The potential for large carnivores to act as biodiversity surrogates in southern Africa

Biodiversity in southern Africa is globally extraordinary but threatened by human activities. Although there are considerable biodiversity conservation initiatives within the region, no one has yet assessed the potential use of large carnivores in such actions. Surrogate approaches have often been suggested as one such way of capitalizing on large carnivores. Here we review the suitability of the large carnivore guild (i.e., brown hyaena Hyaena hyaena, spotted hyaena Crocuta crocutta, cheetah Acinonyx jubatus, leopard Panthera pardus, lion Panthea leo and African wild dog Lycaon pictus) to act as surrogate species for biodiversity conservation in southern Africa. We suggest that the guild must be complete for the large carnivores to fully provide their role as ecological keystones. The potential for large carnivores to act as umbrella and indicator species seems limited. However, self-sustaining populations of large carnivores may be useful indicators of unfragmented landscapes. Moreover, diversity within the large carnivore guild may reflect overall biodiversity. Although the global appeal of the large African carnivores makes them important international flagships, we stress that international conservation funding must be linked to local communities for them to be important also locally. In summary, we suggest that the flagship value of these large carnivores should be used to promote biodiversity conservation in the region, and that the suggested relationship between large carnivore diversity and overall biodiversity is empirically tested. Finally we suggest that direct conservation activities should focus on enhancing the keystone values of large carnivores through complete guild conservation and restoration.     

Seasonal variation in prey preference,diet partitioning and niche breadth in a rich large carnivore guild

Large carnivore community structure is affected by direct and indirect interactions between intra-guild members. Co-existence between different species within a carnivore guild may occur through diet, habitat or temporal partitioning. Since carnivore species are highly dependent on availability and accessibility of prey, diet partitioning is potentially one of the most important mechanisms in allowing carnivores to co-exist. Intra-guild interactions may vary over time as carnivore prey preference and diet overlap can change due to seasonal changes in resource availability. We conducted scat analysis to compare the seasonal changes in prey preference, diet partitioning and niche breadth of four large carnivore species, namely leopard Panthera pardus, spotted hyena Crocuta crocuta, brown hyena Parahyaena brunnea and wild dog Lycaon pictus in central Tuli, Botswana. Large carnivores in central Tuli display a high dietary overlap, with spotted hyena and brown hyena displaying almost complete dietary overlap and the other carnivore species displaying slightly lower but still significant dietary overlap. Dietary niche breadth for both hyena species was high possibly due to their flexible foraging strategies, including scavenging, while leopard and wild dog showed a relatively low niche breadth, suggesting a more specialised diet. High dietary overlap in central Tuli is possibly explained by the high abundance of prey species in the area thereby reducing competition pressure between carnivore species. Our research highlights the need to assess the influence of diet partitioning in structuring large carnivore communities across multiple study sites, by demonstrating that in prey rich environments, the need for diet partitioning by carnivores to avoid competition may be limited.     

Dogs Canis familiaris as carnivores: their role and function in intraguild competition

• 1 Dogs Canis familiaris are the world's most common carnivore and are known to interact with wildlife as predators, prey, competitors, and disease reservoirs or vectors.
• 2 Despite these varied roles in the community, the interaction of dogs with sympatric wild carnivore species is poorly understood. We review how dogs have been classified in the literature, and illustrate how the location and ranging behaviour of dogs are important factors in predicting their interactions with wild prey and carnivores.
• 3 We detail evidence of dogs as intraguild competitors with sympatric carnivores in the context of exploitative, interference and apparent competition.
• 4 Dogs can have localized impacts on prey populations, but in general they are not exploitative competitors with carnivores. Rather, most dog populations are highly dependent on human‐derived food and gain a relatively small proportion of their diet from wild prey. However, because of human‐derived food subsidies, dogs can occur at high population densities and thus could potentially outcompete native carnivores, especially when prey is limited.
• 5 Dogs can be effective interference competitors, especially with medium‐sized and small carnivores. Dogs may fill the role of an interactive medium‐sized canid within the carnivore community, especially in areas where the native large carnivore community is depauperate.
• 6 Dogs can also be reservoirs of pathogens, because most populations around the world are free‐ranging and unvaccinated. Diseases such as rabies and canine distemper have resulted in severe population declines in several endangered carnivores coexisting with high‐density dog populations. Dogs can therefore be viewed as pathogen‐mediated apparent competitors, capable of facilitating large‐scale population declines in carnivores.
• 7 Based on this information, we propose conceptual models that use dog population size and ranging patterns to predict the potential for dogs to be intraguild competitors. We discuss how interactions between dogs and carnivores might influence native carnivore communities.

     

Forest-edge utilization by carnivores in relation to local and landscape habitat characteristics in central European farmland

Rapid changes in agricultural landscape structure and composition affect many different farmland biotas, including carnivores, which are a key element of ecosystem stability, yet little is known about their distribution and habitat use. In this study, we evaluated how habitat characteristics on two different spatial scales (local and landscape scale) affected the forest-edge utilization by small and medium-sized carnivores in fragmented central European farmland. Based on an indirect method for detecting carnivores (scent stations), we sampled 212 forest fragments of different sizes (1–7864 ha) during April to May from 2006 to 2009. Our results indicate that carnivore utilization of forest-edge habitats was driven by landscape rather than local characteristics even though the overall extent of explained variation was small. The most important factors that determined response of the carnivore community were the area of farmland and that of urban land on a landscape scale. The corridor connectivity between small forest fragments and other spatial elements played a crucial role in the occurrence of red fox. Our results suggest that comprehensive studies on multi-species carnivore assemblage using scent station might be useful in evaluating species-specific response to habitat characteristics, especially if large numbers of stations visited by carnivores are available.     

Differences in diet and trophic interactions of Patagonian carnivores between areas with mostly native or exotic prey

The prey base for Patagonian carnivores has been altered greatly over the past 150 years due to widespread overgrazing by livestock, invasions by exotic wildlife, and hunting. On ranches in northern Patagonia carnivores consume mostly exotic species, and native herbivores are ecologically extinct in their role as prey. In this study we compare diets of the culpeo (Lycalopex culpaeus), chilla (L. griseus), puma (Puma concolor), Geoffroy's cat (Leopardus geoffroyi), colocolo (L. colocolo), and hog-nosed skunks (Conepatus chinga and C. humboldtii) in a reserve with a mostly native prey base to their diets on the ranches, and evaluate how differences in prey bases affect trophic interactions among carnivores. Carnivores in the reserve consumed mostly native prey. Dietary overlap among carnivores was not significant on the reserve, but was highly significant on the ranches. This homogenization of diets where densities of native species are reduced could lead to stronger negative interactions among carnivores, altering the composition of the carnivore assemblage to the detriment of the more specialized species. Study of carnivore diets may be a relatively quick way to evaluate the conservation status and ecological functionality of prey assemblages in Patagonia and other areas where these have been altered.     

Key factors and related principles in the conservation of large African carnivores

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The adaptive significance of an environmentally-cued developmental switch in an anuran tadpole

Summary This study investigated the proximate basis of bimodally-distributed, environmentally-induced variation that occurs in natural populations of spade-foot toad tadpoles (Scaphiopus multiplicatus). Most individuals in most populations occur as a small, slowly-developing omnivore morph. In some of these same populations, a varying number of individuals occur as a large, rapidly-developing carnivore morph (Pfennig 1989). Censuses of 37 different natural ponds revealed that the frequency of the faster-developing carnivore morph correlated significantly positively with fairy shrimp density (their chief prey) and pond drying rate. By simultaneously varying two diet components and pond drying regime in artificial pools I found that only fairy shrimp density significantly affected the proportion of carnivores. Separate experiments established that the extent to which tadpoles developed the carnivore morphology correlated with shrimp density, and that morph determination depended on the ingestion of shrimp, not simply their presence. If a critical number of shrimp were ingested, the tadpole developed into a carnivore; if not, the tadpole developed by default into an omnivore. Thus a single cue — shrimp ingestion — triggers alternative ontogenetic trajectories. Using shrimp density to induce morph differentiation enables tadpoles to respond to their environment adaptively as shrimp are most abundant in highly ephemeral ponds, where the faster developing carnivores are favored.     

Changes in home range sizes and population densities of carnivore species along the natural to urban habitat gradient

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Impact of Livestock Husbandry on Small- and Medium-Sized Carnivores in Kalahari Savannah Rangelands

ABSTRACT We analyzed relative sensitivities of small- and medium-sized carnivores to livestock husbandry (stocking rates and predator control) in Kalahari, South Africa, rangelands at a regional scale. We monitored small carnivores using track counts on 22 Kalahari farms across a land-use gradient ranging from low to high stocking rates and also interviewed each farm manager to identify farmers' perception of small carnivores as potential predators for livestock. We recorded 12 species of small- and medium-sized carnivores across 22 Kalahari farms. Stocking rate was the most important driving variable for local carnivore abundance. Abundance of all species was lowest on farms where stocking rate was high. Most farm managers perceived medium-sized carnivores, in particular, African wildcat (Felis silvestris lybica), black-backed jackal (Canis mesomelas), and caracal (Caracal caracal), as potential predators of livestock. Multiple regression analysis shows that black-backed jackal, African wildcat, and caracal were negatively affected by predator control measures, whereas bat-eared fox (Otocyon megalotis), cape fox (Vulpes chama), and small-spotted genet (Genetta genetta) were positively affected. Our results show a need for expanding research and conservation activities toward small- and medium-sized carnivores in southern African savannah rangelands. We, therefore, suggest developing a monitoring program combining passive tracking with indigenous knowledge of local Khoisan Bushmen to monitor carnivore populations, and we recommend additional predator removal experiments that manipulate predator densities.     

An ecological reassessment of the southern African carnivore guild: a case study from Member 4, Sterkfontein, South Africa

The southern African late Pliocene to early Pleistocene carnivore guild was much larger than that of the present day. Understanding how this guild may have functioned is important for the reconstruction of carnivore-hominin interactions and to assess the potential for hominin scavenging in southern Africa. In modern ecosystems, the coexistence of larger carnivore species is constrained by several factors, which include high levels of interspecific competition. Here, the composition of the fossil carnivore guild is examined using Sterkfontein Member 4 (Cradle of Humankind, South Africa) as a case study. Sterkfontein Member 4 contains 10 larger carnivore taxa (body mass > 21.5 kg) and may also contain two Australopithecus species. Two possible causes of higher numbers of carnivore species in the South African fossil record are initially considered. First, that there is a bias introduced through comparing assemblages of differing sizes; second, carnivore biodiversity may have been artificially inflated due to previous taxonomic splitting of carnivore species, such as Crocuta. These possibilities are rejected and modern ecological data are used to construct a simple spatial model to determine how many carnivores could have co-existed. Although the resulting model indicates that the carnivore taxa present in Member 4 could have co-occurred, modern ecological studies indicate that it is highly unlikely that they would have co-existed simultaneously. Considering the complex depositional processes that operate in the southern African cave sites, it is proposed that the larger carnivore guild observed in the Sterkfontein Member 4 fossil assemblage is a palimpsest created by time-averaging. In light of this, we suggest that sites which have a large number of carnivore taxa should be examined for time-averaging, while those sites which have relatively few species may be a better reflection of carnivore communities.     

Use of agroecosystem matrix habitats by mammalian carnivores (Carnivora): a global‐scale analysis

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Eavesdropping in solitary large carnivores: Black bears advance and vocalize toward cougar playbacks

Large carnivore behavioral responses to the cues of their competitors are rarely observed, but may mediate competition between these top predators. Playback experiments, currently limited to interactions involving group‐living large carnivores, demonstrate that attending to cues indicative of the immediate presence of heterospecific competitors plays a substantial role in influencing competition among these species. Group‐living species vocalize regularly to signal to one another, and competitors can readily “eavesdrop” on these acoustic cues. Solitary large carnivores also vocalize to conspecifics, but much less frequently, reducing the ease with which heterospecific competitors can eavesdrop. Eavesdropping could nonetheless play a substantive role in mediating competition among solitary large carnivores if the benefits of responding to the acoustic cues of heterospecific competitors (reducing risk or locating resources) are sufficiently large. Behavioral interactions between solitary large carnivore species are almost never observed, and there have been no experimental tests of their reactions to cues indicative of the immediate presence of other solitary large carnivores. We used an automated playback system to test the responses of a solitary large carnivore (black bear, Ursus americanus) to vocalizations of their similarly solitary competitor (cougar, Puma concolor), presenting both cougar and control vocalizations to free‐living bears foraging along shorelines in British Columbia, Canada. Both mothers with cubs and solitary bears were significantly more likely to advance and vocalize toward cougar than control playbacks, mothers producing one or both of two distinct vocalizations and solitary bears producing just one. Cougars could either represent a potential risk to bears (particularly cubs), or a source of resources, as bears are known to regularly scavenge cougar kills. Our results are consistent with bears eavesdropping on cougars for both these reasons. As with group‐living species, eavesdropping may be common among solitary large carnivores, and may be an important driver of competition between these species.     

Seasonal occupancy of sympatric larger carnivores in the southern San Andres Mountains,south-central New Mexico,USA

We investigated factors facilitating coexistence of pumas (Puma concolor), coyotes (Canis latrans), and bobcats (Lynx rufus) in the arid San Andres Mountains of south-central New Mexico, during the season (winter and spring, prior to the annual monsoon) of greatest resource stress. We established a camera-trapping grid in the San Andres, 2007–2011, and modeled occupancy of the three carnivores as a function of habitat, prey, and presence of the other carnivore species. Species interaction factors were >1.3 for each pair of carnivores, and the presence of the other carnivore species never significantly influenced occupancy of any other carnivore. Similarly, occupancy of the San Andres landscape was positively correlated among all carnivores. Occupancy of pumas was most influenced by proximity of water; coyote occupancy was influenced by terrain ruggedness and presence of medium (primarily lagomorph) prey, and bobcat occupancy was influence primarily small prey and proximity to water. The three carnivores also did not show temporal partitioning in use of habitats. Rather than segregation driven by competition, predation, or despotism, our results appeared to reflect preferences for differing habitat characteristics between ambush and cursorial predators and preferred habitats for travel.     

Reassessment of the environmental model of developmental polyphenism in spadefoot toad tadpoles

Polyphenism is the expression of multiple, discrete phenotypes from one genotype, and understanding the environmental factors that trigger development of alternative phenotypes is a critical step toward understanding the evolution of polyphenism and its developmental control. While much is known about the ecology of the well-known carnivore/omnivore polyphenism in spadefoot toad tadpoles, the environmental cues for the development of the specialized carnivore phenotype are not completely clear. We examined 27 different experimental treatments in two spadefoot toad species and used over 1,000 tadpoles in an attempt to elucidate those cues. While only 44 carnivores developed in these treatments, they were concentrated at cooler water temperatures and a diet that included fairy shrimp. However, while a diet of fairy shrimp promoted carnivore development, it was not necessary for inducing carnivore development at lower and intermediate water temperatures. Evidence also suggested a role for social inhibition that limited the proportion of interacting tadpoles that become carnivores. Tadpoles of Spea multiplicata grew larger at cooler temperatures and larger when their diets included fairy shrimp, whereas tadpoles of S. bombifrons grew larger at warmer temperatures and when their diets did not include fairy shrimp. These results indicate that carnivore induction can occur through different cues and that our current model for carnivore development is too limited. Finally, we argue that the carnivore/omnivore spadefoot system is neither a polyphenism nor a polymorphism but is a continuously distributed plasticity.     

Recent history,current status,conservation and management of native mammalian carnivore species in Great Britain

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Evaluation of Noninvasive Genetic Sampling Methods for Felid and Canid Populations

Abstract: Noninvasive sampling methods provide a means for studying species such as large mammalian carnivores that are difficult to survey using traditional techniques. Focusing on bobcat (Lynx rufus), we compared the effectiveness of noninvasive hair and scat genetic sampling in terms of field sample collection, species identification, and individual identification. We describe a novel hair-snare design and sampling protocol that successfully sampled 4 sympatric carnivore species, bobcat, mountain lion (Felis concolor), coyote (Canis latrans), and gray fox (Urocyon cinereoargenteus), in 3 habitat blocks in coastal southern California, USA. Scat surveys were also successful at sampling bobcats and other carnivores in the area. Hair and scat sampling methods had similar species identification success (81% and 87%, respectively) using mitochondrial DNA amplification and restriction enzyme digestion patterns. Therefore, for studies focused on the distribution and activity of a suite of carnivore species, we recommend a combination of noninvasive methodologies, for example, targeting hair and scat surveys toward species and sites where they are most effective. Because of a higher success rate for scat (85%) than for hair samples (10%) when using 4 microsatellite loci and a multiple-tubes approach to verify individual genotypes, we suggest scat sampling is a better choice for studies that require individual identification of bobcats.     

Spatial and temporal adjustments allowing the coexistence among carnivores in Liguria (N-W Italy)

We investigated the seasonal spatial and temporal co-occurrence of three carnivore species in Liguria region (NW Italy)—the red fox (Vulpes vulpes), the European badger (Meles meles) and the wolf (Canis lupus)—using the information provided by camera-trapping monitoring. Data were collected from January 2013 to January 2015 by positioning camera traps in 200 sample stations. During 3479 trap days, we collected 1048 independent videos of target carnivore species, which revealed a general spatial coexistence among carnivores with some differences in seasonal occurrence of species. The red fox and the European badger showed temporal segregation, as their activity patterns suggested a differential use of night-time in all seasons. Activity patterns of the red fox and the wolf revealed moderate-high overlap and similar density distributions in all seasons except during winter. Coexistence between these species may be allowed by temporal segregation during winter and spatial segregation during spring. Finally, results regarding the European badger and the wolf suggest a moderate temporal segregation with a marked avoidance effect for the European badger induced by the presence of tracks left by wolves. Programmes aimed at carnivore conservation, and management should treat the entire guild, as it has been demonstrated that populations of different carnivores interact with each other in complex ways and that fine-scale mechanisms regulating carnivore assemblage influence different aspects of natural communities.     

Diet choice of large carnivores in northern Cameroon

The diet of African wild dogs, lions and spotted hyenas was studied in the woodland savannah of Faro National Park and surrounding hunting zones in northern Cameroon during the dry season. Faecal analysis revealed that the diet of the three large carnivores overlaps considerably. Frequencies of prey remains of large and medium‐sized antelopes were highest, with Buffon's kob being by far the most common prey species. Carnivores seem to consume the most abundant prey species. No prey item of domestic livestock was found in the faeces. Faecal analysis has proved to be a useful method to reveal diet choice for large carnivores in northern Cameroon. Detailed information on the distribution and density of large carnivores and their prey is needed to give a better picture of the status of carnivores in northern Cameroon and to help resolve conflicts with livestock. Further investigations into the seasonal variation of predation and other factors structuring the large carnivore community in northern Cameroon are recommended.     

Drivers of flea (Siphonaptera) community structure in sympatric wild carnivores in northwestern Mexico

Host identity, habitat type, season, and interspecific interactions were investigated as determinants of the community structure of fleas on wild carnivores in northwestern Mexico. A total of 540 fleas belonging to seven species was collected from 64 wild carnivores belonging to eight species. We found that the abundances of some flea species are explained by season and host identity. Pulex irritans and Echidnophaga gallinacea abundances were significantly higher in spring than in fall season. Flea communities on carnivore hosts revealed three clusters with a high degree of similarity within each group that was explained by the flea dominance of E. gallinacea, P. simulans, and P. irritans across host identity. Flea abundances did not differ statistically among habitat types. Finally, we found a negative correlation between the abundances of three flea species within wild carnivore hosts. Individual hosts with high loads of P. simulans males usually had significantly lower loads of P. irritans males or tend to have lower loads of E. gallinacea fleas and vice‐versa. Additionally, the logistic regression model showed that the presence of P. simulans males is more likely to occur in wild carnivore hosts in which P. irritans males are absent and vice‐versa. These results suggest that there is an apparent competitive exclusion among fleas on wild carnivores. The study of flea community structure on wild carnivores is important to identify the potential flea vectors for infectious diseases and provide information needed to design programs for human health and wildlife conservation.