there was a negative relationship (P < 0.01) between consumption of berries and ungulates, indicating alternative food resources influenced consumption of ungulates on our study sites. Our results will help wildlife managers in Africa identify ungulate species susceptible to jackal predation, and can be used to develop management strategies for reducing jackal predation in areas where it is problematic.     

Behavioural responses of shallow-water benthic marine scavengers to fish carrion: a preliminary study

Observations of the behavioural responses of near-shore marine scavengers to fish carrion were made at two depths (1–2 m, 16–18 m). Gobies and juvenile whelks were the most numerous scavengers, but appeared to consume little biomass. The first scavengers to appear at carrion (seconds/minutes) were swimming forms, later (minutes) joined by fast-moving, crawling portunid crabs. Large scavengers (crabs/starfish/catsharks) arrived after tens of minutes/hours. Scavengers were ‘direct feeders’ on the bait (crabs and some fish) or ‘indirect feeders’ (gobies and whelks) on scraps generated by direct feeders. Scavengers spent little time in aggression. While fish spent relatively low proportions of their time feeding (e.g. Lipophrys pholis: 2.2–15.8%), crabs fed almost continuously (e.g. Carcinus maenas: 97.8–99.3%) before leaving baits. Crab presence depressed fish feeding. Crabs were wasteful feeders that macerated the baits, generating scraps for indirect feeders and attracting more scavengers. Large scavengers consumed most bait.