Illegal hunting of resident and migratory herbivores is widespread in the Serengeti National Park, Tanzania. To devise effective strategies to reduce levels of hunting, information is required on why people are involved in illegal hunting and the role of bushmeat in the local economy. Participation in hunting may be influenced by measures of relative wealth, including livestock ownership, means of generating cash income and access to alternative sources of meat. Data came from 300 individuals responding to a questionnaire in 10 villages, from responses by 359 people in 24 group discussions in another 12 villages, and from 552 people arrested and interviewed in the National Park. A smaller proportion of individual respondents (32%) than group respondents (57%) volunteered that they participated in illegal hunting. Most individual and group respondents were subsistence farmers who considered bushmeat to be a source of protein and a means of generating cash income. Three-quarters of those arrested participated in hunting primarily to generate cash income and a quarter claimed that they only hunted to obtain food. Participation in illegal hunting decreased as wealth in terms of the number of sheep and goats owned increased. People with access to alternative means of generating income or acquiring protein were also less likely to be involved in illegal hunting. Arrested respondents were typically young adult males with low incomes and few or no livestock. Illegal hunting was not reduced by participation in community-based conservation pro-grammes. Results suggested that between 52 000 and 60 000 people participated in illegal hunting within protected areas, and that many young men (approximately 5200) derived their primary source of income from hunting.

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In East Africa, spotted hyenas live in large clans in a highly structured society dominated by females. A clan is a fission-fusion society where members are often solitary or in small groups. Spotted hyenas have a ritualized greeting during which two individuals stand parallel and face in opposite directions. Both individuals usually lift their hind leg and sniff or lick the anogenital region of the other. The unique aspect of greetings between individuals is the prominent role of the erect 'penis' in animals of both sex. Female spotted hyenas have fused outer labiae and a 'pseudo-penis' formed by the clitoris which closely resembles the male penis and can be erected. During greetings subordinates signalled submission with gestures which were not necessarily reciprocated by the dominant participant. Asymmetries were most pronounced in greetings between adult females where the probability of asymmetries increased with the divergence in rank between partners. Greetings between adult females and males were uncommon and restricted to males above median rank, principally the alpha male. Models of primate affiliative behavior assume that benefits derived from social relationships with different individuals are not equal and that individuals are selected to maximize the benefits they receive from social relationships with others. The observed distribution of greetings between partners of different rank matched the predictions of these models. An examination of non-adaptive hypotheses on the evolution of the 'pseudo-penis' demonstrated that the conventional scenario linking (initial) virilization of female genitalia with selection for female dominance does not explain either the initial virilization, nor the evolution of the 'pseudo-penis' to its current form and use. We sketch a new scenario that links (1) initial virilization to the occurrence of neonatal siblicide amongst members of a twin litter, and (2) costs of maintenance, 'pseudo-penile' control over copulation and male submission. Our analysis confirms previous adaptive hypotheses on the function of the 'pseudo-penis' in greetCorrespondence to: M.L. East ings and suggests new hypotheses to account for hitherto unexplained features.

We investigated the reproductive outcomes of male and female mating tactics in the spotted hyena, Crocuta crocuta, a female-dominated social carnivore with high maternal investment, an absence of paternal care and female control over copulation. Paternity was determined using microsatellite profiling of 236 offspring in 171 litters from three clans. We found little evidence that male tactics that sought to coerce or monopolize females were successful. Polyandry and sperm competition appeared to counter effectively pre-copulatory male tactics, such as harassment, monopolization and other tactics, such as infanticide, that were against the evolutionary interests of females, and may have contributed to the stability of the male dominance hierarchy, which operated as a social queue. At least 39% of 54 females mated multiply, and 35% of 75 twin litters were fathered by two sires. Polyandry may also serve to ensure fertilization, compensate for an initial poor-quality mate or ensure fertilization by genetically compatible mates. Female mate choice matched observed patterns of affiliative male-female behaviour, indicating that affiliative behaviour is a successful male mating tactic, and was consistent with the idea that male tenure may serve as an index of male quality, although male fertility may decline with extreme old age.

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Summary1. Over the last three decades the main prey species (wildebeest Connochaetes taurinus , zebra Equus burchelli , Thomson's gazelle Gazella thomsoni , and Grant's gazelle Gazella granti ) of spotted hyaenas Crocuta crocuta in the Ngorongoro Crater, Tanzania, substantially declined in numbers, whereas buffalo Syncerus caffer numbers increased strongly. This provided a 'natural experiment' to investigate how a generalist predator such as the spotted hyaena responds to long-term changes in prey populations. Here we compare data on the feeding ecology of Crater hyaenas from the late 1990s ( period II) with equivalent data from the late 1960s (period I). 2. Hyaenas showed strong hunting preferences for wildebeest calves and gazelle fawns during both periods I and II. Adult buffaloes, prey difficult to hunt for hyaenas, were rarely hunted in either period despite a large increase in buffalo numbers from period I to II. 3. Hyaenas exhibited a functional response to the changes in prey populations by killing more buffalo calves and adult wildebeest during period II, relatively easy prey categories for hyaenas to hunt. 4. The proportion of carcasses scavenged and acquired by kleptoparasitizing other predators increased from period I to II. This was facilitated by an increase in the relative number of lion kills available to hyaenas during the same period. 5. Hyaenas defended a similar proportion of their kills against kleptoparasitic attempts by lions during both periods. Since the ratio of kills hyaenas kleptoparasitized from lions in relation to kills lost to lions increased from period I to period II, hyaenas obtained more food resources from lions during period II than I. 6. The success of hyaenas kleptoparasitizing lions was influenced by the presence of male lions and the ability of hyaenas to recruit sufficient clan members to appropriate carcasses from lions. 7. This study is the first to demonstrate both a functional response of a predator to changes in the size of prey populations and an associated change in foraging behaviour of the predator.