Can Wild Ungulate Carcasses Provide Enough Biomass to Maintain Avian Scavenger Populations? An Empirical Assessment Using a Bio-Inspired Computational Model

Background

The reduction in the amount of food available for European avian scavengers as a consequence of restrictive public health policies is a concern for managers and conservationists. Since 2002, the application of several sanitary regulations has limited the availability of feeding resources provided by domestic carcasses, but theoretical studies assessing whether the availability of food resources provided by wild ungulates are enough to cover energetic requirements are lacking.

Methodology/Findings

We assessed food provided by a wild ungulate population in two areas of NE Spain inhabited by three vulture species and developed a P System computational model to assess the effects of the carrion resources provided on their population dynamics. We compared the real population trend with to a hypothetical scenario in which only food provided by wild ungulates was available. Simulation testing of the model suggests that wild ungulates constitute an important food resource in the Pyrenees and the vulture population inhabiting this area could grow if only the food provided by wild ungulates would be available. On the contrary, in the Pre-Pyrenees there is insufficient food to cover the energy requirements of avian scavenger guilds, declining sharply if biomass from domestic animals would not be available.

Conclusions/Significance

Our results suggest that public health legislation can modify scavenger population trends if a large number of domestic ungulate carcasses disappear from the mountains. In this case, food provided by wild ungulates could be not enough and supplementary feeding could be necessary if other alternative food resources are not available (i.e. the reintroduction of wild ungulates), preferably in European Mediterranean scenarios sharing similar and socio-economic conditions where there are low densities of wild ungulates. Managers should anticipate the conservation actions required by assessing food availability and the possible scenarios in order to make the most suitable decisions.     

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

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

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.     

Large herbivores in sagebrush steppe ecosystems: livestock and wild ungulates influence structure and function

Improving understanding of the connections between vegetation, herbivory, and ecosystem function offers a fundamental challenge in contemporary terrestrial ecology. Using exclosures constructed during the late 1950s, we examined effects of grazing by wild and domestic herbivores on plant community structure, aboveground herbaceous primary production, and nutrient cycling at six sites in semi-arid, sagebrush rangelands during 2001-2002 in Colorado, USA. Enclosures provided three treatments: no grazing, grazing by wild ungulates only, and grazing by wild and domestic ungulates. Excluding all grazing caused an increase in shrub cover (F = 4.97, P = 0.033) and decrease in bare ground (F = 4.74, P = 0.037), but also a decrease in plant species richness (F = 6.19, P = 0.018) and plant diversity (F = 7.93, P = 0.008). Effects of wild ungulate grazing on plant cover and diversity were intermediate to the effects of combined domestic and wild grazing. Aboveground net primary production was higher in both grazed treatments than in the ungrazed one (F(wild + domestic) = 2.98, P = 0.0936 and F(wild only) = 3.55, P = 0.0684). We were unable to detect significant effects of grazing on other ecosystem states and processes including C:N ratios of standing crops, N mineralization potential, or nitrification potential. Best approximating models revealed positive correlation between N availability and herbaceous cover and a negative correlation between herbaceous primary production and the ratio of shrub-herb cover and plant diversity. We conclude that ungulate herbivory, including both wild and domestic ungulates, had significant effects on plant community structure and ecosystem function during this 42-year span. Responses to the wild ungulate treatment were consistently intermediate to responses to the no grazing and wild + domestic grazing treatments. However, we were unable to detect statistical difference between effects of wild ungulates alone and wild ungulates in combination with livestock.     

Functional differences in scavenger communities and the speed of carcass decomposition

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

The role of griffon vultures Gyps africanus and Gyps ruppellii as scavengers

In the Serengeti, predator kills formed from 8 to 45 % of the carcasses that griffon vultures fed upon, the remaining carcasses coming from animals that had died from other causes. But since vultures obtained only very small amounts of food from predator kills, they had to rely for their food supply chiefly on the carcasses that were derived from other causes. In the Serengeti the large proportion of migratory ungulates prevent the mammalian predators from building up to a population size that can be responsible for a large proportion of ungulate mortality. It is estimated that the mammalian predators only take about one quarter of the potential food supply available for carnivores. The remaining ungulates die from various other causes, and so provide food for scavengers which varies in abundance and location with the seasons. This is a food supply which mammalian scavengers cannot easily use but griffon vultures, with their adaptations for gliding flight, are able to exploit it. These vultures therefore exploit a basically different food supply from predators and probably evolved as scavengers of migratory ungulate populations.     

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.     

The feeding habits of wolves in relation to large prey availability in northern Italy

We investigated wolf feeding habits in relation to the abundance of wild and domestic ungulates to test the hypothesis that large prey are preferred and that their abundance affects the use of other food categories and diet breadth. We determined diet composition by scat analysis from December 1987 to December 1992. The research was carried out in three study areas located in northern Italy and characterised by marked differences in wild and domestic ungulate abundance. In study area A (low wild and domestic ungulate availability) fruits, livestock, other vertebrates and wild ungulates made up the bulk of the diet (71% in volume). In area B (high availability of livestock) wolf diet was mainly based on sheep and wild boars (80% in volume). In study area C (high availability of wild ungulates) wild ungulates were the main food of wolves (90% in volume). Significant differences were found among study areas in the mean percentage volume of all food categories and in particular for wild ungulates, livestock, other vertebrates and fruits (p < 0.0001 in all cases). Diet breadth decreased in areas with high availability of large wild and domestic herbivores. The use of livestock species was lower where there was high abundance, richness and diversity of the wild ungulate guild. Selection for wild ungulate species was partially affected by their abundance: however other factors as prey social behaviour, adaptability to the habitat (for introduced species), and body size could have an important role in species selection by wolves. In particular in area C wild boars were selected for, roe and red deers avoided, and fallow deers and mouflons used as available. Livestock species were used in relation to their abundance and accessibility, in particular sheep were selected for and cattle avoided; but if calves bom in the pastures were considered as the only available cattle, they were selected for and sheep were used as available. Large and in particular wild herbivores were found to be of great importance for the wolf population maintenance in northern Italy, one of the most important recovery areas of Mediterranean wolves.     

Dung Beetle Conservation: Effects of Habitat and Resource Selection (Coleoptera: Scarabaeoidea)

There is increasing concern about the conservation of dung beetles which are threatened by several peculiar dangers world-wide, both at the population and community level. A noticeable threat in Europe is the dramatic reduction in the number of cattle and sheep grazing in the open, which is associated with both intensive agriculture and the progressive reforestation of previously pastured areas. We studied dung beetle habitat and resource preferences at La Mandria Park (north-west Italy) which is a mosaic of open and wooded patches where domestic (cows and horses) and wild ungulates (deer and wild boar) co-exist. Scarabaeidae were numerically dominant, accounting for 61.5% of the approximately 3000 individuals sampled (Aphodiidae accounted for 32.5% and Geotrupidae for only 6%). However, when species richness was considered, Aphodiidae were dominant, with 17 of the 27 species found (Scarabaeidae with eight and Geotrupidae with two). Assuming a null hypothesis of equal probability of colonizing any habitat or faeces, we found that most species were significantly associated with one of the four dung types or with one of the two habitats considered. On average, Scarabaeidae preferred cattle dung and open habitats whereas most Aphodiidae used deer lumps and wooded habitats. In spite of the precise ecological choices observed at La Mandria, surveys from other European areas suggest that both habitat and food selection are quite flexible. From a conservation viewpoint, the ability of coprophagous beetles to choose herbivore faeces according to their availability and to select habitats that satisfy their own microclimate requirements may certainly be useful in preserving biodiversity. Agroecology, which implies some woodland and hedgerow management practices associated with pastoralism, could be the right approach to the management of the agricultural landscape. Conversely, in reforested areas, while wild ungulate populations should be preserved, some form of human disturbance, particularly clearings used for pastoralism, should also be maintained.     

Experimental evidence that livestock grazing intensity affects the activity of a generalist predator

Grazing by domestic ungulates has substantial impacts on ecosystem structure and composition. In grasslands of the northern hemisphere, livestock grazing limits populations of small mammals, which are a main food source for a variety of vertebrate predators. However, no experimental studies have described the impact of livestock grazing on vertebrate predators. We experimentally manipulated sheep and cattle grazing intensity in the Scottish uplands to test its impact on a relatively abundant small mammal, the field vole (Microtus agrestis), and its archetypal generalist predator, the red fox (Vulpes vulpes). We demonstrate that ungulate grazing had a strong consistent negative impact on both vole densities and indices of fox activity. Ungulate grazing did not substantially affect the relationship between fox activity and vole densities. However, the data suggested that, as grazing intensity increased i) fox activity indices tended to be higher when vole densities were low, and ii) the relationship between fox activity and vole density was weaker. All these patterns are surprising given the relative small scale of our experiment compared to large red fox territories in upland habitats of Britain, and suggest that domestic grazing intensity causes a strong response in the activity of generalist predators important for their conservation in grassland ecosystems.     

Managing open habitats by wild ungulate browsing and grazing: A case‐study in North‐Eastern Germany

Question: Can wild ungulates efficiently maintain and restore open habitats? Location: Brandenburg, NE Germany. Methods: The effect of wild ungulate grazing and browsing was studied in three successional stages: (1) Corynephorus canescens‐dominated grassland; (2) ruderal tall forb vegetation dominated by Tanacetum vulgare; and (3) Pinus sylvestris‐pioneer forest. The study was conducted over 3 yr. In each successional stage, six paired 4 m²‐monitoring plots of permanently grazed versus ungrazed plots were arranged in three random blocks. Removal of grazing was introduced de novo for the study. In each plot, percentage cover of each plant and lichen species and total cover of woody plants was recorded. Results: Wild ungulates considerably affected successional pathways and species composition in open habitats but this influence became evident in alteration of abundances of only a few species. Grazing effects differed considerably between successional stages: species richness was higher in grazed versus ungrazed ruderal and pioneer forest plots, but not in the Corynephorus sites. Herbivory affected woody plant cover only in the Pioneer forest sites. Although the study period was too short to observe drastic changes in species richness and woody plant cover, notable changes in species composition were still detected in all successional stages. Conclusion: Wild ungulate browsing is a useful tool to inhibit encroachment of woody vegetation and to conserve a species‐rich, open landscape.     

Foraging of lynxes in a managed boreal-alpine environment

Foraging of Eurasian lynxes Lynx lynx was studied with telemetry and snow tracking in central Norway. In all habitats and at all seasons, medium-sized ungulates (roe deer Capreolus capreolus , reindeer Rangifer tarandus and domestic sheep Ovis aries) dominated the diet (81% of ingested biomass estimated from faeces). Mountain hares Lepus timidus and galliform birds comprised the remainder of the diet (15% and 3%, respectively). Lynxes with different life history status did not differ in prey choice, but adult males utilised carcasses of ungulate prey considerably less (16% of the edible parts) than did females with offspring (80%) and subadults (58%.). Forest habitats in lowlands and adjacent to cultivated fields were the most favourable foraging habitats (indexed as the prey encounter rate per km lynx track) primarily owing to the presence of roe deer. Two family groups tracked in winter killed 0.2 ungulate per day. The importance of agricultural land as a foraging habitat and the dominance of livestock in the diet in remoter areas indicate that the lynx has responded to agriculture and reindeer husbandry during the past century by switching from smaff game to ungulates.     

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.     

How to fit the distribution of apex scavengers into land‐abandonment scenarios? The Cinereous vulture in the Mediterranean biome

Aim

Farmland abandonment or “ecological rewilding” shapes species distribution and ecological process ultimately affecting the biodiversity and functionality of ecosystems. Land abandonment predictions based on alternative future socioeconomic scenarios allow foretell the future of biota in Europe. From here, we predict how these forecasts may affect large‐scale distribution of the Cinereous vulture (Aegypius monachus), an apex scavenger closely linked to Mediterranean agro‐grazing systems.

Location

Iberian Peninsula.

Methods

Firstly, we modelled nest‐site and foraging habitat selection in relation to variables quantifying physiography, trophic resources and human disturbance. Secondly, we evaluate to what extent land abandonment may affect the life traits of the species and finally we determined how potential future distribution of the species would vary according to asymmetric socioeconomic land‐abandonment predictions for year 2040.

Results

Cinereous vultures selected breeding areas with steep slopes and low human presence whereas foraging areas are characterized by high abundance of European rabbits (Oryctolagus cuniculus) and wild ungulates. Liberalization of the Common Agricultural Policy (CAP) could potentially transform positively 66% of the current nesting habitat, favouring the recovery of mature forest. Contrarily, land abandonment would negatively affect the 63% of the current foraging habitat reducing the availability of preferred food resources (wild European rabbit). On the other hand, the maintenance of the CAP would determine lower frequencies (24%–22%) of nesting and foraging habitat change.

Main conclusions

Land abandonment may result into opposite effects on the focal species because of the increase in nesting habitats and wild ungulates populations and, on the other hand, lower availability of open areas with poorer densities of European rabbits. Land‐abandonment models’ scenarios are still coarse‐grained; the apparition of new human uses in natural areas may take place at small‐sized and medium‐sized scales, ultimately adding complexity to the prediction on the future of biota and ecosystems.

     

Overabundant wild ungulate populations in Europe: management with consideration of socio-ecological consequences

1. Throughout recent years, ungulates have experienced significant increases in numbers and geographic range sizes in Europe, becoming locally overabundant populations in some regions. Changes in legislation regarding poaching, abandonment of land and re-naturalisation of habitats, and decreasing numbers of hunters, among other things, have led to alarming scenarios in wild ungulate biology. Although ungulates bring some financial benefits for ecosystems and society through tourism and hunting, the problems associated with populations that are no longer controlled can outweigh the advantages. Damage to forestry and agriculture, ungulate-vehicle collisions, and diseases are among the most concerning problems related to ungulate overabundance.
2. To deal with these problems and to decide on the best management strategy to apply, it is essential to have tools available to monitor populations with an integrative approach based on ecological change indicators, and to assess population and ecosystem status. Furthermore, in a globalised world, people’s opinions matter, and sociological studies regarding human perception of wild mammals must take place in order to allow proper management, including consideration of people’s expectations as well as animal and ecosystem needs. Successful and unsuccessful management strategies have already been attempted, and the knowledge of consequences over time enables an adaptive approach.
3. Management of ungulate populations is a complex subject, and each case should be studied, analysing the cost_performance balance of measures to be taken, and ensuring ongoing financial means to carry out and continue with successful ecosystem management strategies. Multidisciplinary teams should be built, including biologists, veterinarians, stakeholders, sociologists, and others, to deal with the management of European wild ungulate populations.

     

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.     

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

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

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

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

Food searching in griffon vultures

Observations are presented on food searching in griffon vultures. To locate food directly, griffon vultures do not use a sense of smell but rely on vision. However, most birds locate carcasses indirectly—by watching the activities of neighbouring birds. The method of food searching is described and the number of birds which arrive at a carcass is shown to depend on the amount of food that is available. Birds do not hold feeding ranges, but travel widely during food searching, one bird travelling 180 km in 6 days. The density of searching birds varies greatly according to the density of ungulates in an area, birds being most numerous over high ungulate concentrations. The altitude at which birds search also varies, birds flying at higher altitudes over areas of low ungulate density and at lower altitudes over ungulate concentrations. These variations in searching density and height are shown to affect the efficiency of food searching. Carcasses are located rapidly and the food consumed quickly in high ungulate density areas, while in low ungulate density conditions carcasses can take a long time to be located. It is concluded that griffon vultures are more likely to find food and are better able to compete with mammalian competitors by searching over migratory ungulate herds than over areas containing resident ungulate species. Their adaptations for gliding flight enable these birds to follow these migratory ungulates throughout the year.     

Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white‐tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5–6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures – none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m²) and greater woody species richness at the subplot scale (1 m²) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white‐tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing‐species richness curve, but have rarely been documented in forests.