Codon usage in the vertebrate hemoglobins and its implications

A study of codon usage in vertebrate hemoglobins revealed an evolutionary trend toward elevated numbers of CpG codon boundary pairs in mammalian hemoglobin alpha genes. Selection for CpG codon boundaries countering the generally observed CpG suppression is strongly suggested by these data. These observations parallel recently published experimental results that indicate that constitutive expression of the human alpha-globin gene appears to be determined by regulatory information encoded within the structural gene. The possibility is raised that, in the absence of selection, CpG decay can be used to date the evolutionary origin of a mammalian alpha pseudogene from its active alpha gene.      

Fission of a Spotted Hyena Clan: Consequences of Prolonged Female Absenteeism and Causes of Female Emigration

We describe the fission of one large East African clan of spotted hyenas (Crocuta crocuta). During an 8-mo period from late 1989 to early 1990, 8 of 22 adult females vanished from the clan, along with their juvenile offspring. 6 of these 8 adult females formed a new clan in an area adjacent to the parent clan's home range. The 6 adult female dispersers belonged to three mid- or low-ranking matrilines in the parent clan. Two juvenile males returned to the parent clan for temporary visits, and three female members of one matriline rejoined the parent clan after absences of over 8 mo. Returning females, but not males, were targets of severe aggression by adult female residents of the parent clan, and by juvenile residents of both sexes. Returning females fell from their previous mid-ranking positions to the bottom of the adult female hierarchy. Our data suggest that clan fission was promoted by the simultaneous occurrence of low food availability, an unusually high density of higher-ranking conspecific competitors, and the availability of neighboring vacant habitat.     

Risk-taking in free-living spotted hyenas is associated with anthropogenic disturbance,predicts survivorship,and is consistent across experimental contexts

Anthropogenic disturbance can have important influences on the fitness and behaviors of wild animals, including their boldness when exposed to risky conditions. We presented spotted hyenas (Crocuta crocuta) from two populations, each exposed to a different level of human activity, with a life-size model hyena representing an intruder from another clan. The high-disturbance population lived adjacent to human settlements, and the low-disturbance population inhabited a relatively undisturbed part of the same national park in Kenya. The mock intruder was presented to individual hyenas to assess their reactions to an alien hyena, and to determine whether their reactions varied with their exposure to anthropogenic activity. We found that human disturbance was indeed associated with hyena risk-taking behavior in response to the model intruder. Hyenas tested in the low-disturbance area exhibited more risk-taking behaviors by approaching the mock intruder more closely, and spending more time near it, than did their counterparts living in high-disturbance areas. Hyenas that spent less time in close proximity to the model had greater survivorship than those that spent more time in close proximity to it, regardless of disturbance level. Furthermore, the individual differences in risk-taking measured here were consistent with those obtained previously from the same animals using a different set of experimental manipulations. However, the experimentally induced behaviors were not consistent with naturally occurring risk-taking behaviors in proximity to lions; this suggests that risk-taking behaviors are consistent within individuals across experimental contexts, but that exposure to lions elicits different responses. Although our results are consistent with those from earlier tests of anthropogenic disturbance and boldness in spotted hyenas and other predators, they differ from results obtained from birds and small mammals, which are generally found to be bolder in areas characterized by human disturbance.     

Energetic and endocrine mediation of natal dispersal behavior in Belding's ground squirrels

Natal dispersal, the permanent departure of an individual from its birth site, is sex biased in most mammals, with males dispersing at higher rates or over greater distances than do female conspecifics. Because dispersal movements may be energetically expensive, their occurrence should theoretically be influenced by energy availability. Moreover, the male bias typical of mammalian dispersal suggests that this behavior might be mediated by gonadal androgens. Using free-living Belding's ground squirrels (Spermophilus beldingi) as subjects, we provisioned juveniles with extra food to evaluate energetic influences on male dispersal behavior. Provisioning increased body mass and body fat of juvenile males and caused them to disperse at younger ages, but did not affect blood glucose levels. Dispersing males were fatter than same-aged males that had not yet dispersed. Moreover, body fat of provisioned and unprovisioned males did not differ when evaluated relative to the week during which they dispersed, suggesting that there may be a fat threshold for dispersal. In a second experiment, we measured plasma concentrations of testosterone (T) in provisioned and unprovisioned, free-living juveniles to evaluate the hypothesis that male dispersal behavior is activated by concurrent high levels of T. We observed no increase in plasma T associated with dispersal by juvenile males, no sex differences in circulating T among juveniles, and no effects of food provisioning on juvenile T levels. In a third experiment with free-living S. beldingi, we concurrently altered early androgen exposure by treating females with T at birth and manipulated energy availability by food provisioning. Perinatal T-treatment increased the likelihood of dispersal among juvenile females. Provisioning increased body mass and body fat of juveniles and caused males and T-treated females to disperse at significantly younger ages than either their unprovisioned counterparts or the few provisioned control females that dispersed. These results suggest that early T exposure in this species determines the probability of dispersal, whereas the amount of energy an individual has stored as fat strongly influences the timing of dispersal. Early T exposure also appears to cause the timing of dispersal to respond to energy availability and body fat in a male-typical way, possibly by organizing masculine mass and fat thresholds for dispersal.     

Behavioural structuring of relatedness in the spotted hyena (Crocuta crocuta) suggests direct fitness benefits of clan-level cooperation

Spotted hyenas (Crocuta crocuta) are gregarious carnivores that live in multigenerational social groups, called clans, containing one to several matrilines. Members of multiple matrilines within a clan cooperate during dangerous interactions with inter- and intraspecific competitors. The evolution of cooperation may be influenced by relatedness between individuals, which in turn is influenced by reproductive skew and mate choice, dispersal and territorial behaviours. Behavioural data exist for spotted hyenas, but corresponding data on patterns of relatedness are unavailable; this lack of data makes it difficult to assess the relative importance of selection pressures favouring cooperative behaviour within and among groups. Therefore we conducted a longitudinal analysis of relatedness within a single large clan of spotted hyenas, as well as a cross-sectional analysis of relatedness among hyenas from multiple clans. Within a clan, patterns of relatedness reflected known pedigree relationships, and relatedness was higher within than among matrilines, even across generations. Although mean within-matriline relatedness varied among matrilines, it did not decline with matriline rank. On average, clan members were not related closely, due to high levels of male-mediated gene flow among clans, and relatedness declined very slightly across clan borders. Low mean relatedness within clans suggests that spotted hyenas cooperate with unrelated clan-mates against close paternal kin in other clans. Our data also suggest that spotted hyenas must derive large net direct fitness benefits from group living and cooperation.     

Mechanisms of maternal rank 'inheritance' in the spotted hyaena, Crocuta crocuta

Maternal rank 'inheritance', the process by which juveniles attain positions in the dominance hierarchy adjacent to those of their mothers, occurs in both cercopithecine primates and spotted hyaenas. Maternal rank is acquired in primates through defensive maternal interventions, coalitionary support and unprovoked aggression ('harassment') directed by adult females towards offspring of lower-ranking individuals. Genetic heritability of rank-related traits plays a negligible role in primate rank acquisition. Because the social lives of Crocuta and cercopithecine primates share many common features, we examined whether the same mechanisms might operate in both taxa to promote maternal rank 'inheritance'. We observed a large clan of free-living spotted hyaenas in Kenya to test predictions of four mechanistic hypotheses. Hyaena rank acquisition did not appear to be directly affected by genetic heritability. Unprovoked aggression from adult female hyaenas was not directed preferentially towards low-ranking cubs. However, high-ranking mothers intervened on behalf of their cubs more frequently and more effectively than low-ranking mothers. Maternal interventions and supportive coalitions appeared to reinforce aggression directed at 'appropriate' conspecific targets, whereas coalitionary aggression directed at cubs apparently functioned to extinguish their aggressive behaviour towards 'inappropriate' targets. Young hyaenas and primates thus appear to 'inherit' their mothers' ranks by strikingly similar mechanisms. Copyright 2000 The Association for the Study of Animal Behaviour.     

Topological effects of network structure on long‐term social network dynamics in a wild mammal

Social structure influences ecological processes such as dispersal and invasion, and affects survival and reproductive success. Recent studies have used static snapshots of social networks, thus neglecting their temporal dynamics, and focused primarily on a limited number of variables that might be affecting social structure. Here, instead we modelled effects of multiple predictors of social network dynamics in the spotted hyena, using observational data collected during 20 years of continuous field research in Kenya. We tested the hypothesis that the current state of the social network affects its long‐term dynamics. We employed stochastic agent‐based models that allowed us to estimate the contribution of multiple factors to network changes. After controlling for environmental and individual effects, we found that network density and individual centrality affected network dynamics, but that social bond transitivity consistently had the strongest effects. Our results emphasise the significance of structural properties of networks in shaping social dynamics.     

Innovative problem solving by wild spotted hyenas

Innovative animals are those able to solve novel problems or invent novel solutions to existing problems. Despite the important ecological and evolutionary consequences of innovation, we still know very little about the traits that vary among individuals within a species to make them more or less innovative. Here we examine innovative problem solving by spotted hyenas (Crocuta crocuta) in their natural habitat, and demonstrate for the first time in a non-human animal that those individuals exhibiting a greater diversity of initial exploratory behaviours are more successful problem solvers. Additionally, as in earlier work, we found that neophobia was a critical inhibitor of problem-solving success. Interestingly, although juveniles and adults were equally successful in solving the problem, juveniles were significantly more diverse in their initial exploratory behaviours, more persistent and less neophobic than were adults. We found no significant effects of social rank or sex on success, the diversity of initial exploratory behaviours, behavioural persistence or neophobia. Our results suggest that the diversity of initial exploratory behaviours, akin to some measures of human creativity, is an important, but largely overlooked, determinant of problem-solving success in non-human animals.