Masculinized otoacoustic emissions in female spotted hyenas (Crocuta crocuta)

In humans and rhesus monkeys, click-evoked otoacoustic emissions (CEOAEs) are stronger in females than in males, and there is considerable circumstantial evidence that this sex difference is attributable to the greater exposure to androgens prenatally in males. Because female spotted hyenas are highly androgenized beginning early in prenatal development, we expected an absence of sexual dimorphism in the CEOAEs of this species. The CEOAEs obtained from 9 male and 7 female spotted hyenas confirmed that expectation. The implication is that the marked androgenization to which female spotted hyenas are exposed masculinizes the cochlear mechanism responsible for CEOAEs. The CEOAEs measured in 3 male and 3 female hyenas that had been treated with anti-androgenic agents during prenatal development were stronger than the CEOAEs of the untreated animals, in accord with the implied inverse relationship between prenatal androgen exposure and the strength of the cochlear mechanisms producing CEOAEs. The CEOAEs of three ovariectomized females and two castrated males were essentially the same as those for the untreated females and males, suggesting that there is little or no activational effect of hormones on CEOAE strength in spotted hyenas. Distortion product OAEs (DPOAEs) also were measured. Those sex differences also were generally small (as they are in humans), and the effects of the anti-androgen agents were inconsistent. Thus, prenatal androgen exposure apparently does affect OAEs, but the effects appear to be greater for the reflection-based cochlear mechanism that underlies CEOAEs than for the nonlinear cochlear mechanism underlying DPOAEs.     

Adrenocorticol responsiveness to immobilization stress in spotted hyenas (Crocuta crocuta)

• 1.1. The adrenocortical responsiveness to an induced Stressor was monitored in free-living spotted hyenas belonging to a number of social and reproductive categories.
• 2.2. No significant differences between sexes, or changes in mean cortisol concentrations during serial sampling within the sexes, could be demonstrated.
• 3.3. The extreme individual variance in temporal cortisol profiles recorded in this study is inexplicable, as it was not related to differences in immobilization procedure, sex, age, reproductive or social category.
• 4.4. Females, which are the dominant sex in this species, generally showed larger percentage increases in cortisol concentrations during serial sampling.
• 5.5. Significant correlations between initial cortisol concentrations, as well as cortisol responsiveness and androgen concentrations, in a number of social and reproductive categories, suggest that these categories do not provide sufficient resolution for the identification of specific dominance-related trends.

     

Sex differences in the play behavior of immature spotted hyenas, Crocuta crocuta

Social, locomotor, and object play were studied in a colony of five male and five female peer-reared spotted hyenas during 12 1-hr tests while the animals were 13-19 months of age. Animals were tested in both same-sex and mixed-sex groups and were stimulated to play by the introduction of fresh straw and sawdust bedding. Each test was videotaped and the frequency of each type of play was determined by a time sampling procedure. Females played more frequently than males, however, the category of play which was elevated depended upon the social context during testing. In same-sex tests the frequency of vigorous social play displayed by females markedly exceeded that by males, but no comparable sex difference appeared in mixed-sex tests. Females engaged in locomotor play more frequently than males in mixed-sex tests, but no comparable sex difference appeared in same-sex tests. No sex difference in object play was observed. Two male and two female hyenas were gonadectomized prior to the initiation of the present sequence of tests. The results suggest that gonadectomy during the prepubertal period does not affect the frequency of play behavior. However, the small sample sizes preclude any conclusive determination of the effects of these gonadectomies on play.     

Male spotted hyenas (Crocuta crocuta) queue for status in social groups dominated by females

A long-term study of immigrant male spotted hyenas (Crocuta crocuta) living in large multimale/multifemale groups (clans) demonstrated that males acquire social status by queuing. Maximumlikelihood estimates of parameters of a stochastic queuingmodel that assessed queuing discipline confirmed that immigrantmales respected the convention that their positions in a queueof typically 15 or more individuals was determined by theirsequence of arrival. Levels of aggression among males were low;males did not attempt to improve their social status throughphysical contests. Size and body mass did not influence malesocial status. The stability of queues was insured by an increasein the rate at which males formed coalitions against othermales as they rose in social status and by coalitions between high-ranked males and dominant females. High-ranked, long-tenuredmales chiefly consorted with ("shadowed") and focused theiraffiliative behavior on females of high reproductive valueand disrupted attempts by subordinate males to associate withthese females. High-ranked males also supported females againstlower-ranked males that harassed them. In contrast, lower-ranked,short-tenured males focused their affiliative behavior on young adult females and rarely shadowed or defended females. Malesthat did not disperse from their natal clan (nondispersers)quickly acquired top rank in the male social hierarchy. Irrespectiveof the social status acquired from their mother when young,nondisperser adult males submitted to all adult females.     

Patterns of growth of the mandibular corpus in spotted hyenas (Crocuta crocuta) and cougars (Puma concolor)

Differences in jaw morphology among adult carnivorans are well established, but the ontogenetic mechanisms by which these differences arise are largely unexplored. Mandibular ontogeny in Crocuta crocuta and Puma concolor is analysed biomechanically using principles of beam theory. In each species, the development of cross-sectional properties of the mandibular corpus associated with rigidity under loading follows a biphasic pattern of growth. In early postnatal growth, deposition of cortical bone appears to be constrained by the overall weaker tissue with which juvenile skeletons are constructed and by the need to volumelrically accommodate the developing teeth within their bony crypts. Thus, this stage of growth is characterized by a net periosteal deposition of bone and a swelling of the medullary cavity. In late postnatal growth, the constraints on endosteal deposition of bone are relieved as the permanent teeth erupt; thus, cortical thicknesses increase sharply by periosteal expansion as well as medullary contraction. Finally, it is noted that basic differences in jaw construction between Crocuta and Puma appear to develop prenatally as they are largely in place at birth. Hence, postnatal development enhances, but does not soley contribute to, the biomechanical differences in the jaws of these species.     

Coprologic survey of parasites of spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve,Kenya

Seventy fecal samples from spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve, Kenya were examined for parasite eggs and oocysts using sugar flotation. A total of nine parasite genera were identified, and all samples were positive for at least one parasite species. Most individuals were infected with Ancylostoma sp. and Spirometra sp., and these species had the highest median intensity of infection. Other parasites identified include Isospora sp., Taeniidae, Spirurida, Toxocara sp., Mesocestoides sp., Dipylidium sp., and Trichuris sp.     

Antibodies to canine and feline viruses in spotted hyenas (Crocuta crocuta) in the Masai Mara National Reserve

Spotted hyenas (Crocuta crocuta) are abundant predators in the Serengeti ecosystem and interact with other species of wild carnivores and domestic animals in ways that could encourage disease transmission. Hyenas also have a unique hierarchical social system that might affect the flow of pathogens. Antibodies to canine distemper virus (CDV), feline immunodeficiency virus (FIV), feline panleukopenia virus/canine parvovirus (FPLV/CPV), feline coronavirus/ feline infectious peritonitis virus (FECV/IPV), feline calicivirus (FCV), and feline herpesvirus 1 (FHV1) have been detected in other Serengeti predators, indicating that these viruses are present in the ecosystem. The purpose of this study was to determine whether spotted hyenas also had been infected with these viruses and to assess risk factors for infection. Serum samples were collected between 1993 and 2001 from 119 animals in a single clan for which behavioral data on social structure were available and from 121 hyenas ill several other clans. All animals resided in the Masai Mara National Reserve. Antibodies to CDV, FIV, FPLV/CPV, FECV/FIPV, FCV, and FHV1 were present in 47%, 3.5%, 81%, 36%, 72%, and 0.5% of study hyenas, respectively. Antibody prevalence was greater in adults for FIV and FECV/FIPV, and being a female of high social rank was a risk factor for FIV. Hyenas near human habitation appeared to be at lower risk to have CDV, FIV, and FECV/FIPV antibodies, whereas being near human habitation increased the risk for FPLV/CPV antibodies. Canine (distemper virus and FECV/FIPV antibody prevalence varied considerably over time, whereas FIV, FPLV/CPV, and FCV had a stable, apparently endemic temporal pattern. These results indicate that hyenas might play a role in the ecology of these viruses in the Serengeti ecosystem. The effect of these viruses on hyena health should be further investigated. The lower prevalence of CDV antibody-positive hyenas near human habitation suggests that reservoirs for CDV other than domestic dogs are present in the Serengeti ecosystem.     

Of arcs and vaults: the biomechanics of bone‐cracking in spotted hyenas (Crocuta crocuta)

The ability to break open large bones has evolved independently in only three groups of carnivorous mammals, all of which have robust teeth, vaulted foreheads, and pronounced sagittal crests. One unusual skull feature, present in bone‐cracking members of the family Hyaenidae, is a caudally elongated frontal sinus, hypothesized to function in resistance to bending and stress dissipation during bone‐cracking. In the present study, we used finite element (FE) analysis to examine patterns of stress distribution in the spotted hyena (Crocuta crocuta) skull during unilateral biting, and inquire about the functional role of the fronto‐parietal sinus in stress dissipation. We constructed and compared three FE models: (1) a ‘normal’ model of an adult Crocuta skull; (2) a model in which the caudal portion of the fronto‐parietal sinus was filled with bone; and (3) a model in which we flattened the sagittal crest to resemble the plate‐like crests of other mammals. During biting, an arc of stress extends from the bite point up through the vaulted forehead and along the sagittal crest. Our results suggest that pneumatization of the hyena's skull both enhances its ability to resist bending and, together with the vaulted forehead, plays a critical role in evenly dissipating stress away from the facial region. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 246–255.     

Social intelligence in the spotted hyena (Crocuta crocuta)

If the large brains and great intelligence characteristic of primates were favoured by selection pressures associated with life in complex societies, then cognitive abilities and nervous systems with primate-like attributes should have evolved convergently in non-primate mammals living in large, elaborate societies in which social dexterity enhances individual fitness. The societies of spotted hyenas are remarkably like those of cercopithecine primates with respect to size, structure and patterns of competition and cooperation. These similarities set an ideal stage for comparative analysis of social intelligence and nervous system organization. As in cercopithecine primates, spotted hyenas use multiple sensory modalities to recognize their kin and other conspecifics as individuals, they recognize third-party kin and rank relationships among their clan mates, and they use this knowledge adaptively during social decision making. However, hyenas appear to rely more intensively than primates on social facilitation and simple rules of thumb in social decision making. No evidence to date suggests that hyenas are capable of true imitation. Finally, it appears that the gross anatomy of the brain in spotted hyenas might resemble that in primates with respect to expansion of frontal cortex, presumed to be involved in the mediation of social behaviour.     

Pleistocene Chinese cave hyenas and the recent Eurasian history of the spotted hyena,Crocuta crocuta

The living hyena species (spotted, brown, striped and aardwolf) are remnants of a formerly diverse group of more than 80 fossil species, which peaked in diversity in the Late Miocene (about 7–8 Ma). The fossil history indicates an African origin, and morphological and ancient DNA data have confirmed that living spotted hyenas (Crocuta crocuta) of Africa were closely related to extinct Late Pleistocene cave hyenas from Europe and Asia. The current model used to explain the origins of Eurasian cave hyena populations invokes multiple migrations out of Africa between 3.5–0.35 Ma. We used mitochondrial DNA sequences from radiocarbon‐dated Chinese Pleistocene hyena specimens to examine the origin of Asian populations, and temporally calibrate the evolutionary history of spotted hyenas. Our results support a far more recent evolutionary timescale (430–163 kya) and suggest that extinct and living spotted hyena populations originated from a widespread Eurasian population in the Late Pleistocene, which was only subsequently restricted to Africa. We developed statistical tests of the contrasting population models and their fit to the fossil record. Coalescent simulations and Bayes Factor analysis support the new radiocarbon‐calibrated timescale and Eurasian origins model. The new Eurasian biogeographic scenario proposed for the hyena emphasizes the role of the vast steppe grasslands of Eurasia in contrast to models only involving Africa. The new methodology for combining genetic and geological data to test contrasting models of population history will be useful for a wide range of taxa where ancient and historic genetic data are available.     

Sex-specific androgen binding in spotted hyaena (Crocuta crocuta) plasma

• 1.1. A charcoal adsorption assay demonstrated a large variance in androgen binding ability in female spotted hyaenas.
• 2.2. A positive correlation between plasma androgen binding ability and ovarian steroid concentrations was demonstrated in adult females.
• 3.3. The strong plasma binding affinity for testosterone and dihydrotestosterone (DHT) (nM) together with the lack of cortisol and weaker oestradiol-17β binding suggests that a specific androgen binding substance, possibly a protein, is present in adult females of this species.
• 4.4. The lack of high affinity binding in male spotted hyaenas is unusual and deserves further investigation.
• 5.5. Some androgen binding in all, including males and immature animals suggests that albumin may bind some plasma androgens in this species.

     

Patterns of den occupation by the spotted hyaena (Crocuta crocuta)

Spotted hyaenas utilize isolated natal dens (NDs) and communal dens (CDs) for rearing their cubs. Here we describe patterns of natal and CD occupation by hyaenas belonging to one well‐studied clan in the Maasai Mara National Reserve during a 10‐year period. Locations of 98 den sites that were used as natal or CDs by hyaenas in the study clan were digitized in a Geographic Information System, and the duration of use of each den site, frequency of re‐use, and distances involved in den moves were quantified. Hyaenas moved their CD monthly on average. Most CD sites were occupied only once during the study, but several sites were used repeatedly. On rare occasions, the movement of hyaenas to a new den site could be attributed to a disturbance event at the CD, but factors regularly prompting hyaenas to move to new CD sites were unclear. High‐ranking female hyaenas were more likely to rear their cubs from birth in a CD than low‐ranking females. Low‐ranking females almost always utilized isolated NDs for the first few weeks of a litter's development, and low‐ranking females transferred their cubs over longer distances than did high‐ranking females.     

Hunting rates and hunting success in the spotted hyena (Crocuta crocuta)

Hunting group size, hunting rates and hunting success were monitored over a seven-year period among members of one large clan of spotted hyenas ( Crocuta crocuta ) inhabiting the Masai Mara National Reserve, in south-west Kenya. Prey availability varied seasonally in this study area, and hyenas tended to hunt whichever prey species were most abundant during each month of the year. Over 75% of 272 hunting attempts were made by lone hyenas, even when they hunted antelope three times their own body mass, such as wildebeest and topi. Of all prey that were commonly hunted, only zebra were usually hunted in groups. Approximately one-third of all hunting attempts resulted in prey capture. Although no significant sex differences were observed in juvenile or adult hunting rates, low-ranking adult females hunted at significantly higher hourly rates than did higher-ranking females. Hunting success was not influenced by the social rank of hunters, but hunting group size and hunter's age strongly influenced success. Young hyenas were poor hunters, and did not achieve adult competency levels until they were 5–6 years old.