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The present is a palaeobiological and taphonomic analysis of a Marine Isotope Stages (MIS) 4–3 (Late Pleistocene) assemblage of animal remains and hominin artefacts from layers 7–5 of Biśnik Cave, Czestochowa Upland, Poland. The analysis indicates that the bone assemblage is the result of a time-averaged palimpsest of both biotic and episodic abiotic events, the former consisting of many successive generations of animals and hominins that frequented the cave, and the latter including hydraulic winnowing. In fact, the taphonomic history of the fossil assemblage from Biśnik Cave's layers 7–5 is partially obscured by the overprint of hydraulic winnowing, which purportedly removed a certain amount of the original specimens. Besides evidence of cave bear deaths from non-violent, hibernation-related mortality and of occupation by generations of denning wolves and hyaenas, there is a wealth of flint artefacts, alongside remains of a few fireplaces and of a structure built in the cave by hominins to partition the cave chambers. The studied layer contains an impressive number of shed antlers, primarily of the red deer Cervus elaphus. Crocuta crocuta spelaea is normally held responsible for such accumulations of shed antlers in various European caves; Biśnik Cave's layers 7–5 will therefore simply add to the list. However, the role of accumulator of shed antlers attributed to the Pleistocene spotted hyena does not match the behaviour of its modern counterpart and seems not accounted for metabolically. The only reasonable alternative is that the antlers were collected by hominins. From this alternative perspective the cave would have functioned as a warehouse, where naturally shed antlers were stored as raw material, potentially to be shaped into tools and/or employed as tools to make other tools. The palaeobiological and taphonomic analysis presented here provides new insights into the succession of pre- and postdepositional events that involved the bone remains accumulated in the cave, as well as into the interactions between the animals and hominins of the time. More importantly, if hominins, and not hyaenas, were responsible for the amassment of the shed antlers in Biśnik Cave, this study raises doubts as to the hyaenid or human origin of other similar cave accumulations of shed antlers throughout Europe.

This paper focuses on new findings from Middle Paleolithic Geula Cave, Israel, located in the northern part of Mt. Carmel. The cave, consists of several small chambers that are remnants of a larger cave system, initially excavated between 1958 and 1964. The occupation at the cave was ascribed to Middle Paleolithic modern humans. In 2016, a salvage excavation was conducted in areas of the cave that were not previously explored. Analyses of the new excavation revealed a consistency in lithic technology throughout the new excavation areas, with an emphasis on Levallois production using mainly bidirectional and centripetal core preparation modes. The faunal study identified intensive hyena and porcupine activities in small chambers, probably at the back part of the cave, while fractured bones in association with flint artifacts were found at what appears as the living area in the cave. Luminescence ages indicated that Geula Cave was occupied as early as 175 ka, but that the major occupation was from 120 to 100 ka. The major occupation at Geula chronologically overlaps with the Skhul and Qafzeh Caves thus reinforcing the notion that Homo sapiens dominated the southern Levant during early MIS 5. This study demonstrates the importance of reinvestigating and reevaluating past excavated prehistoric sites and their contents to enhance our understanding of the regional, cultural and biological history.

The dire wolf was one of the successful top predators in North America during the Pleistocene. It is best known from the southern part of North America, and it even immigrated to South America. Fossils of dire wolves are very rare north of 42° North latitude in North America. That distribution supported the belief that the low temperatures and ice sheets in the higher latitudes of North American formed an insurmountable barrier for the dispersal of dire wolves. Here we report the first record of the dire wolf fossil in Eurasia. The fossil is a partial mandibular fragment with a tooth recovered from a Late Pleistocene underwater sand mine site near the city of Harbin in Northeastern China. Other mammalian fossils from the same site suggest that this dire wolf coexisted with the typical Mammuthus-Coelodonta fauna of Eurasia. The newly discovered specimen has a huge m1; much larger than gray wolves and other large canids from the same region. The massive m1 trigonid accounts for a high percentage of its overall length. The m1 talonid is reduced, but retains a small entoconid. The m2-3 alveoli suggest that the two teeth were smaller than those of the gray wolf. The combination of these morphologies is present only in the dire wolf. When the body mass spectra of medium and large-bodied carnivores from Asia and North America were compared, the Asian dire wolf occupies a position nearly overlapping the spotted hyena (Crocuta crocuta ultima). During the Late Pleistocene, C. crocuta ultima was one of the most widely distributed and dominant carnivores in Asia. Competition from hyenas may have kept dire wolf Asian populations at very low levels leading to their rarity as fossils rare in Asia.

The giant short-faced hyena, Pachycrocuta brevirostris, was the largest and most spectacular bone-cracking hyena that ever existed. Although already extinct long ago, it was extensively distributed across Eurasia during the Early and Middle Pleistocene. P. brevirostris is remarkable for a series of iconic craniodental adaptations in sway with a commitment to scavenging off carcasses of other large herbivores, such as the powerfully built skull and massive premolars, yet the most eye-catching aspect is the enormous size as emphasized by its vernacular name. The largest hitherto known skull of this species is the holotype from Sainzelles in Auvergne, France, discovered over a century ago. We hereby report a new discovery from Jinyuan Cave in Dalian, China, which slightly surpasses the holotype in craniodental dimension, posing a new record in Pachycrocuta body size. Three skull fragments of the same individual unearthed from the site are described in detail herein and compared with other P. brevirostris discoveries from both Europe and Asia. Thorough comparative studies show that the newly discovered materials are very close to the holotype from Sainzelles both metrically and morphologically, which fully indicates that the European morphotype, namely Pachycrocuta brevirostris brevirostris, surprisingly appeared in Northeast Asia as well, and further challenges the conventional concept of the population or subspecies. The giant hyenas from Asia were originally proposed as independent species, such as Pachycrocuta licenti or P. sinensis, but now broadly accepted as conspecifics of the European Pachycrocuta brevirostris, forming only different subspecies. The occurrence of P. b. brevirostris in Northeast Asia forcefully demonstrates that the differentiation is more of temporal than geographical significance, i.e., the subspecies of Eurasia basically constituted an ancestor-descendant sequence, from licenti to sinensis via brevirostris. Moreover, represented by the newly discovered specimens, the forerunner of P. b. brevirostris with transitional features appeared in Northeast Asia as early as 2.0 Ma, which implies the population of Europe most likely originated and dispersed from Northeast Asia. A hypothesis of “out of Northeast Asia” is hereby proposed for P. b. brevirostris. In the present paper, a series of diagnostic craniodental characters, such as carnassial morphology, the accessory cuspids of the premolars and the shape of the skull profile, are also discussed and re-valuated to investigate their potential significance in phylogenetic analysis.

This paper presents a Multi-threaded version of the Spotted Hyena Optimizer algorithm with thread-crossing techniques (MT-SHO) to improve the ability of the algorithm to explore the search space. The original algorithm is inspired by the hunting behavior of the spotted hyena. Along the different sections of the work, we explain in detail how the original algorithm simulates the spotted hyena’s behavior to optimize highly complex mathematical functions and how we handle the procedures and results of the multi-threaded version, with thread-crossing techniques that improve the ability to explore and exploit the search space by letting threads learn between them. We present the experiments used to determine the best value of the parameters used in the parallel version of the algorithm and to prove that our proposal obtains significantly good results we compare the results obtained by evaluating 24 benchmark functions with the results published for the original algorithm as well as other metaheuristic algorithms.

Spotted hyena optimizer (SHO) is a novel metaheuristic optimization algorithm based on the behavior of spotted hyena and their collaborative behavior in nature. In this paper, we design a spotted hyena optimizer for training feedforward neural network (FNN), which is regarded as a challenging task since it is easy to fall into local optima. Our objective is to apply metaheuristic optimization algorithm to tackle this problem better than the mathematical and deterministic methods. In order to confirm that using SHO to train FNN is more effective, five classification datasets and three function-approximations are applied to benchmark the performance of the proposed method. The experimental results show that the proposed SHO algorithm for optimization FNN has the best comprehensive performance and has more outstanding performance than other the state-of-the-art metaheuristic algorithms in terms of the performance measures.

Palaeoenvironmental information on Marine Isotope Stage 3 (MIS 3) coastal Latium is sparse, mainly based on studies of individual faunal assemblages or long pollen records from lake sediments, often of insufficient resolution to aid in palaeoenvironmental reconstruction. This study describes in detail the Late Pleistocene faunal assemblage from layers SU11 and SU12 of Cava Muracci (Cisterna di Latina, central Italy), the first of which is a partially-preserved hyena den. The first multi-disciplinary palaeoenvironmental reconstruction of coastal Latium between 34–44 ka BP is provided combining palaeoecological inferences from a previous pollen analytical study of hyena coprolites with the palaeontological study described here. The results indicate a temperate climate and a landscape characterised by the coexistence of at least three habitats within a short distance between the coastline and the inland mountains, suitable for a wide variety of species. A wooded hilly area populated by wolfs, aurochs, cervids and other small vertebrates; wide steppe and grasslands inhabited by wild horses, rhinoceros and cave hyenas; lastly, wetland habitats along coasts and rivers populated by wild boars and several amphibians and reptiles.

Salivary hormone analyses provide a useful alternative to fecal and urinary hormone analyses in non-invasive studies of behavioral endocrinology. Here, we use saliva to assess cortisol levels in a wild population of spotted hyenas (Crocuta crocuta), a gregarious carnivore living in complex social groups. We first describe a novel, non-invasive method of collecting saliva from juvenile hyenas and validate a salivary cortisol assay for use in this species. We then analyze over 260 saliva samples collected from nearly 70 juveniles to investigate the relationships between cortisol and temporal and social variables in these animals. We obtain evidence of a bimodal daily rhythm with salivary cortisol concentrations dropping around dawn and dusk, times at which cub activity levels are changing substantially. We also find that dominant littermates have lower cortisol than either subordinate littermates or singletons, but that cortisol does not vary with age, sex, or maternal social rank. Finally, we examine how social behaviors such as aggression or play affect salivary cortisol concentrations. We find that inflicting aggression on others was associated with lower cortisol concentrations. We hope that the detailed description of our methods provides wildlife researchers with the tools to measure salivary cortisol in other wild carnivores.HIGHLIGHTSWe validated methods for collecting and analyzing saliva from wild carnivores.We documented a bimodal daily rhythm in juvenile spotted hyena salivary cortisol.Cortisol varied among juvenile hyenas based on litter size and intra-litter rank.Inflicting aggression on others was associated with lower cortisol concentrations.

This paper proposes a multi-objective version of recently developed Spotted Hyena Optimizer (SHO) called Multi-objective Spotted Hyena Optimizer (MOSHO). It is used to optimize the multiple objectives problems. In the proposed algorithm, a fixed-sized archive is employed for storing the non-dominated Pareto optimal solutions. The roulette wheel mechanism is used to select the effective solutions from archive to simulate the social and hunting behaviors of spotted hyenas. The proposed algorithm is tested on 24 benchmark test functions and compared with six recently developed metaheuristic algorithms. The proposed algorithm is then applied on six constrained engineering design problems to demonstrate its applicability on real-life problems. The experimental results reveal that the proposed algorithm performs better than the others and produces the Pareto optimal solutions with high convergence.

In the present study, we report brown hyena tooth marks on australopiths from Sterkfontein's Plio-Pleistocene-age Member 4 (South Africa). Classic taphonomic analyses and the implementation of new techniques, including Geometric Morphometrics and Machine Learning, are combined to identify the modifying agent and provide the first direct evidence of hyenid scavenging on australopiths. This hypothesis adds a new perspective to the relationships between carnivores and hominins in the Cradle by expanding on previous hypotheses proposing that leopards were the primary predator and bone accumulator of early hominin remains at South African palaeocaves.

The giant, short-faced hyena Pachycrocuta brevirostris was the largest Hyaenidae ever existed and the one that perfectly embodied the distinctive bone-cracking adaptations of this mammal family. Its dispersal into Europe is regarded as a biochronological marker of the Late Villafranchian at ∼2.0 Ma, and its potential ecological interactions with other carnivorans and early Homo populations diffusing Out of Africa have given rise to extensive discussions. Nevertheless, our comprehension of the extinction of P. brevirostris remains vague. Here, we first critically evaluate the European fossil record of the species and then we review the whole Epivillafranchian and Galerian Hyaenidae record, including P. brevirostris, Crocuta crocuta and “Hyaena” prisca. Biometric comparisons with other extinct and extant bone-cracking hyenas are carried out. In contrast to a common view, we recognize that there is neither evidence of a persistence of P. brevirostris beyond the Early-Middle Pleistocene boundary, nor of a coexistence between the giant hyena and C. crocuta. The replacement between the two species, which was also accompanied by the arrival of “H.” prisca, occurred at ∼0.8 Ma and can serve as a marker of the Epivillafranchian–Galerian turnover, part of the faunal renewal that reflects the response of mammal communities to the Early–Middle Pleistocene Transition. Moreover, we clarified that Pliocrocuta perrieri and “H.” prisca were different species, and that the latter was relatively more widespread than often assumed, being recorded from localities spanning in age almost the whole Middle Pleistocene and even the early Late Pleistocene.

Hunting preferences reveal a great deal about the life of Paleolithic humans, and may reflect changes in human demography, technology, and adaptations to changing environments. However, the effects of hunting preferences and environmental availability are often conflated, stressing the need for comparisons to other predators that exploited the same environment. Manot Cave (Israel), preserved rich Early Upper Paleolithic (46-33 ka) human occupations, along with repeated spotted hyena (Crocuta crocuta) occupations, allowing us to compare anthropogenic and biogenic bone assemblages within the same space and time frame. We focused on the faunal remains retrieved in the middle of the cave (Area D), and conducted detailed taphonomic and zooarchaeological analyses. The Area D archaeofauna was dominated by Mesopotamian fallow deer (Dama mesopotamica) and mountain gazelle (Gazella gazella), which showed high abundance of carnivore damage. This and the carnivore-to-ungulate ratio, the presence of juvenile hyena and numerous coprolites match the criteria of a hyena den, confirming that the bone assemblage was created mainly by hyenas. Manot Area D thus reveals hyena prey choice in the Upper Paleolithic Galilee, which we then compared with human prey choice. Our results showed that hyena prey assemblages in Manot and elsewhere in the Levant were Dama-dominated while human assemblages were dominated by Gazella, demonstrating that hyenas and humans hunted different animal size groups, possibly in different habitats. We interpret this phenomenon as resulting from two possible scenarios: the emergence of projectile technology which may favor hunting in open environments, and niche partitioning derived by human-hyena competition. Hyenas were abundant and hunted unselectively while the anthropogenic record presents population turnovers and some dietary diversification. Whereas both scenarios are difficult to test directly with the available evidence, we argue that either one explains quite well the late Pleistocene archaeofaunal patterns in the Levantine record.

This paper presents a novel metaheuristic algorithm named as Spotted Hyena Optimizer (SHO) inspired by the behavior of spotted hyenas. The main concept behind this algorithm is the social relationship between spotted hyenas and their collaborative behavior. The three basic steps of SHO are searching for prey, encircling, and attacking prey and all three are mathematically modeled and implemented. The proposed algorithm is compared with eight recently developed metaheuristic algorithms on 29 well-known benchmark test functions. The convergence and computational complexity is also analyzed. The proposed algorithm is applied to five real-life constraint and one unconstrained engineering design problems to demonstrate their applicability. The experimental results reveal that the proposed algorithm performs better than the other competitive metaheuristic algorithms.

Recent application of photogrammetric and geometric morphometric approaches to the study of cut marks on bones has yielded positive results in discriminating different types of tools and even some raw materials. Here, we apply this analytical technique to the study of carnivore tooth scores. The goal is twofold: on the one hand, we intend to differentiate carnivore types and on the other one, we show the application of this approach to a sample of tooth scores from long bones documented at the modern assemblage of the Olduvai Carnivore Site (OCS). Previous taphonomic work at OCS suggested that this bone assemblage constituted a good evidence of a carcass accumulation behavior by lions, followed by hyena ravaging. The application of these 3D techniques to the selected sample of tooth marks shows that lions, as well as spotted hyenas, did indeed impart marks on the OCS assemblage. This reinforces the dual nature of the assemblage and the implication of lions in its formation.

Coprolites are commonly identified in the Pleistocene archaeo-palaeontological record. They have often been described as indirect evidence for the presence of carnivores (usually hyenids) during the formation of a depositional sequence. However, coprolites are a much larger source of information that can provide data relating to factors affecting an archaeological assemblage, including its taphonomic history and palaeoecology. In this paper, two fossil coprolite accumulations dating to the late Early Pleistocene are described: Level TD6.1 of Gran Dolina (Sierra de Atapuerca, Burgos) and Unit II of la Mina (Barranc de la Boella, Tarragona). The results highlight the morphological homogeneity of these hyena coprolites, despite a considerable variation in size. The presence of microspherulites is clearly identified in all of the analysed hyena coprolites. Their composition is rich in elements characteristic of bone-origin, such as calcium (Ca) and phosphorus (P). A major difference, however, differentiating the remains from these sites was a higher frequency of bone fragments in the la Mina coprolites, regarding to TD6.1. Pollen, spores, and palynomorphs are scarce in the samples from both sites, which in fact share similar ecological characteristics such as the presence of pine, wild grasses, and coprophilous fungi. Hyena coprolites are relatively easy to differentiate from those of other taxa; however, they show intra-specific similarities in their micro- and macro-morphology and composition, despite their wide size range. The species of hyenids responsible for the coprolites is not possible to identify based exclusively on the size range, and the importance of a multidisciplinary study of the latrines and the coprolites is discussed as a source of palaeoecological and taphonomic information. In this study, coprolites are analysed, compared, and attributed to a hyenid, identified as Crocuta crocuta in the case of TD6.1, but without specific attribution in the case of la Mina., The research at Barranc de la Boella has been carried out with the financial support of the Spanish Ministerio de Economía y Competitividad projects: CGL2012-38358; CGL2012-38434-C03-03; CGL2012-36682, and HAR2012-32548. The Generalitat de Catalunya, AGAUR agency, provided projects 2014SGR-901, 2014PBR-899 and the framework of CERCA Programme/Generalitat de Catalunya. Financial support for Barranc de la Boella fieldwork and archaeological excavations has been provided by the Ajuntament de La Canonja the Diputació de Tarragona and the Departament de Cultura (Servei d'Arqueologia i Paleontologia) (2014/100574) of the Generalitat de Catalunya. The research at Sierra de Atapuerca has been carried out with the financial support of the Spanish Ministerio de Economía y Competitividad and the Fondo Europeo de Desarrollo Regional (FEDER): CGL2015-65387-C3-1-P (MINECO/FEDER).

El Harhoura 2 cave (Temara, Morocco) has yielded Aterian and Iberomaurusian lithic arti-facts 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 car-nivores, 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.

Four new saber-tooth cat ( Homotherium ) sites in Germany with new dental and postcranial bone material are different in their taphonomic context: 1. The Archaeological Middle Palaeolithic (MIS 9e-Interglacial) Schoningen Lake site with remains of a cub carcass, 2. The Middle Palaeolithic (MIS 5e-9) Archaeological/cave bear den site of Balve Cave yielding a lower canine tooth of an older individual, 3. The Zoolithen Cave (MIS 3–9) cave bear/hyena den with one distal half humerus of an adult, 4. The Ketsch open air Rhine River terrace site which has provided another distal humerus of an adult saber-tooth cat. Whereas only the Schoningen site is precisely dated as Holsteinian Interglacial (approx. 330.000–315.000 BP), all other material seems to come from the same Middle Pleistocene warm period, or few younger Saalian interstadials (MIS 7a, e) deposits, and did not extend over the last MIS 7 glacial into the Late Pleistocene. Homotherium as hyena-like slow moving cat seems to have disappeared within the Saalian due to competition with other scavengers like Ice Age spotted and brown hyenas ( Crocuta crocuta praespelaea/ultima and Pachycrocuta brunnea mosbachensis ). The juvenile saber-tooth cat cub from Schoningen might be in archaeological context or represent only a carnivore kill. At the Zoolithen Cave, the single bone must have been imported into a hyena prey bone assemblage. The situation is possibly similar at the two other sites Ketsch and Balve Cave. The formerly described Schoningen “saber-tooth cat” humerus is revised, such as other opposite as lion humeri described material from different European sites. The presence of the well-developed supracondylar ridge distinguishes Homotherium well from Middle/Late Pleistocene lions Panthera leo (e.g. spelaea, fossilis ). The Schoningen lion humrus has been chew-cut first most probably by a stripped hyena whose cutting scissor teeth produced a diagonal bite cut and P 4 /M 1 impact marks around the trochlea. 1–2 mm small, mostly triangular-oval bite marks on the lion humerus shaft compacta results from a second scavenger and not from “Neanderthal tool use”. Those bite mark sizes are produced mainly of the upper molar teeth of a the red wolf Cuon alpinus subsp. (or small fox Vulpes praecorsac ), which were present in the region within the Holsteinian/Saalian.

The striped hyena (Hyaena hyaena Linnaeus, 1758) is one of the large carnivores of forest and grassland ecosystems across Africa and Asia and is currently threatened throughout its range. Hyena habitats are shrinking, due to the increase of human-dominated landscapes. The aim of this study was to identify hyena habitat use as indicated by visitation frequency in the human-dominated landscape of lowland Nepal. We followed a camera trap protocol for the estimation of hyena presence and visitation frequency. We used a generalized additive modelling approach with zero-inflated Poisson distribution to evaluate the effects of environmental variables on capture frequency. Our results showed that the two most important and significant predictors of hyena frequency data were canopy volume and maxD (the maximum among the shortest distances to water sources, agriculture land and villages). The highest hyena frequencies were in sites with low canopy volume, i.e. riverbeds and grasslands, whereas lower frequencies were observed in dense forests, i.e. acacia and riverine forests. Additionally, hyena frequency increased with increasing maxD up to 5–6 km and then dropped sharply at larger maxD values. Overall, our study suggests that hyenas prefer open landscapes along rivers and human settlements, where the following combination of conditions is met: adequate cover, access to food and water. The preservation of natural open landscapes close to human settlements and rugged hilly areas seems crucial for the long-term conservation of hyenas.

During the Middle Stone Age (MSA), hominins occupied a number of cave sites in southern Africa. Faunal remains of large mammals are commonly found at these MSA sites. Additionally, a number of faunal accumulations have been excavated that were collected by carnivores such as leopards and hyenas. Here we use the ratios of animal remains from MSA sites in the Eastern and Western Cape provinces of South Africa to investigate the contribution of specimens accumulated by carnivores versus hominins. In particular, we calculate the carnivore – ungulate, leopard, leopard – hyena, Raphicerus and hyrax indices. We also calculate a Modified Raphicerus index. Our results suggest that two of these ratios, namely the carnivore – ungulate and leopard indices are useful to demonstrate differences between carnivore and anthropogenic accumulations. High values of the carnivore – ungulate ratio indicate samples accumulated by hyenas, whereas those assemblages with high values of the leopard index, point to leopards as the main accumulator of faunal remains. Conversely, the carnivore – ungulate and leopard ratio values are low for samples accumulated mostly by humans.

Local attitudes towards wildlife species are key for the coexistence between humans and wildlife. To assess how economic, social, and psychological factors affect human perception of wildlife species we conducted questionnaire surveys in two ecologically and culturally distinct rural areas of Northern Tanzania (Mbulu Plateau highlands and Rift Valley lowlands). Using responses of 356 individuals we determined local preferences for population sizes of 31 wildlife species. For five of these species or species groups (rodents, hyena, birds of prey, African elephant, jackal), more than 30% of participants desired a population decline. To investigate correlates for desired population reductions of these species, we ran species-specific (five listed species and African lion) and study area-specific generalized linear mixed models that accounted for spatial autocorrelation. Using these models we assessed relationships between the likelihood of respondents desiring a population decline and six hypothesized explanatory variables: gender; age; ethnicity, and wealth of participant; perceived frequency of negative interactions with; and fear of each species. In both the highland and lowland and for all species, participants that perceived higher instances of negative interactions with wildlife were more likely to prefer smaller future populations, but relationships between perceived frequency of negative interactions and attitudes were often non-linear. People who feared a species were also more likely to desire a population decline. Other variables (gender, age, ethnicity, wealth) showed species- and area-level variations, and we observed substantial spatial variation in expressed attitudes towards species. Thus, negative attitudes towards wildlife appear to be mainly associated with past (negative) experiences and fear, but not necessarily with associated costs or socio-demographic variables. To ensure coexistence between wildlife and humans, we suggest (1) wildlife damage prevention through technical measures and (2) educational initiatives to increase positive attitudes towards wildlife.

The study aims to develop a model for quantifying competitive relations within carnivore guilds. Previous methods were able to calculate competition or niche overlap, but were not applicable to the entire guild. They only permit a pairwise view among the guild members and neither the amount of the resource nor the role of the resource in the diet of each guild member is considered. The model we introduce is based on the theory of competition-free carrying capacity (CFCC). This CFCC value represents the maximum population size under the assumption that the guild member would exploit its essential resources alone in the absence of competitors. Because the actual population size is known, the percentage to which the guild member exploits its CFCC can be calculated. The loss of CFCC to other guild members is called competition effect. The effects can be calculated by a set of equations and the characterization of each guild member by its actual population size, prey mass spectrum, and daily meat intake. The model is applied to the large carnivore guilds in the Serengeti (Tanzania) and the Kruger National Park (South Africa) to show how a guild of large carnivores is structured. The guild structures of the Serengeti and the Kruger National Park are different with respect to the roles the guild members play. The spotted hyena ( Crocuta crocuta ) plays the dominant role by exploiting most of its CFCC and having strongest competition effects to the other guild members in the Serengeti. In the Kruger guild, this position is occupied by the lion ( Panthera leo ).

Visceral pentastomiasis (porocephalosis) caused by Armillifer armillatus larvae was incidentally diagnosed in a female striped hyena (Hyaena hyaena) of unknown age which died unexpectedly in 2013. The hyena had been imported from Tanzania 8years earlier and have been since then in a zoo in Chiang Mai, northern Thailand. Pathological examination revealed visceral nymph migrans of pentastomes throughout the intestine, liver, diaphragm, omentum and mesentery, spleen, kidneys, and urinary bladder. Polymerase chain reaction and sequencing that targeted the pentastomid-specific 18S rRNA gene determined 100% identity with reference sequence for A. armillatus, suggesting that its ova can infect the hyena to serve as an intermediate host for the parasite. Further studies to identify the source of infection, its risk factors, and host range for A. armillatus are important to determine its zoonotic potential and to better prevent and manage the disease to protect animal and human health.

Archaeological interpretations of the agent of bone breakage are important for reconstructing site formation. Impact flakes, or bone flakes created during bone breakage that exhibit features analogous to flakes produced during lithic reduction, are often used as evidence of hominin marrow acquisition. However, it has long been acknowledged that carnivores, mainly hyenas, also create flakes with these features, but the body of literature on these flakes is virtually non-existent. Here, we present and analyze impact flakes from hammerstone- and hyena-created assemblages and compare them to published research on hammerstone-generated flakes in experimental and archaeological contexts. Impact flake frequencies in the experimental hyena assemblage occur at half the rate as found in the hammerstone created assemblages. However, hyena created flakes do display typical impact flake features at similar rates to those found in the hammerstone-created assemblage. Finally, we compare some frequencies of impact flakes found in other experiments and at archaeological sites. Due to equifinality, impact flakes can only be used as supporting evidence rather than central evidence for understanding hominin marrow access, especially when multiple agents of bone accumulation are present.

Serum samples from three species of living terrestrial top predators were analysed for six groups of persistent organic pollutants (POPs), such as polychlorinated biphenyls (PCBs), dichlorodiphenyltrichloroethane and its metabolites (DDTs), chlordane compounds (CHLs), hexachlorocyclohexane isomers (HCHs), hexachlorobenzene (HCB) and polybrominated diphenyl ethers (PBDEs). The study included three carnivore species: lion (Panthera leo), hyena (Hyena brunnea) and cheetah (Acinonyx jubatus). All samples were collected from healthy living animals between 2004 and 2005. Most of the samples (wild lions (n = 50) and hyenas (n = 11)) were collected from various locations within the Kruger National Park and Addo National Park (South Africa), while captive lions (n = 6) and cheetahs (n = 3) were collected from the Antwerp Zoo (Belgium). In general, relatively low levels of POPs were found in the studied species, varying widely within species and locations. Median concentrations of POPs were higher in captive lions (PCBs: 505 pg/mL; DDTs: 270 pg/mL; HCHs: 72 pg/mL; HCB: 34 pg/mL; CHLs: 24 pg/mL; PBDEs: 8 pg/mL) compared to wild lions (DDTs: 274 pg/mL; HCHs: 44 pg/mL; CHLs: 7.9 pg/mL; PCBs: 2.1 pg/mL; HCB: < LOQ; PBDEs: < LOQ). In the wild animals, POPs accumulated in the following order: DDTs > HCHs > CHLs > HCB > PCBs > PBDEs, while in the captive animals, the order was: PCBs > DDTs > CHLs > HCHs > HCB > PBDEs, suggesting differences in the diet of these animals. Furthermore, wild hyena contained significantly higher (p < 0.05) median levels of POPs compared to wild lions, possibly reflecting differences in metabolic capacity and/or feeding habits, together with an uneven distribution of POPs in the area where the animals lived. No previous data are available to compare for similar terrestrial top predators, such as lion, hyena and cheetah. To our knowledge, this is first study on POPs in these three species. The low POP levels found in this study were several orders of magnitude lower than those for other carnivore species, such as polar bears, grizzly bears, brown bears and wolves worldwide. The present study has revealed the need for expanding research and monitoring on occurrence, levels and disposition of POPs in the top predators of the terrestrial environment.

Hyenas are effective hunters and will consider humans as potential prey if the need and opportunity arise. This study describes the circumstances of hyena attacks, the patterns of injuries sustained, and reconstruction in a resource-poor setting. As part of a charitable surgical mission to Ethiopia in 2012, 45 patients with facial deformities were reviewed, of whom four were victims of hyena attacks. A semi-structured interview was performed to ascertain the circumstances of the attack and the subsequent consequences. The age of the victims at the time of attack varied from 5 to 50 years. The attacks occurred when the victims were alone and vulnerable and took place in outdoor open spaces, during the evening or at night. The initial lunge was made to the facial area; if the jaws closed on the facial bones they were crushed, but in all cases the soft tissues were grasped and torn from the underlying bone. Reconstruction was dictated by the extent of soft tissue loss but could normally be obtained by use of local or regional flaps. Hyenas have been shown to attack humans in a predictable way and cause injuries that typically involve the soft tissues of the face.

From MIS 5 to 3, Western Europe has seen important climate changes that shaped the environments inhabited by past human societies. As a consequence, climate is often considered as a key factor in explaining the cultural and biological events of the Middle and Early Upper Paleolithic (e.g. changes in lithic technology and mobility strategies during the Mousterian, Neanderthal extinction, emergence of “modern” behavior, onset of the Upper Paleolithic). However, the exact impact of Upper Pleistocene global climatic changes on terrestrial ecosystems is still poorly documented. Remarkably, large predators can be affected by changes in herbivore communities because they result in fluctuations of the ecosystem's carrying capacity. Paleoenvironmental reconstructions that include data on faunal dynamics are still rare, so that questions concerning human–environment interactions cannot be properly discussed. Building on a review of modern analogs for ungulate biomass, this paper tries to reconstruct changes in carrying capacity of terrestrial ecosystems in southwestern France. In order to assess the importance of these biomass fluctuations for large predators, cave hyena paleodemographic shifts are estimated through Bayesian analyses of the chronological distribution of hyena den assemblages. Comparison between these two proxies (ungulate biomass modern analogs and cave hyena paleodemography) allows for an adequate discussion of the impact of ungulate biomass fluctuations on large predators. Results show that fluctuations in ungulate biomass are noticeable, with an increase at the end of the Mousterian and a significant drop at the beginning of the Aurignacian. These changes likely had a pronounced impact on large predator populations, as interesting parallels can be identified between the demographic history of hyenas and reconstructed fluctuations in carrying capacity. These results bring new insights on the role played by MIS 3 environmental changes in the demise of the last Neanderthals and the emergence of the Upper Paleolithic in southwestern France.

Cova del Gegant has been subject of several publications, focused on the Neanderthal remains identified, stratigraphical context and small vertebrates. However, a systematic taphonomical research of the macro mammal assemblage has not been conducted yet. Here we present the archaeozoological analysis of the material recovered in two sectors (SG and J) adjacent to Gallery 1 (GL-1), which sediments yielded the Neanderthal mandible. Numerous evidences of carnivore involvement are observed in the assemblage, such as tooth marks, a significant quantity of hyena remains, coprolites and deciduous teeth. Their occurrences together with the topography of the passages suggest that humans were not the main agent responsible for the faunal accumulation. These passages seem to have acted as storage and breeding spaces for hyenas during the Upper Pleistocene.

This review/research paper summarizes data on development of the external genitalia of the spotted hyena, a fascinating mammal noted for extreme masculinization of the female external genitalia. The female spotted hyena is the only extant mammal that mates and gives birth through a pendulous penis-like clitoris. Our studies indicate that early formation of the phallus in both males and females is independent of androgens; indeed the phallus forms before the fetal testes or ovaries are capable of synthesizing androgens. Likewise, pre- and postnatal growth in length of the penis and clitoris is minimally affected by “androgen status”. Nonetheless, several internal morphologies, as well as external surface features of the phallus, are androgen-dependent and thus account for dimorphism between the penis and clitoris. Finally, estrogens play a critical role in penile and clitoral development, specifying the position of the urethral orifice, determining elasticity of the urethral meatus, and facilitating epithelial-epithelial fusion events required for proper formation of the distal urethra/urogenital sinus and prepuce. Accordingly, prenatal inhibition of estrogen synthesis via administration of letrozole (an aromatase inhibitor) leads to malformations of the glans as well as the prepuce (hypospadias). The effects of prenatal androgens, anti-androgens and impaired estrogen synthesis correlated with the tissue expression of androgen and estrogen receptors.

European leopard sites in Europe demonstrate Early/Middle Pleistocene out of Africa lowland, and Late Pleistocene Asian alpine migrations being driven by climatic changes. Four different European Pleistocene subspecies are known. The final European Late Pleistocene “Ice Age leopard” Panthera pardus spelaea ( Bachler, 1936 ) is validated taxonomically. The skull shows heavy signs of sexual dimorphism with closest cranial characters to the Caucasian Panthera pardus ciscaucasica (Persian leopard). Late Pleistocene leopards were distributed northernmost, up to S-England with the youngest stratigraphic records by skeletons and cave art in the MIS 2/3 (about 32,000–26,000 BP). The oldest leopard painting left by Late Palaeolithics (Aurignacians/Gravettians) in the Chauvet Cave (S-France) allows the reconstruction of the Ice Age leopard fur spot pattern being close to the snow or Caucasian leopards. The last Ice Age glacial leopard habitat was the mountain/alpine boreal forest (not mammoth steppe lowland), where those hunted even larger prey such as alpine game (Ibex, Chamois). Into some lairs, those imported their prey by short-term cave dwelling (e.g. Baumann's Cave, Harz Mountains, Germany). Only Eurasian Ice Age leopards specialized, similar as other Late Pleistocene large felids (steppe lions), on cave bear predation/scavenging partly very deep in caves. In Vjetrenica Cave (Dinarid Mountains, Bosnia Herzegovina), four adult leopards (two males/two females) of the MIS 3 were found about two km deep from the entrance in a cave bear den, near to one cave bear skeleton, that remained articulated in its nest. Leopards died there, partly being trapped by raising water levels of an active ponor stream, but seem to have been killed possibly either, similar as for lions known, in battles with cave bears in several cave bear den sites of Europe (e.g. Baumann's Cave, Wildkirchli Cave, Vjetrenica Cave). At other large cave sites, with overlap of hyena, wolf and dhole dens at the cave entrances, leopard bones with bite damages indicate their remains to have been imported and consumed by predators in alpine regions due to reduced prey availability. The best models for the competition/taphonomy of large predators – felids, hyenids, canids – within large cave bear dens of Europe is represented in combination of the Zoolithen Cave and Vjetrenica Cave taphonomy.

The Cueva del Camino site (Pinilla del Valle, Madrid) represents the most complete MIS 5 record from the Iberian Peninsula (away from the Mediterranean margin), including a large accumulation of fossilized remains of small and large vertebrates and two human teeth. The presence of carnivores (mainly hyenas) and humans suggests that the site should be interpreted as a spotted hyena den, a human occupation, or both. During an earlier phase of excavation undertaken during the 1980s, an anthropic origin was suggested for the accumulation at the site. However, research was resumed in 2002, leading to an increase in the number of vertebrate remains recovered, as well as the recognition of new vertebrate species. These have now been incorporated into the site’s list of fauna. In addition, new palaeobotanical, geochronological and stratigraphic data have been recorded and analysed, and the human teeth identified as being of Neanderthal origin. Floristic data (pollen and charcoal remains) obtained for the north sector of this site indicate an open landscape with Pinus sylvestris-nigra as the main arboreal taxon. The available evidence suggests this accumulation to be the result of spotted hyena activity during a warm phase of Marine Isotope Stage 5 (MIS 5) in an environment in which fallow deer was the most abundant herbivore.

Crocuta crocuta spelaea ( Goldfuss, 1823 ) remains (NISP = 50) are present in the early to middle Upper Pleistocene Emscher River terrace open air den site along the Rhine–Herne Canal near Bottrop (Westphalia, NW Germany). The population includes bones from cubs and bones with pathological features from old animals but is predominantly made up of adult hyenas (NISP = 3820) found within the glacial mammoth steppe fauna of the Munsterland Bay Lowlands. A larger quantity of woolly rhinoceros ( Coelodonta antiquitatis ) bones (NISP = 1601) are present than in other bone-rich open air hyena den sites, of which about 67% are damaged. This damage can be shown to have resulted mainly from hyena activities, and shows repetitions of the same three stages of destruction on the massive woolly rhinoceros bones. Those bone shafts that the hyenas were unable to break were left with irregular jagged margins. The taphonomy of bones from the Bottrop open air site indicates that the rhinoceros body parts had only been transported over short distances, in contrast to those from the Perick Caves hyena den where longer transport distances resulted in a higher proportion of limbs compared to other body parts. The large woolly rhinoceros was an important second megafauna prey for hyenas in the lowland areas (after the woolly mammoth), in contrast to the nearby Sauerland Karst mountain areas that are rich in hyena dens and their associated bone assemblages, although the higher proportions of rhinoceros bones present is also to some extent a result of the incomplete destruction of their massive bones. The proportions of woolly mammoth and woolly rhinoceros remains in the hyena prey bone assemblages gradually decrease from the lowlands towards the nearby Sauerland Karst mountain regions. In these mountainous boreal forest regions cave bears were instead the main food source for hyenas, as result of the scarcity or absence of mammoth steppe fauna. The large bone accumulations along the Emscher River terraces near Bottrop can mostly be attributed to the activities of hyenas, which were responsible for the repeated incomplete state of bone preservation. The bone accumulations are predominantly of leg bones and include a relatively high proportion of hyena bones which, together with the faunal composition and the size of the accumulations, indicate a mixed long-term use of the terraces along the Emscher River as an extensive communal hyena den and prey storage site, and also probably somewhere in the area, as a birth den. Some of the bone material may also have been accumulated by natural processes, or even by Neanderthals.

Associated and partially articulated fossil remains of the endemic reduced size elephant Palaeoloxodon mnaidriensis have been recently discovered at San Teodoro Cave, a large Late Pleistocene hyena den in Sicily. The skeletal elements belong to both cranial and post-cranial portions, and are represented by a semi-complete mandible, a tusk fragment, a cervical vertebra, three thoracic vertebrae, a rib fragment, a scapula, a distal epiphysis of radius, a pyramidal bone, a III metacarpal bone, a coxal bone fragment, a femur shaft, two symmetrical pairs of tibias and fibulas, a patella and an astragalus. The bones were recovered mixed with remains belonging to other taxa, numerous hyena coprolites and with a juvenile elephant mandible in the perinatal stage. On the basis of anatomical representation, spatial distribution and development stages, the partially articulated elephant bones from the San Teodoro Cave belong to a single adult individual, perhaps a female, which probably entered the cave before its death, and was afterwards scavenged and disarticulated by hyenas. This is the first finding of a partially articulated elephant skeleton from a fossil hyena den, and may attest to the intermittent use of the cave by hyenas and elephants.

The use of enzyme immunoassays (EIAs) to measure faecal glucocorticoid metabolites (fGCM) is a useful non-invasive technique to monitor adrenocortical activity in vertebrates. The first objective of this study was to validate an ‘in-house’ EIA (cortisol-3-CMO) for the measurement of fGCM concentrations in spotted hyenas. High-performance liquid chromatography (HPLC) was used to characterise fGCM in samples from a captive hyena that received an i.v. injection of [3H] cortisol. All HPLC fractions were analysed with the EIA for the presence and quantities of radiolabelled fGCM. Radiolabelled fGCM consisted of substances with a higher polarity than cortisol and substances of lower polarity that eluted between cortisol and corticosterone. Authentic radiolabelled cortisol was not detected. The EIA measured substantial amounts of immunoreactivity corresponding to the radioactive peaks. It also detected a significant increase in fGCMs after an adrenocorticotropic hormone (ACTH) challenge in two other captive animals and a significant increase in fGCMs in a fourth captive animal after anaesthesia. The second objective was to investigate an age effect on fGCM: we conducted pairwise comparisons of fGCM concentrations in individual free-ranging juvenile spotted hyenas when less than 6 months of age and when between 6 and 24 months of age. We expected juveniles to experience a more unpredictable and therefore more stressful environment when younger than when older. When younger, juveniles had significantly higher fGCM concentrations than when they were older. Our results demonstrate that our assay can be used to assess adrenocortical activity in spotted hyenas.

Hyaenas are relevant members of the Plio-Pleistocene carnivore guilds in Africa and Eurasia. The spotted hyena Crocuta crocuta is an important element of the dispersal events from Africa and Asia to Europe occurred at the Early–Middle Pleistocene transition (about 0.8 Ma). The presence of Crocuta crocuta remains from the Ponte Galeria formation at Casal Selce (Roma, Italy) from sands of the Ponte Galeria Formation referable to the earliest Middle Pleistocene (MIS 18), close to the Brunhes/Matuyama boundary, is one of the earliest occurrence of the genus in Europe.

Late Pleistocene Ice Age Crocuta crocuta spelaea ( Goldfuss, 1823 ) hyenas from the open-air gypsum karst site Westeregeln (Saxony-Anhalt, central Germany) is dated into the early to middle Late Pleistocene. Hyena clans apparently used the karst for food storage and as “commuting den”, where typical high amounts (15% of the NISP) of hyena remains appear, also faecal pellets in concentrations for den marking purposes. Additionally small carnivores Meles, Vulpes and Mustela appear to have used some cavities as dens. Several hundreds of lowland “mammoth steppe fauna” bones (NISP = 572) must have been accumulated primarily by hyenas, and not by Neanderthals at the contemporary hyena/human camp site. Abundant caballoid horse remains of “E. germanicus Nehring, (1884) ” are revised by the holotype and original material to the small E. c. przewalskii horse. Woolly rhinoceros Coelodonta antiquitatis remains are also abundant, and were left in several cases with typical hyena scavenging damages. Rangifer tarandus (11%) is mainly represented by numerous fragments of shed female antlers that were apparently gathered by humans, and antler bases from male animals that were collected and chewed in few cases (only large male antlers) by hyenas. The large quantities of small reindeer antlers must have been the result of collection by humans; their stratigraphic context is unclear but such large quantities most probably resulted from schamanic activities. The hyena site overlaps with a Middle Palaeolithic Neanderthal camp, as well as possibly with a later human Magdalenian site.

Adult external genitalia (ExG) are the endpoints of normal sex differentiation. Detailed morphometric analysis and comparison of adult mouse ExG has revealed 10 homologous features distinguishing the penis and clitoris that define masculine vs. feminine sex differentiation. These features have enabled the construction of a simple metric to evaluate various intersex conditions in mutant or hormonally manipulated mice. This review focuses on the morphology of the adult mouse penis and clitoris through detailed analysis of histologic sections, scanning electron microscopy, and three-dimensional reconstruction. We also present previous results from evaluation of “non-traditional” mammals, such as the spotted hyena and wallaby to demonstrate the complex process of sex differentiation that involves not only androgen-dependent processes, but also estrogen-dependent and hormone-independent mechanisms.

During the last Ice Age of central Europe, cave bears hibernated deep inside their caves and Ice Age spotted hyenas used the cave entrance areas as dens. Steppe lions were also sporadic cave dwellers in order to hunt the herbivorous cave bears or steal prey from hyena dens, or for antagonistic and territorial reasons. Steppe lion skeletons recently discovered in the midst of cave bear skeletons, deep inside the Ursilor Cave (Romania), and steppe lion remains from other European caves, in particular a large population from the Zoolithen Cave in Germany, provide evidence of active predation by lions on cave bears. However, it was not mainly lions that fed on the cave bears: the large quantities of damaged cave bear bones, including incomplete long bones that repeatedly show similar types of damage, crushed long bones, and damaged skulls, reveal that hyenas were the primary scavengers on cave bear carcasses and were largely responsible for the destruction of their carcasses and bones. Predation and scavenging on cave bears (mainly by these two Ice Age predators but also by wolves and leopards) particularly in mountainous boreal forest environments that were devoid of the mammoth steppe fauna, explains the large quantities of fragmented cave bear bones that have been found all over Eurasia. The stress caused by these carnivores, deprived of their specialized ecological niche, may have led the herbivorous cave bears to hibernate as deeply as possible within their hibernation caves, in an attempt to protect themselves against predation. The felid predators were sometimes killed inside the caves, probably in conflicts with adult cave bears or even with hyenas, and remained as complete or only partly disarticulated carcasses.

A detailed taphonomic analysis of the large mammal assemblage from 1998 to 2006 excavations at San Teodoro Cave is presented, taking into account the stratigraphic context of the deposits. Three not strictly contemporary fossiliferous levels having different lithological features have been detected, here named B-I, B-II, and B-III. Fossil remains are prevalently accumulated in B-I and B-II. The three levels are characterized by evidence of Crocuta crocuta spelaea occupation, represented by their skeletal remains, coprolites, and distinctive damages on the bones, similar to fossil and modern spotted hyena dens from Europe and Africa. A differential distribution of coprolites and small digested bones, probably due to different humidity conditions, has been recognized in B-I and B-II, and can be related to different topographic locations within the cave or to different climate conditions during the sedimentation phases. The very low density of fossil remains in B-III, which is the oldest level, could indicate an area that was less inhabited by hyenas, probably due to geomorphological conditions. Taphonomic comparison of the three fossiliferous levels of the San Teodoro Cave deposits points to a long-term, perhaps cyclic, occupation of the cave by hyenas and confirms the cave as one of the most important Pleistocene hyena dens in Europe.

The giant hyena Pachycrocuta brevirostris was the largest bone-cracking carnivore that ever existed. With the mass of a lioness, it had massive limbs with shortened distal bones and a heavy, powerfully built mandible with robust, well-developed premolars. All these features reflect its adaptation for dismembering ungulate carcasses, transporting large pieces of them without dragging to the denning site and fracturing bones. This paper estimates the relative contribution of hunting and scavenging to the diet of this extinct hyena, using a combined biomechanical and taphonomic approach. Analysis of the bone-cracking behavior of P. brevirostris was based on the abundance of skeletal elements in the large mammals assemblage from Venta Micena (Guadix-Baza basin, southeast Spain), a locality currently interpreted as an early Pleistocene hyena den. The distribution of major limb bones of ungulates among complete elements, isolated epiphyses and diaphyses were analyzed using contingency tables and correspondence analysis. Results obtained showed that the bones with greater marrow contents (femur, humerus and tibia) were preferentially fractured by the hyenas, while those others with less nutritional value (radius and metapodials) were better represented as complete elements in the assemblage. The quantitative analysis of the preservational state of skeletal elements allowed testing specific patterns of bone modification by the giant hyenas, such as a proximodistal sequence of consumption for humerus and tibia, thus revealing the highly specialized bone-cracking behavior of P. brevirostris. Regression equations adjusted with modern carnivores for body size on craniodental and postcranial measurements provide an average estimate of mass of ∼110 kg for the giant hyena. The high moment arms for masseter and temporalis muscles indicate a substantial strength for bone fracturing with the well-developed premolar teeth. Jaw depth provided resistance against dorsoventral loads during bone-cracking activities. However, the moment arm of resistance for an object positioned at the canines reveals a loss of bite strength compared with spotted hyenas and thus less predatory abilities. These results are in agreement with the scavenging niche deduced for P. brevirostris from taphonomic analysis.

Bone remains and a trackway of Pantheraichnus bottropensis nov. ichg. ichnsp. of the Late Pleistocene lion Panthera leo spelaea ( Goldfuss, 1810 ) have been recovered from Bottrop and other open air sites in northern Germany. Some of these bones are from open air hyena den sites. A relative high proportion of lion bones (20%) exhibit bite, chew or nibble marks, or bone crushing and nibbling caused by a large carnivore. Repeated patterns of similar bone damage have been compared to bone remains found at hyena dens in gypsum karst areas and cave sites in northern Germany. Ice Age spotted hyenas have been the main antagonists and the main scavengers on lion carcasses. The remains appear to have been imported often by hyenas into their communal dens, supporting the theory of strong hyena-lion antagonism, similar to the well documented antagonism between modern African lions and spotted hyenas. Most of the lion bones from the open air hyena den at Bottrop are probably a result of such antagonism, as are the rare remains of these carnivores found within large hyena prey bone accumulations along the Pleistocene rivers. The Emscher River terrace also has the largest quantity of hyena remains from open air river terrace sites in northern Germany. Their cub remains, and incomplete chewed prey bones from mammoths and woolly rhinoceroses, typical of hyena activity, underline the character of these sites as cub-raising and communal dens, where their prey was accumulated along the riverbanks in a similar manner to modern African hyenas.

A relatively high amount of hyena Crocuta crocuta spelaea (Goldfuss, 1823) remains at the Palaeolithic archaeological cave site Balve Cave (Sauerland Karst, NW Germany) and many chewed prey bones indicate periodic repeated use by hyenas. They reared their cubs and imported many prey remains. Most probably, Ice Age spotted hyenas occupied the Balve Cave for only a few years but at several times, which is sufficient time for a high amount of accumulated bones of any kind of mega mammals, of which the best den indicators are bones of the woolly rhinoceros. Comparisons with incomplete Coelodonta antiquitatis and other megafauna bones at seven other hyena den caves in the Sauerland Karst of northwestern Germany which have no human remains or impact show similar bone assemblages and bone preservation. The separation of human and animal prey remain is only possible in some cases, where cutting or hitting marks on bones can be observed. Generally it seems that humans smashed large mammoth bones into pieces for the use as burning material, resulting in “bone coal” layers. In the Late Pleistocene, fewer cave bears, then hyenas and finally mainly Neanderthalians (Micoqien/Mousterien) used the cave. In the Early to Early Late Weichselian, the first modern humans of the Aurignacien settled at the cave entrance. After the maximum cold period, this site was reused by Late Magdalenians when hyenas were already extinct in northern Europe.

Risk is a fundamental driver of prey behaviour, but the importance of risk in the decision-making processes of top predators is poorly understood. We investigated the functions of vigilance behaviour in wild spotted hyaenas. These large carnivores have few predators but face frequent threats of attack and food theft by conspecifics. We made repeated observations of vigilance behaviour by hyaenas when they were resting, feeding and nursing, and examined variation due to age, group size, social rank, and factors related to predation risk. We used these observations to test whether vigilance functions in hyaenas to detect prey, mates, conspecific threats or interspecific threats, and whether functions differ between juveniles and adults. We found that adults were more vigilant than juveniles in all contexts. Vigilance by adults was greatest when nursing, but nursing females reduced their vigilance as cubs matured. Among both age groups, vigilance decreased with increasing group size, and hyaenas were less vigilant when feeding on high-quality food than when feeding on low-quality food. Social rank did not affect vigilance in either age group. Individual adults, but not juveniles, showed consistent patterns of vigilance within, but not between, behavioural contexts. Surprisingly, our results indicate that the primary function of vigilance in hyaenas is to detect interspecific threats. Furthermore, vigilance was context specific, and adult hyaenas showed individual variation in vigilance within behavioural contexts. These findings illustrate the relative importance of particular risks in shaping vigilance behaviour of a top carnivore.

9 páginas, 5 figuras., This article examines the role of the Late Pleistocene climatic changes in the disappearance of the European populations of spotted hyenas. A species distribution model was built using both current and past environmental requirements of the species. Model projections were made with climatic scenarios provided by the GENESIS 2.0 General Circulation Model (126 ka, 42 ka, 30 ka and 21 ka). Those projections indicate (1) that during the Late Pleistocene warm scenarios spotted hyenas should have been widespread in Europe, and (2) that during the last glacial maximum their potential climatically suitable geographic distribution diminished in size. The decrease in the potential climatic distribution was strictly restricted to Northern Europe. Climatic conditions in Southern Europe during the Late Pleistocene remained within the spotted hyena climatic tolerance. Hence, climate changes could have directly affected the Northern distribution of the species during the last glaciations. However, climate change alone is not sufficient to have caused the disappearance of the spotted hyena populations in Southern Europe. That is, other factors, such as prey abundance or human ecological impacts, in addition to climatic change, are needed to completely account for extinction of the European spotted hyena., The first author (S. Varela) was supported by a doctoral fellowship from the Fundación Atapuerca. J. Rodríguez was supported by the Spanish MEC project (CGL2009-12703-C03-01) and by the JCyLGrupos de Excelencia program (GR249/2008).

Late Pleistocene spotted hyenas hunted Przewalski horses in spring to early summer, as documented by foetal horse skeleton remains of Equus ferus cf. przewalskii Poljakoff 1881, found between 3569 megamammal bones in the hyena den site Srbsko Chlum-Komin of the Bohemian Karst (Czech Republic). The main prey of hyenas Crocuta crocuta spelaea (Goldfuss 1823) was this small horse, well distributed in Central Europe. 51% (NISP = 1.800) of the megafauna in the Srbsko Chlum-Komin Cave are horse bones, consisting of mainly complete and in a few cases cracked and chewed dominantly leg and cranial remains. Carcass body parts must have been imported into the hyena prey storage and cub-raising den cave. Similar high amounts of horse remains and taphonomic situations are reported from other Late Pleistocene hyena den caves in Central Europe, such as Rochelot Cave (France). Den sites indicate an active selected hunt of mainly adult horses. Hyenas left partly articulated legs or complete distal bones and many isolated teeth at their dens, the latter resulting from consumption of the thin-walled horse skulls. In periods of megafauna species abundance, hyenas used mainly the meat leaving only a few chewed or cracked horse bones (Srbsko Cave or Rochelot Cave), whereas in poor feeding times or in cub-raising dens, bone consumption was higher resulting in many fragmented and chewed horse bones (Nad Kacakem Cave, Czech Republic). Whereas African spotted hyenas hunt zebras and even modern donkeys successfully, in comparison to modern spotted hyena dens in Amboseli (Kenya) and Syokimau (Botswana), it seems as if the Ice Age spotted hyena Crocuta crocuta spelaea of Europe hunted very similar equine prey, in this case the Przewalski-horse, especially in hilly and mountainous regions, where the largest prey (mammoth and woolly rhino) were rare or absent.

Data of high resolution for reconstructions of archaeological site formation processes can be obtained only by the use of precise excavation methods and comprehensive recovery techniques using fine-mesh water screening, followed by meticulous sorting and quantitative studies of all the small organic components from the screens. These methods are generally used at human occupation sites, but are not often employed at paleontological or mixed human-carnivore sites. At the site of Bois Roche, an Upper Pleistocene hyena den, we have used these meticulous recovery techniques in combination with absolute dating, geoarchaeological observations, taphonomic analyses, studies of microvertebrates and pollen analysis. Correlation of the site environmental data from pollen and microvertebrate assemblages with the vegetation record of a deep-sea core taken in the Bay of Biscay have allowed us to constrain the ages provided by ESR dating of bovid teeth and to reconstruct the physical environment of the site with a degree of confidence rarely achieved by the use of fossil remains from archaeological sites alone. Prior to our excavations, the site was interpreted as containing a mixed assemblage with evidence of hyena activities and human occupation in situ. Quantitative analysis of the small organic components present in large quantities in the sediments (deciduous hyena teeth, coprolite fragments, digested small bone fragments) prove, more than all other lines of evidence, that hyenas were in residence at Bois Roche, thus providing diagnostic criteria for distinguishing between occasional visitors and the main inhabitants of mixed human-carnivore sites. The analytical procedures presented in this paper have a value beyond the significance of the Bois Roche site because they can be applied to other sites.

The late Miocene/early Pliocene ‘E’ Quarry deposits at Langebaanweg, South Africa, have yielded fossil material of at least four hyena species ( Ikelohyaena abronia , Hyaenictitherium namaquensis , Chasmaporthetes australis , Hyaenictis hendeyi ). The co-occurrence of four closely-related hyaenids at Langebaanweg raises interesting questions about niche partitioning that have not been sufficiently addressed. In the only previous systematic attempt to tackle this issue, an analysis of m1 lengths, m1 trigonid lengths and p3 widths found evidence for community-wide character displacement in the first two variables. Here, a geometric morphometric approach is used to compare p2, p3, p4 and m1 (carnassial) crown form to better understand feeding niche partitioning among these four hyena species. While results indicate that all four species are very similar in crown shape, I. abronia and H. namaquensis tend to have slightly shorter, broader cheek teeth than C. australis and H. hendeyi . This suggests that I. abronia and H. namaquensis may have been less carnivorous than C. australis and H. hendeyi . The biggest difference between the species however relates to crown size. The four species exhibit community-wide character displacement with respect to p3 and p2 crown size, but not p4 crown size. While box plots based on crown log centroid size suggest that these hyenas also exhibit community-wide character displacement with respect to m1 crown size, this could not be statistically tested due to small sample size. These results support a previous suggestion that community-wide character displacement was in operation among the ‘E’ Quarry hyenas and probably played a major role in niche partitioning and in reducing inter-specific competition.

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 lupus spelaeus 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.

The faunal assemblages excavated by Mary Leakey in Bed II of Olduvai Gorge, Tanzania, have, like the more well-known Bed I assemblages, traditionally been interpreted as the result of hominid butchering activities in the lake margin and riverine settings of the paleo-Olduvai Basin. A reexamination of all of Leakey's Bed I sites has shown that hominids played little or no role in the formation of all but one of those faunal assemblages, a finding that prompted the reanalysis of the Bed II sites presented here. We expand upon a previous taphonomic study that provided systematic data for HWK East Levels 1-2, MNK Main, and BK. In addition to these assemblages, we provide data on HWK East Levels 3-5, FC West, TK, and SHK. Our data contradict previous interpretations of MNK Main as a hominid accumulation but uphold the contention that BK represents a primarily hominid accumulation reflecting early access to carcasses. The small and poorly preserved assemblages from FC West and TK are difficult to link unambiguously to either hominids or carnivores. Site MNK Main and HWK East Levels 3-5 appear to be death arenas where carcasses accumulated via natural deaths and/or serial predation. Site SHK is severely biased by selective retention and therefore little can be said of its formational history. Nevertheless, no hominid modifications were documented in this assemblage. Comparisons with other Olduvai sites indicate a more conspicuous hyena taphonomic signal during Bed II times than Bed I times, which appears to mirror the changing configuration of the large carnivore guild. These findings also beg the question of what activities were being carried out by hominids with the stone tools discarded at these sites. Although it seems clear that hominids were utilizing stone tools to carry out subsistence activities unrelated to carcass butchery, more excavation and techniques such as phytolith analysis should be employed to explore alternative explanations.

Thirteen strains of Streptococcus equi subsp. ruminatorum from free-ranging spotted hyenas ( Crocuta crocuta ) and plains zebras ( Equus burchelli ) in Tanzania were characterized by biochemical and molecular-biological methods. Although the colony appearance of the S.e. ruminatorum wildlife strains differed from that of the S.e. ruminatorum type strain CECT 5772 T , all biochemical reactions of the wildlife strains were similar to those of the type strain. In addition, all wildlife strains produced hyaluronidase and were capable of hydrolysing arginine, three strains (23%) synthesized acetoin, but only eight strains (62%) produced acid from ribose. rep-PCR indicated that different clones of S.e. ruminatorum were distributed among the hyena and zebra populations in the study area. Identical rep-PCR patterns in hyena and zebra strains suggest that a direct transmission of S.e. ruminatorum between these species may occur. The presence of a M-like protein (SrM) gene was demonstrated in all S.e. ruminatorum strains including the type strain. Sequencing of the M-like protein gene revealed a hypervariable region within the deduced amino acid sequence. Most of the strains clustered with previously described strains based on the hypervariable region of the S.e. zooepidemicus SzP protein. Sequencing also demonstrated that identical SrM protein sequences were shared among S.e. ruminatorum strains from different host species.

Fecal hormone analysis is a useful tool for frequent, non-invasive sampling of free-living animals. Estrogens fluctuate throughout life among reproductive states in female animals, and intensive repetitive sampling can permit accurate assessment of female reproductive condition. This type of repetitive sampling is difficult in large carnivores, including the spotted hyena (Crocuta crocuta). Patterns of estrogen secretion in captive and free-living hyenas are virtually unknown. Here we present validation of an enzyme-immunoassay to measure fecal estrogen (fE) concentrations in wild and captive spotted hyenas. Results from high-performance liquid chromatography indicate that an antibody specific for estradiol exhibits high immunoreactivity with our extracted samples. Fecal extract displacement curves paralleled our estradiol standard curve within the range of 20 - 80% antibody binding. Additionally, animals treated with luteinizing hormone-releasing hormone showed a measurable rise in fE concentrations. Finally, once we controlled for effects of time of day of sample collection from wild hyenas, patterns in fE concentrations resembled those in plasma estradiol, including higher levels of fE in mature than immature females, and higher levels of fE during late than early pregnancy. Together, these results suggest that fE concentrations reflect circulating estrogens in spotted hyenas.