T. Weingrill

Market forces predict grooming reciprocity in female baboons

We argue that grooming is a commodity that female primates can trade, either for itself or in exchange for other services (sensu biological markets theory) and that the decision to do either will depend on the degree of competition within a social group. We test this using data from four chacma baboon troops, living in two populations that differ markedly in the degree of contest competition. As predicted by the predominance of grooming dyads in which females are closely ranked there was, in all four troops, a positive correlation between the time invested by one partner and that by the other. In addition, as predicted, the allocation of time was more closely matched in troops where grooming could not be exchanged for anything else. In troops where resource competition was high, we found in one of two troops a positive relationship between rank distance and the discrepancy in time allocation, with the lower ranking of the partners contributing more grooming.

Day length, latitude and behavioural (in)flexibility in baboons (Papio cynocephalus ursinus)

Annual cycles in day length are an important consideration in any analysis of seasonal behaviour patterns, since they determine the period within which obligate diurnal or nocturnal animals must conduct all of their essential activities. As a consequence, seasonal variation in day length may represent an ecological constraint on behaviour, since short winter days restrict the length of the time available for foraging in diurnal species (with long summer days, and thus short nights, a potential constraint for nocturnal species). This paper examines monthly variation in activity patterns over a 4-year study of chacma baboons (Papio cynocephalus ursinus) at De Hoop Nature Reserve, South Africa. Time spent feeding, moving, grooming and resting are all significant positive functions of day length, even before chance events such as disease epidemics and climatically mediated home range shifts have been accounted for. These results provide strong support for the idea that day length acts as an ecological constraint by limiting the number of daylight hours and thus restricting the active period at certain times of year. Day length variation also has important implications across populations. Interpopulation variation in resting time, and non-foraging activity in general, is a positive function of latitude, with long summer days at temperate latitudes apparently producing an excess of time that cannot profitably be devoted to additional foraging or social activity. However, it is the short winter days that are probably of greatest importance, since diurnal animals must still fulfil their foraging requirements despite the restricted number of daylight hours and elevated thermoregulatory requirements at this time of year. Ultimately this serves to restrict the maximum ecologically tolerable group sizes of baboon populations with increasing distance from the equator. Seasonal variation in day length is thus an important ecological constraint on animal behaviour that has important implications both within and between populations, and future studies at non-equatorial latitudes must clearly be mindful of its importance.

MALE CONSORTSHIP BEHAVIOUR IN CHACMA BABOONS: THE ROLE OF DEMOGRAPHIC FACTORS AND FEMALE CONCEPTIVE PROBABILITIES

There is evidence for a general relationship between male dominance rank and mating success in primates, although the strength of this relationship differs among species. In chacma baboons (Papio hamadryas ursinus) male rank is found to be of more importance than in the other savannah baboon subspecies. However, even though the priority-of-access model explains the proportion of time spent in consortship for males of different rank in chacmas, highest-ranking males usually consort less often than expected. In this study, conducted in the Drakensberg Mountains of Natal and at De Hoop in the Western Cape, we investigated why dominant males in four study troops consorted only between 50% and 75% of days that they were expected to consort according to the priority-of-access model. Consortship success of highest-ranking males was primarily dependant on the number of available oestrous females in a troop. This was likely due to costs involved in consorting which limit the amount of days that a male could spend in consortship. Females pass through several cycles before conceiving and highest-ranking males were observed to consort more often on the conceptive cycle compared to the nearest nonconceptive cycle, but this was only true for males that were already resident for several months. Recently immigrated males that became highest-ranking often consorted during nonconceptive female cycles, while older, lower-ranking males consorted during the conceptive cycles. We propose that males with longer residency have more information about reproductive state of females and thus higher reproductive success than recently immigrated males.

INFANTICIDE AND THE VALUE OF MALE-FEMALE RELATIONSHIPS IN MOUNTAIN CHACMA BABOONS

Summary Associations between females and males over relatively long periods of time are common among savannah baboons (Papio cynocephalus). It seems clear that a female can benefit from close proximity to a male, since males are powerful partners in conflict situations with conspecifics and predators. For a male, proposed benefits of an association with a female are: (a) increased chances of mating with a female in the future through a positive effect on female choice and (b) increased fitness of the offspring sired with the female. In this study, data from a Drakensberg mountain chacma baboon troop (P. c. ursinus) were used to show that male-female associations were mainly between pregnant or lactating females and the putative fathers of their offspring. In general, associations had no effect on male consort success. One observed and one suspected infanticide occurred during the study, suggesting that the main benefit of male-female associations derives from infanticide avoidance. An immigrated male was observed killing an infant sired prior to his residence and was suspected of killing another infant sired during his residence. I suggest that an unusual high degree of paternity certainty and long alpha-male tenure made infanticide an adaptive reproductive strategy for the highestranking male even after several months of residence in the group (infants were killed five and ten months after male immigration). While the highest-ranking male did not often interact with his infant, only the other two fathers carried their respective inferred offspring. This

Effect of resource competition on the long-term allocation of grooming by female baboons: evaluating Seyfarth's model

Abstract Seyfarth (1977, Journal of Theoretical Biology, 65, 671–698) proposed an influential model that explained the long-term patterning of grooming relationships between female primates in terms of an interaction between the idealized grooming objectives of females and competition for valuable grooming partners. A critical test of the model requires a demonstration not only that competition for partners exists, but also that females do have an underlying target for the amount of grooming that they would like to receive relative to the amount they need to give. As it is not possible to stipulate a priori what this target is, or to detect it directly, we assessed its applicability by making a set of predictions, within the framework of Seyfarths model, as to how observed grooming patterns should change with changes in the intensity of resource competition. We tested these predictions with data from a troop of baboons, Papio hamadryas ursinus, for which the within-bout structure of grooming has already been shown to be sensitive to changes in resource competition (Barrett et al. 2002, Animal Behaviour, 63, 1047–1053). We found no evidence of competition for grooming partners and our results do not support the predictions of the modified model with respect to the existence of underlying grooming objectives. The findings that the grooming of female baboons is more diverse and that the mean rank distance separating partners increases when resource competition is greater, together with the absence of rank effects on long-term patterns, are, however, supportive of the recent biological market approach to social interactions. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Female baboons do not raise the stakes but they give as good as they get

We used data from four chacma baboon, Papio cynocephalus ursinus, troops, living in two populations, to test the raise the stakes (RTS) strategy of reciprocity. Female baboons did not raise the stakes either within or across grooming bouts. Instead they time-matched grooming contributions and divided grooming into short episodes. In addition, analysis of the grooming behaviour of frequently versus infrequently grooming dyads did not reveal differences in grooming patterns predicted by the RTS strategy. We suggest time constraints preclude the escalation of grooming bout length as required by RTS; the data were more consistent with a strategy of give as good as you get. However, this strategy could not explain all the patterns observed, and we conclude that biological market theory represents a more appropriate framework for investigating female grooming dynamics than dyadic games based on the iterated prisoners dilemma. We suggest that competitive altruism among individuals acts as a market force influencing an individuals value as a grooming partner. Copyright 2000 The Association for the Study of Animal Behaviour.

Indices of environmental temperatures for primates in open habitats.

Studies of thermoregulation in primates are under-represented in the literature, although there is sufficient evidence to suggest that temperature represents an important ecological constraint. One of the problems in examining thermoregulation in primates, however, is the difficulty in quantifying the thermal environment, since shade temperatures, solar radiation, humidity and wind speed all serve to alter an animal’s ‘perceived’ temperature. Since animals respond to their perceived temperature, we need methods to account for each of these factors, both individually and collectively, if we are to understand the integrated impact of the thermal environment on primates. Here, we present a review of some thermal indices currently available. Black bulb temperatures can account for the effect of solar radiation, with wind chill equivalent temperatures and the heat index providing quantifiable estimates of the relative impact of wind speed and humidity, respectively. We present three potential indices of the ‘perceived environmental temperature’ (PET) that account for the combined impact of solar radiation, humidity and wind speed on temperature, and perform a preliminary test of all of the climatic indices against behavioural data from a field study of chacma baboons (Papio cynocephalus ursinus) at De Hoop Nature Reserve, South Africa. One measure of the perceived environmental temperature, PET2, is an effective thermal index, since it enters the models for feeding and resting behaviour, and also accounts for levels of allogrooming. Solar radiation intensity is an important factor underlying these relationships, although the wind chill equivalent temperature and humidity enter the models for other behaviours. Future studies should thus be mindful of the impact of each of these elements of the thermal environment. A detailed understanding of primate thermoregulation will only come with the development of biophysical models of the thermal characteristics of the species and its environment. Until such developments, however, the indices presented here should permit a more detailed examination of the thermal environment, allowing thermoregulation to be given greater precedence in future studies of primate behaviour.