Late Pleistocene leopards as a bone accumulator: taphonomic results from S’Espasa cave and other Iberian key sites

Abstract

The Pleistocene faunal accumulations documented in caves have commonly been attributed to the activity of humans or carnivores. According to the palaeontological and archaeological literature, cave hyena (Crocuta spelaea) was the main known bone accumulator in karstic environments. However, in recent times, the role of leopards as bone accumulators has been revealed, and recent research has identified this behaviour in the Iberian Pleistocene. Moreover, there are other caves where leopard could have been claimed as an accumulator such as S’Espasa. In this work we present its taphonomic study. This cave was compared with the actualistic studies of leopards. Besides, the site of S’Espasa was compared with the other leopard dens in the Iberian Peninsula. These sites present faunal assemblages composed mainly by leopard (Panthera pardus) and Iberian wild goat (Capra pyrenaica), the bones of this ungulate present a similar pattern of bone modification by carnivores, skeletal survival rate, and bone breakage. These features indicate that goats could have been accumulated by leopards. With the data from this work and the previous ones, we try to establish a pattern that will help in the future to identify other accumulations created by this big cat.     

The effect of competing carnivores on the feeding behaviour of leopards (Panthera pardus) in an African savanna

Knowledge of competition dynamics among Africa’s large carnivores is important for conservation. However, investigating carnivore behaviour in the field can be challenging especially for species that are difficult to access. Methods that enable remote collection of data provide a means of recording natural behaviour and are therefore useful for studying elusive species such as leopards (Panthera pardus). Camera traps and Global Positioning System (GPS) collars are powerful tools often used independently to study animal behaviour but where their data are combined, the interpretation of a species’ behaviours is improved. In this study we used data from baited camera trap stations to investigate the feeding habits of leopards at Malilangwe Wildlife Reserve, Zimbabwe. We investigated the influence of spotted hyenas, lions and other competing leopards on the feeding duration of leopards using Generalized Linear Mixed Effects Modelling. To test the influence of competing predators on resting distances from bait sites, eight leopards were fitted with GPS collars. Results showed that leopards spent the shortest time feeding on the baits in the presence of competing male leopards compared to other predators while lion presence caused animals to rest farthest from bait sites. Interaction analysis indicated that small‐bodied leopards spent significantly shorter durations feeding when spotted hyenas were present. Our findings demonstrate that competition from guild carnivores has negative impacts on the food intake of leopards, which may have implications for fitness and survival. This study provides a snapshot of the competition dynamics at bait sites which may give insight to ecosystem level interactions among large carnivores in savanna ecosystems.     

The largest European lion Panthera leo spelaea (Goldfuss 1810) population from the Zoolithen Cave,Germany: specialised cave bear predators of Europe

Remains of 13 individuals with 3/1 male/female ratio of the extinct Upper Pleistocene lion Panthera leo spelaea (Goldfuss, 1810) from the Zoolithen Cave near Burggeilenreuth (Bavaria, Germany) include the holotype skull and all paratype material. The highest mortality rate for the Zoolithen Cave lions is in their reproductive adult ages. Bite marks on lion bones or skulls are results of hyena activities, or rare cannibalism of lions under stress situations. Lions were possibly also killed in battles with cave bears during predation on hibernating bears in winter times. This cave bear hunt specialisation in caves overlaps with the ecological behaviour of cave bear feeding by Ice Age-spotted hyenas. Both largest Ice Age predators, lions and hyenas, had to specialise on feeding herbivorous cave bears in boreal forest mountainous cave rich regions, where the mammoth steppe megafauna prey was absent. This cave bear hunt by felids, and scavenging by hyenas and other large carnivores such as leopards and wolves explains why cave bears hibernated deep in to the European caves, for protection reasons against predators. Within such lion–cave bear and even lion–hyena conflicts in the caves lions must have been killed sometimes, explaining mainly the skeleton occurrences in different European caves.     

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy,palaeobiology and sexual dimorphism

Remains of the steppe lion Panthera leo spelaea (Goldfuss) from historical digs in the Bilstein Caves of Warstein (Sauerland, NW Germany) are described. Their age seems to be from the Early Weichselian periods (Upper Pleistocene). Whereas the Bilstein cave was inhabited by cave bears at that time only a few hyena prey remains, were most likely imported into the cave entrance by hyenas. Bite and crush marks on a few bones of Bison priscus, Bos primigenius, Cervus elaphus, a rhinoceros Coelodonta antiquitatis vertebra and even several chewed cave bear bones prove the hyena presence which is similar to other caves in the Sauerland hyena den cave rich region. Additionally some larger wolves subspecies Canis lupusspelaeus bones were found, but only few Crocuta crocuta spelaea remains are present. After taphonomic comparisons to six other hyena and cave bear den caves of northern Germany, this cave can be classified as a cave bear den, which was briefly used by hyenas only for food storage or commuting or cave bear predation site in one part of the Cave. The lion material refers at least to one young adult lioness, one more adult female and two male lions; therefore, at minimum, the remains of four adult individuals are represented. The absence of juvenile lion material, in contrast to cave bear cub remains in the Bilstein Caves, proves that P. leo spelaea did not use this and all other caves in the region to raise their cubs. The bone material from the Bilstein Caves would prove the same hyena-lion antagonism conflict being recently proven for the Perick Caves, Balve Cave or Martins Cave well. Other situations in caves such as the Keppler Cave and the Bilstein Cave initially show the more complex taphonomic situation of lion remains in European caves, especially in cave bear dens, where they seem to have hunted periodically cave bears, such as it is already proven for hyenas in the Sauerland Karst and other caves of Europe.     

Ancient mitochondrial genomes from Chinese cave hyenas provide insights into the evolutionary history of the genus Crocuta

Cave hyenas (genus Crocuta) are extinct bone-cracking carnivores from the family Hyaenidae and are generally split into two taxa that correspond to a European/Eurasian and an (East) Asian lineage. They are close relatives of the extant African spotted hyenas, the only extant member of the genus Crocuta. Cave hyenas inhabited a wide range across Eurasia during the Pleistocene, but became extinct at the end of the Late Pleistocene. Using genetic and genomic datasets, previous studies have proposed different scenarios about the evolutionary history of Crocuta. However, causes of the extinction of cave hyenas are widely speculative and samples from China are severely understudied. In this study, we assembled near-complete mitochondrial genomes from two cave hyenas from northeastern China dating to 20 240 and 20 253 calBP, representing the youngest directly dated fossils of Crocuta in Asia. Phylogenetic analyses suggest a monophyletic clade of these two samples within a deeply diverging mitochondrial haplogroup of Crocuta. Bayesian analyses suggest that the split of this Asian cave hyena mitochondrial lineage from their European and African relatives occurred approximately 1.85 Ma (95% CI 1.62–2.09 Ma), which is broadly concordant with the earliest Eurasian Crocuta fossil dating to approximately 2 Ma. Comparisons of mean genetic distance indicate that cave hyenas harboured higher genetic diversity than extant spotted hyenas, brown hyenas and aardwolves, but this is probably at least partially due to the fact that their mitochondrial lineages do not represent a monophyletic group, although this is also true for extant spotted hyenas. Moreover, the joint female effective population size of Crocuta (both cave hyenas and extant spotted hyenas) has sustained two declines during the Late Pleistocene. Combining this mitochondrial phylogeny, previous nuclear findings and fossil records, we discuss the possible relationship of fossil Crocuta in China and the extinction of cave hyenas.     

Which predators are responsible for faunal accumulations at the Late Pleistocene layers of El Harhoura 2 Cave (Témara,Morocco)?

El Harhoura 2 cave (Témara, Morocco) has yielded Aterian and Iberomaurusian lithic artifacts associated with faunal remains. Both humans and carnivores occupied this cave and non-human predator modifications occurred mainly at the end of the Late Pleistocene. Diverse faunal taxa have been identified, with a predominance of gazelles and various carnivores, particularly canids. The location of the cave and of the excavation area, at the bottom of a cliff of low elevation and in the entrance of the cave, does not correspond to a protected area for large raptor nests or a natural trap. Considering the consumed species, the type of carnivore remains, the skeletal representation of prey, taphonomic alterations such as tooth marks, semi-digested bones and destruction sequences, large canids would be the main cause for faunal modifications. However, North African fossil data attributed to hyena activities present similar results. This paper highlights the difficulty of discriminating between potential accumulators/consumers due to a lack of taphonomic reference data. This study thus demonstrates the necessity of compiling fossil records and neotaphonomic reference data for North African medium-large predators in order to better understand the taphonomic history of North African archaeological and paleontological sites.     

Effects of temporary closure of a national park on leopard movement and behaviour in tropical Asia

Across Asia protected areas serve as refuges for carnivores inside human-dominated landscapes. However, the creation of hard edges around reserve boundaries where conflicts with humans arise and disturbance from human activities inside the reserves may affect carnivore behaviour and ecology. Thailand’s largest protected area, Kaeng Krachan National Park (2915 km²) receives >100,000 visitors annually while maintaining an intact assemblage of prey species for large carnivores, making it a potentially important site for population recovery of leopards (Panthera pardus), tigers (Panthera tigris) and dholes (Cuon alpinus). We assessed the abundance of leopards and their prey base, and their response to changes in levels of human activity after an unexpected flooding event that resulted in the park being closed to visitors for >6 months. Using camera-traps, we identified 6 individual leopards and used spatially explicit capture-recapture (SECR) methods, incorporating humans and prey as covariates, to test for factors affecting the detection probability of leopards before and after the park closure. Leopard density was unchanged between the two periods, however the movement and activity patterns were clearly different. In the absence of tourist activity, leopards tended to move more frequently, leopard detection rates increased by 70% and activity shifted towards being more diurnal. The consequences of these changes in behaviour may include improved health, reproduction and survival. A management strategy involving seasonal closure of parks may serve to alleviate pressure on leopards and other carnivores. We recommend using information on abundance of large carnivores and their prey species, and human disturbance as the key indicators for long-term monitoring and management of protected areas in Southeast Asia.     

Étude taphonomique, paléoécologique et archéozoologique des faunes de grands mammifères de la seconde moitié du Pléistocène inférieur de la grotte du Vallonnet (Roquebrune-Cap-Martin, Alpes-Maritimes, France)

Vallonnet cave (Alpes-Maritimes, France) provided archaic lithic tools together with a big amount of large mammals of the end of lower pleistocene. At the crossing of diverse ecological niches in time and space, three main levels and within them several smaller units were distinguished. The poorly mineralized bones witnessed a huge diagenetic fragmentation. The presence of the animals in the cave seems to be linked to the reasons of their death. The exhaustive and systematic observation of the bones, like the bears probably used to die inside the cave. Some other carnivores especially hyenas, used the cave as a nursery. Man also occupied the cave and had an activity mainly on large cervid bones and on some other species as well. The very progressive deposit gave a way for carnivores and porcupine to gnaw the bones on the ground before their burial. The succession of actions (man, carnivores and porcupine) on the same bones is characteristic on Vallonnet large mammals bones.     

Small and large mammals from the Ciota Ciara cave (Borgosesia,Vercelli, Italy): An Isotope Stage 5 assemblage

The Ciota Ciara cave is located within the Monte Fenera karst system (Borgosesia, Vercelli, Italy) at 670 m a.s.l. The cave entrance presents a deposit with Mousterian quartz and flint industry. The faunal remains from Stratigraphic Units 13, 14 and 103 are the subject of this work and are presented here as a whole for the first time. The large mammal assemblage is dominated by Ursus spelaeus. In addition, a few remains of carnivores such as Panthera leo spelaea, Panthera pardus, Meles meles have been found together with ungulates such as Cervus elaphus and Rupicapra rupicapra. The small mammal assemblage is characterized by a high biodiversity, especially in bats, by the dominance of Clethrionomys glareolus and by a relatively large number of Pliomys coronensis, a species that seems to disappear from the Italian Peninsula at the end of Marine Isotope Stage 5 or during the beginning of MIS 4. The changes in frequency of the small and large mammals between the two S.U. suggest a change from a relatively cold-humid (S.U. 14) to a warmer-still humid climate (S.U. 13). Although no radiometric dates are available yet, the small and large mammal assemblages, the presence of Pliomys coronensis and the climate change inferred by the variation of the small mammal frequencies allow us to correlate these two units of the Ciota Ciara cave to a relatively warm moment of Marine Isotope Stage 5.     

Phylogenetic characterisation of Taenia tapeworms in spotted hyenas and reconsideration of the “Out of Africa” hypothesis of Taenia in humans

The African origin of hominins suggests that Taenia spp. in African carnivores are evolutionarily related to the human-infecting tapeworms Taenia solium, Taenia saginata and Taenia asiatica. Nevertheless, the hypothesis has not been verified through molecular phylogenetics of Taenia. This study aimed to perform phylogenetic comparisons between Taenia spp. from African hyenas and the congeneric human parasites. During 2010–2013, 233 adult specimens of Taenia spp. were collected from 11 spotted hyenas in Ethiopia. A screening based on short DNA sequences of the cytochrome c oxidase subunit 1 gene classified the samples into four mitochondrial lineages designated as I–IV. DNA profiles of nuclear genes for DNA polymerase delta (pold) and phosphoenolpyruvate carboxykinase (pepck) showed that lineages II and III can be assigned as two independent species. Common haplotypes of pold and pepck were frequently found in lineages I and IV, suggesting that they constitute a single species. Morphological observations suggested that lineage II is Taenia crocutae, but the other lineages were morphologically inconsistent with known species, suggesting the involvement of two new species. A phylogenetic tree of Taenia spp. was reconstructed by the maximum likelihood method using all protein-coding genes of their mitochondrial genomes. The tree clearly demonstrated that T. crocutae is sister to T. saginata and T. asiatica, whereas T. solium was confirmed to be sister to the brown bear tapeworm, Taenia arctos. The tree also suggested that T. solium and T. arctos are related to two species of Taenia in hyenas, corresponding to lineages I + IV and III. These results may partially support the African origin of human-infecting Taenia spp., but there remains a possibility that host switching of Taenia to hominins was not confined to Africa. Additional taxa from African carnivores are needed for further testing of the “Out of Africa” hypothesis of Taenia in humans.     

A survey for Echinococcus spp. of carnivores in six wildlife conservation areas in Kenya

To investigate the presence of Echinococcus spp. in wild mammals of Kenya, 832 faecal samples from wild carnivores (lions, leopards, spotted hyenas, wild dogs and silver-backed jackals) were collected in six different conservation areas of Kenya (Meru, Nairobi, Tsavo West and Tsavo East National Parks, Samburu and Maasai Mara National Reserves). Taeniid eggs were found in 120 samples (14.4%). In total, 1160 eggs were isolated and further analysed using RFLP-PCR of the nad1 gene and sequencing. 38 of these samples contained eggs of Echinococcus spp., which were identified as either Echinococcus felidis (n = 27) or Echinococcus granulosus sensu stricto (n = 12); one sample contained eggs from both taxa. E. felidis was found in faeces from lions (n = 20) and hyenas (n = 5) while E. granulosus in faeces from lions (n = 8), leopards (n = 1) and hyenas (n = 3). The host species for two samples containing E. felidis could not be identified with certainty. As the majority of isolated eggs could not be analysed with the methods used (no amplification), we do not attempt to give estimates of faecal prevalences. Both taxa of Echinococcus were found in all conservation areas except Meru (only E. felidis) and Tsavo West (only E. granulosus). Host species identification for environmental faecal samples, based on field signs, was found to be unreliable. All samples with taeniid eggs were subjected to a confirmatory host species RLFP-PCR of the cytochrome B gene. 60% had been correctly identified in the field. Frequently, hyena faeces were mistaken for lion and vice versa, and none of the samples from jackals and wild dogs could be confirmed in the tested sub-sample. This is the first molecular study on the distribution of Echinococcus spp. in Kenyan wildlife. The presence of E. felidis is confirmed for lions and newly reported for spotted hyenas. Lions and hyenas are newly recognized hosts for E. granulosus s.s., while the role of leopards remains uncertain. These data provide the basis for further studies on the lifecycles and the possible link between wild and domestic cycles of cystic echinococcosis in eastern Africa.     

A clan of Late Pleistocene hyenas,Crocuta crocuta spelaea (Goldfuss 1823), from the Rösenbeck Cave (Germany) and a contribution to cranial shape variability

The remains of a large population of Late Pleistocene Ice Age spotted hyenas (Crocuta crocuta spelaea Goldfuss 1823) are described from the Rösenbeck Cave in the Sauerland Karst of Germany. They include four skulls and 79 other skeletal parts, mainly from adult to senile animals, making this an important Late Pleistocene hyena cave‐den site in Europe. The skulls have been compared with 30 other hyena skull specimens from open air and cave‐den sites in central Europe (Germany, Austria, the Czech Republic and Romania) in order to achieve an understanding of sexual dimorphism in the crania of Ice Age spotted hyenas from the Upper Pleistocene cold period (Weichselian/Wuermian), and the types of injuries that they acquired during their lifetimes. Three different types of cranial shape have been distinguished, one of which appears to have been a consequence of pathologies that developed in response to injuries caused by bites received during the animal's lifetime, as a result of either intraspecies fights or fights with lions. Although cave bears penetrated to great depths within the Rösenbeck Cave for hibernation purposes, hyenas appear to have utilized only a short section of the cave that branched off directly from the entrance area. Hyena cub material is scarce, suggesting that this area was used as a communal den rather than for cub rearing. Bones exhibiting gnaw marks, representing prey imported by hyenas, are also rare but include horse (Equus caballus przewalskii) and reindeer (Rangifer tarandus) remains. The scarcity of bones from hyena prey suggests that this cave was not used as a food storage site. Some Ursus spelaeus cave bear remains, including skulls, show evidence of having been gnawed, chewed and cracked by hyenas, indicating that the hyenas periodically fed on cave bear carcasses in a specialization response to the mammoth steppe megafauna absence of the boreal mountain forest regions. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 191–220.     

Les faunes des sites du Pléistocène supérieur du Caucase méridional : Grotte de Saka?hia, Grotte d’Ortvala et Grotte du Bronze (République de Géorgie)

The paleontological analysis of the fauna from three Late Pleistocene localities in Southern Caucasus (Saka?hia, Ortvala, and Bronze cave) confirmed the presence of Ursus spelaeus, Canis lupus, Equus ferus, Rhinocerotidae, Cervus elaphus, Capreolus capreolus, Alces alces, Bison priscus, Capra caucasica, Ovis ammon and Sus scrofa. The study also permitted the identification of new taxa for these localities. There may be the presence of reindeer (Rangifer tarandus) at Saka?hia, however it has not yet been confirmed. Moreover, remains of aurochs (Bos primigenius) were identified, in particular at Bronze cave where its abundance is exceptional. Indeed, Bos and Bison are generally very rare in Southern Caucasus. The analysis of the proportion of the different species in each locality revealed the two types of occupations of the caves, one dominated by hominids and the other by carnivores. At Saka?hia, where fauna is dominated by the presence of cave bear, the cave was seasonally occupied by Neanderthal groups. On the other hand, Bronze cave corresponds to a habitat of hunters, which occupied the site for longer periods.     

Micromammals (rodents and insectivores) from the early Late Pleistocene cave site of Bois Roche (Charente, France): Systematics and paleoclimatology

We present the study of rodents and insectivores from the 1995 season of excavation at the early Late Pleistocene site of Bois Roche (Charente, France). The site is a small cave with a low ceiling, used as a den by hyenas. It contains a large herbivore assemblage (mainly bovids and equids) with a smaller representation of medium-size taxa (cervids), many microvertebrates, abundant coprolites and deciduous hyena teeth, and a few lithic artifacts (Middle Paleolithic) introduced into the cave by gravity and slope wash. The rodent fauna consists of Eliomys quercinus cf. quercinus, Apodemus sp., Microtus gregalis, Microtus oeconomus, Arvicola terrestris and Dicrostonyx torquatus. Insectivores are represented by Neomys cf. fodiens. The most common species is M. gregalis which forms 93.7% of the total MNI. The micromammal fauna suggests an open landscape, with some vegetation mainly of the steppe or tundra type, with areas of water and more wet vegetation and some areas with trees. The micromammal association of Bois Roche is characteristic of a cold or very cold dry climate.     

Carnivores from Los Rincones,a leopard den in the highest mountain of the Iberian range (Moncayo,Zaragoza, Spain)

Carnivores are a common element in Pleistocene fossil assemblages. However, they are not so abundant in terms of the numbers of remains. Here, we present a palaeontological study of the carnivores from the Late Pleistocene (MIS 3) of Los Rincones (Spain), one of the few deposits accumulated by leopards. One hundred and ten leopard remains have been recovered. This carnivore is not the only inhabitant of the cave, and 175 remains belonging to Ursus arctos have also been recovered, making it one of the sites with the greatest number of brown bear remains in the Iberian Peninsula. The large number of leopard remains has allowed us to make a detailed morphological and biometrical study that has enabled us to classify the remains within the subspecies Panthera pardus spelaea. The European Ice Age leopard inhabited Europe during Upper Pleistocene and it presents some similarities with Panthera uncia. A study of the scarce remains of Canis lupus indicates that this was similar in size to Canis lupus maximus; the scarcity of the remains prevents us from assigning our remains to this subspecies. A study of the brown bear remains indicates that it is similar to other populations in the north of the Iberian Peninsula with this chronology.     

The Crocuta crocuta spelaea (Goldfuss 1823) population and its prey from the Late Pleistocene Teufelskammer Cave hyena den besides the famous Paleolithic Neandertal Cave (NRW,NW Germany)

Remains from at least seven individuals of the Late Pleistocene Ice Age spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) from the Teufelskammer Cave in the Neandertal valley (North Rhine-Westphalia, northwest Germany) are described. The small cave was a well-frequented hyena den of the Early to Middle Late Pleistocene which was only 100 m from the famous small Feldhofer Cave, where the first Neandertal human skeleton was found. The high amount of hyena bone material (37%) and its strongly chewed and incomplete prey remains of the mixed mammoth steppe and boreal forest megafauna prove one more of 11 recently known hyena den caves in the Rhenish Massif. Hyenas and cave bears have used the cave, but Neandertal humans lived possibly not at the same time in the same valley. Although hyenas occupied mainly the smaller caves such as the Teufelskammer Cave, humans preferred large portal cave entrances such as in the Neandertal valley with the Small Feldhofer Cave.     

The TD6.3 faunal assemblage of the Gran Dolina site (Atapuerca,Spain): a late Early Pleistocene hyena den

Ethological studies have shown that besides human groups, large-medium carnivores have bone-collecting habits. The research developed since the last half of the twentieth century has attempted to characterise the carnivore’s accumulations and to identify them in the archaeo-paleontological record. At present, we have diagnostic criteria that define the accumulations produced by hyenids (mainly, Crocuta crocuta), thereby allowing us to differentiate them from the other accumulating agents. The faunal assemblage recovered at the Early Pleistocene TD6.3 layer of the Gran Dolina site is characterised by the presence of typical elements described in hyena dens: presence of small carnivores remains, high bone breakage, low epiphysis survival and a high frequency (>30%) of specimens with carnivore induced modifications, including large amounts of digested bones. However, attritional mortality profiles, hyena’s cubs remains, mid-shaft bone cylinders or differential anatomical composition among different weight-sizes, have not been observed or are ambiguous. In addition, anthropic evidences are scarce and concentrated in the uppermost section of the layer. TD6.3 is the result of an accumulation produced by hyenas using the cave as a den, in alternation with sporadic occupations by hominin groups. TD6.3 shows that hyena fossil accumulations may present great anatomical and taxonomic variability.     

Rabies Virus and Canine Distemper Virus in Wild and Domestic Carnivores in Northern Kenya: Are Domestic Dogs the Reservoir?

Rabies virus (RV) and canine distemper virus (CDV) can cause significant mortality in wild carnivore populations, and RV threatens human lives. We investigated serological patterns of exposure to CDV and RV in domestic dogs (Canis familiaris), African wild dogs (Lycaon pictus), black-backed jackals (Canis mesomelas), spotted hyenas (Crocuta crocuta), striped hyenas (Hyaena hyaena) and African lions (Panthera leo), over a 10-year period, in a Kenyan rangeland to assess the role domestic dogs may play in the transmission dynamics of these two important canid pathogens. Observed patterns of RV exposure suggested that repeated introduction, rather than maintenance, occurred in the wild carnivore species studied. However, RV appeared to have been maintained in domestic dogs: exposure was more likely in domestic dogs than in the wild carnivores; was detected consistently over time without variation among years; and was detected in juveniles (≤1-year-old) as well as adults (>1-year-old). We conclude that this domestic dog population could be a RV reservoir. By contrast, the absence of evidence of CDV exposure for each carnivore species examined in the study area, for specific years, suggested repeated introduction, rather than maintenance, and that CDV may require a larger reservoir population than RV. This reservoir could be a larger domestic dog population; another wildlife species; or a “metareservoir” consisting of multiple interconnected carnivore populations. Our findings suggest that RV risks to people and wild carnivores might be controlled by domestic dog vaccination, but that CDV control, if required, would need to target the species of concern.     

Les petits ongulés de la grotte acheuléano-moustérienne du Lazaret (Nice,France) – Origine anthropique ou carnivore ?

Lazaret Cave contains archaeological deposits contemporaneous with Acheulean and Mousterian cultures, corresponding to marine isotopic stage 6 (MIS 6). Archaeological excavations at the site revealed twenty-nine archaeostratigraphic units with recurring, relatively long occupations by Homo heidelbergensis groups. These groups preferentially hunted red deer and ibex, which were abundant in all the units. In contrast, other species, such as the chamois and the roe deer, are poorly represented. In this article, we present the results of the archaeozoological and taphonomic analyses of small ungulate bones in order to determine how these remains were deposited in the cave. Opportunistic anthropogenic hunting is brought to light in some units, explaining the low proportion of these species at the site. In addition, these species only provide low nutrient levels for the needs of the group. The accumulation of remains by carnivores is also observed, mainly by the wolf, which is abundant in the upper units.     

Peri-urban spotted hyena (<Emphasis Type="Italic">Crocuta crocuta</Emphasis>) in Northern Ethiopia: diet,economic impact,and abundance

Global declines of carnivores are related to difficult integration with human land use, in particular conflicts caused by livestock depredation. Spotted hyenas (Crocuta crocuta) live in remarkably close proximity to humans in the degraded and prey-depleted Enderta district, northern Ethiopia. Their diet and interaction with people were investigated in sub-districts close to the regional capital, Mekelle. We interviewed 1,686 randomly selected households from three sub-districts, Debri, Aynalem, and Felege Selam, about livestock management and incidence of depredation from 2005 to 2009. Livestock loss amounted to 492 heads over 5 years; an annual mean of 0.6% worth US$ 7,042. We also performed a survey giving a minimum population estimate of 60 hyenas in the three sub-districts; all but four were found in church forests where they are traditionally tolerated and protected. A total of 1,200 hyena scats were analyzed to determine prey species; the diet contained only domestic species, with sheep being by far the most common prey species. About 5.5% of fecal analysis contained human hairs. We conclude that hyenas depended entirely on domestic prey species, partly through depredation but more importantly through scavenging on (peri-) urban waste. Under the particular local circumstances, continued coexistence appears possible, provided that damage remains tolerable.