Ecology,feeding competition and social structure in baboons

Predictions of the model of van Schaik (1989) of female-bonding in primates are tested by systematically comparing the ecology, level of within-group contest competition for food (WGC), and patterns of social behaviour found in two contrasting baboon populations. Significant differences were found in food distribution (percentage of the diet from clumped sources), feeding supplant rates and grooming patterns. In accord with the model, the tendencies of females to affiliate and form coalitions with one another, and to be philopatric, were strongest where ecological conditions promoted WGC. Group fission in the population with strong WGC was “horizontal” with respect to female dominance rank, and associated with female-female aggression during a period of elevated feeding competition. In contrast, where WGC was low, females’ grooming was focused on adult males rather than other females. Recent evidence suggests that group fission here is initiated by males, tends to result in the formation of one-male groups, and is not related to feeding competition but to male-male competition for mates. An ecological model of baboon social structure is presented which incorporates the effects of female-female competition, male-male competition, and predation pressure. The model potentially accounts for wide variability in group size, group structure and social relationships within the genus Papio. Socio-ecological convergence between common baboons and hamadryas baboons, however, may be limited in some respects by phylogenetic inertia. Received: 22 April 1994/Accepted after revision: 9 December 1995     

The evolution of female social relationships in nonhuman primates

Considerable interspecific variation in female social relationships occurs in gregarious primates, particularly with regard to agonism and cooperation between females and to the quality of female relationships with males. This variation exists alongside variation in female philopatry and dispersal. Socioecological theories have tried to explain variation in female-female social relationships from an evolutionary perspective focused on ecological factors, notably predation and food distribution. According to the current “ecological model”, predation risk forces females of most diurnal primate species to live in groups; the strength of the contest component of competition for resources within and between groups then largely determines social relationships between females. Social relationships among gregarious females are here characterized as Dispersal-Egalitarian, Resident-Nepotistic, Resident-Nepotistic-Tolerant, or Resident-Egalitarian. This ecological model has successfully explained differences in the occurrence of formal submission signals, decided dominance relationships, coalitions and female philopatry. Group size and female rank generally affect female reproduction success as the model predicts, and studies of closely related species in different ecological circumstances underscore the importance of the model. Some cases, however, can only be explained when we extend the model to incorporate the effects of infanticide risk and habitat saturation. We review evidence in support of the ecological model and test the power of alternative models that invoke between-group competition, forced female philopatry, demographic female recruitment, male interventions into female aggression, and male harassment. Not one of these models can replace the ecological model, which already encompasses the between-group competition. Currently the best model, which explains several phenomena that the ecological model does not, is a “socioecological model” based on the combined importance of ecological factors, habitat saturation and infanticide avoidance. We note some points of similarity and divergence with other mammalian taxa; these remain to be explored in detail. Received: 30 September 1996 / Accepted after revision: 20 July 1997     

To breed or not to breed—food competition and other factors involved in female breeding decisions in the pair-living nocturnal fork-marked lemur (<Emphasis Type="Italic">Phaner furcifer</Emphasis>)

Differential access to food resources is thought to be the main determinant of differences in female reproductive success but is poorly studied in both pair-living and nocturnal primates. The modes of food competition within and between families were investigated following the principles proposed by the ecological model using 3 years of field data from seven fork-marked lemur (Phaner furcifer) families. The major food resources were identified from year-round feeding observations and the strength and mode of competition were inferred from differences in physical condition. The most important food resource of fork-marked lemurs were tree exudates which occurred in small, defendable food patches, characterized by fast depletion and rapid renewal. These characteristics led to strong within-group contest and scramble competition, which were found to yield a positive dominance effect and a negative group-size effect on female net energy gain. Differential physical condition, however, did not translate directly into differential reproductive success. Low female fertility was best predicted by large family size associated with delayed dispersal by previous offspring. Although there is no obvious benefit from full-grown offspring in their territory, adults tolerate delayed natal dispersal, probably because dispersal poses extraordinary costs for the offspring. These costs are likely to accrue from decreased foraging efficiency in unfamiliar habitats because exudate feeding requires very rigid feeding itineraries. In conclusion, the presented evidence for group-size effects on reproductive success in pair-living females opens a new area for research on the costs and benefits of delayed dispersal and female reproductive decisions.This revised version was published in September 2003 with corrections to the Authors Present address.An erratum to this article can be found at Communicated by E.H.M. Sterck     

Socioecological influences on the reproductive success of female mountain gorillas (<Emphasis Type="Italic">Gorilla beringei beringei</Emphasis>)

Over the past few decades, socioecological models have been developed to explain the relationships between the ecological conditions, social systems, and reproductive success of primates. Feeding competition, predation pressures, and risk of infanticide are predicted to influence how female reproductive success (FRS) depends upon their dominance rank, group size, and mate choices. This paper examines how those factors affected the reproductive success of female mountain gorillas (Gorilla beringei beringei) of the Virunga Volcanoes, Rwanda from 1967–2004. Reproductive success was measured through analyses of interbirth intervals, infant survival, and surviving infant birth rates using data from 214 infants born to 67 females. Mountain gorillas were predicted to have “within-group scramble” feeding competition, but we found no evidence of lower FRS in larger groups, even as those groups became two to five times larger than the population average. The gorillas are considered to have negligible “within-group contest” competition, yet higher ranked mothers had shorter interbirth intervals. Infant survival was higher in multimale groups, which was expected because infanticide occurs when the male dies in a one-male group. The combination of those results led to higher surviving birth rates for higher ranking females in multimale groups. Overall, however, the socioecological factors accounted for a relatively small portion of the variance in FRS, as expected for a species that feeds on abundant, evenly distributed foliage.     

Agonistic relationships between female mountain gorillas (Gorilla gorilla beringei)

Females in some mountain gorilla groups can be ranked on the basis of decided, non-aggressive approach-retreat interactions. However, data on four gorilla groups show that females do not form clear agonistic dominance hierarchies. Most aggressive interactions between females were undecided, and most dyads had undecided dominance relationships (Fig. 2). Females ignored aggression or responded to it aggressively more often than they responded submissively (Fig. 3). Moreover, directional inconsistency in aggression (including escalated contests) was high, and in some groups females showed significant bidirectionality in initiation of aggression (i.e., the more often they received aggression from partners, the more often they directed aggression at those partners; Fig. 4). Assigning ranks on the basis of aggressive interactions has limited power to explain variation in rates of initiating aggression and in responses and outcomes to aggression. Aggression was most common during feeding, but usually did not interrupt feeding bouts. It was proportionately more common in contexts that seem to involve competition for access to males (Figs 5, 6). This is consistent with the argument that females depend crucially on male services. As predicted by current models of female primate social relationships, agonistic relationships between gorilla females are unlike those typical in cercopithecine primates. They resemble those of some other primates and some equids in which contest competition for food is relatively unimportant and ecological costs of female transfer are low.     

Within-group spatial position and vigilance: a role also for competition? The case of impalas (<Emphasis Type="Italic">Aepyceros melampus</Emphasis>) with a controlled food supply

Theory predicts that individuals at the periphery of a group should be at higher risk than their more central conspecifics since they would be the first to be encountered by an approaching terrestrial predator. As a result, it is expected that peripheral individuals display higher vigilance levels. However, the role of conspecifics in this “edge effect” may have been previously overlooked, and taking into account the possible role of within-group competition is needed. Vigilance behavior in relation to within-group spatial position was studied in impalas (Aepyceros melampus) feeding on standardized patches. We also controlled for food distribution in order to accurately define a “central” as opposed to a “peripheral” position. Our data clearly supported an edge effect, with peripheral individuals spending more time vigilant than their central conspecifics. Data on social interactions suggest that it was easier for a foraging individual to defend its feeding patch with its head lowered, and that more interactions occurred at the center of the group. Together, these results indicate that central foragers may reduce their vigilance rates in response to increased competition. Disentangling how the effects of competition and predation risk contribute to the edge effect requires further investigations.     

Competition and group size in Thomas’s langurs (Presbytis thomasi): the folivore paradox revisited

Among primates, group size is highly variable. The standard ecological model assumes that better predation avoidance as group size increases favours living in larger groups, whereas increased travel costs and reduced net food intake due to within-group competition for resources set the upper limit. Folivorous primates, however, tend to defy this generalisation in that some live in small groups despite low costs of feeding competition. To resolve this ’folivore paradox’, it has been suggested that folivore group size is limited by social factors such as male harassment or infanticide, or that females can disperse more easily and thus maintain group size near optimum levels. In this paper, we examine the effects of group size on home range size, day-journey length, activity budget and diet in wild Thomas’s langurs (Presbytis thomasi), which live in one-male multi-female groups with a limited life cycle. We examined only data from the stable middle tenure phase when factors such as the strength of the breeding male or the way in which groups were formed did not influence ranging and activities. During this phase, group size affected day-journey length and home range size, and had a minor effect on diet, but did not influence time spent feeding or resting, allogrooming or birth rates. Hence the upper limit to group size during the middle tenure phase in Thomas’s langurs is not set by feeding competition. The folivore paradox is not due to frequent female dispersal in Thomas’s langurs. The timing of female dispersal is not as expected if it serves to keep group sizes near the ecological optimum, and groups seem to be below this optimum. Instead, female reproductive success is presumably maximised in small to mid-sized groups because larger groups show a clear trend to experience higher risk of take-over, often accompanied by infanticide. Because females can redistribute themselves among nearby groups when groups reorganise each time a new male starts up a new group, females can keep the group small. Thus, a social factor, risk of infanticide, seems to provide the selective advantage to small group size in Thomas’s langurs. Received: 29 July 1999 / Revised: 17 November 1999 / Accepted: 15 October 2000     

Production and perception of situationally variable alarm calls in wild tufted capuchin monkeys (Cebus apella nigritus)

Many mammalian and avian species produce conspicuous vocalizations upon encountering a predator, but vary their calling based on risk urgency and/or predator type. Calls falling into the latter category are termed “functionally referential” if they also elicit predator-appropriate reactions in listeners. Functionally referential alarm calling has been well documented in a number of Old World monkeys and lemurs, but evidence among Neotropical primates is limited. This study investigates the alarm call system of tufted capuchin monkeys (Cebus apella nigritus) by examining responses to predator and snake decoys encountered at various distances (reflecting differences in risk urgency). Observations in natural situations were conducted to determine if predator-associated calls were given in additional contexts. Results indicate the use of three call types. “Barks” are elicited exclusively by aerial threats, but the call most commonly given to terrestrial threats (the “hiccup”) is given in nonpredatory contexts. The rate in which this latter call is produced reflects risk urgency. Playbacks of these two call types indicate that each elicits appropriate antipredator behaviors. The third call type, the “peep,” seems to be specific to terrestrial threats, but it is unknown if the call elicits predator-specific responses. “Barks” are thus functionally referential aerial predator calls, while “hiccups” are better seen as generalized disturbance calls which reflect risk urgency. Further evidence is needed to draw conclusions regarding the “peep.” These results add to the evidence that functionally referential aerial predator alarm calls are ubiquitous in primates, but that noncatarrhine primates use generalized disturbance calls in response to terrestrial threats.     

Do males and females differ in the feeding of large and small siblings? An experiment with the bluethroat

Males and females have been reported to differ in their feeding of large and small siblings in several species of birds. According to recent hypotheses, this phenomenon may be related to a sexual conflict over avian hatching patterns. We designed an experiment to test for the existence of such a sex difference by manipulating nestling size hierarchies of the bluethroat (Luscinia s. svecica) in two directions; half the broods were “asynchronized” to yield large size-differences within broods and the other half were “synchronized” to yield small size-differences. In all broods, nestlings were categorized as being either large or small according to body mass. We recorded male and female food distribution by video early (day 4 after hatching) and late (day 8) in the nestling period. Males and females did not differ in their distribution of food among different-sized nestlings. With large size-differences, both males and females fed large nestlings nearly twice as often as small ones. In contrast, when the size-differences were small, food was more evenly distributed among nestlings. Early in the nestling period, males fed more nestlings during each feeding visit than did females. Our finding that male and female bluethroats do not differ in the feeding of large and small siblings is in contrast to most previous studies. Variation in costs and benefits to males and females from feeding different-sized nestlings, and restrictions to parental choice due to nestling interactions, may explain interspecific variation. Received: 27 June 1997 / Accepted after revision: 26 January 1998     

Do bumble bee queens choose nest sites to maximize foraging rate? Testing models of nest site selection

We proposed “foundress-max” hypothesis that a bumble bee foundress chooses her nest site to maximize her energy intake rate from nectar. To examine the hypothesis, we estimated the maximum energy intake rate at each site in the study area and compared the distribution of the maximum energy intake rates with those of actual nest sites. We also calculated rank correlations of the maximum energy intake rate with the number of nest-searching foundresses at 54 sites. The nest locations supported the foundress-max hypothesis, but the number of nest-searching foundresses did not. This could be attributed to the density of food sites: many food sites may attract many foundresses. Therefore, we subsequently proposed “foundress-sum” hypothesis that a foundress chooses her nest site to maximize the sum of energy intake rates. The nest locations supported the foundress-max hypothesis more than the foundress-sum hypothesis. A profitable food site would affect foundresses’ nest site selection.     

Individual dispersal status influences space use of conspecific residents in the common lizard, Lacerta vivipara

The effects of immigration on the behaviour of residents may have important implications for the local population characteristics. A manipulative laboratory experiment with yearlings of the common lizard (Lacerta vivipara) was performed to test whether the introduction of dispersing or philopatric individuals influences the short-term spacing behaviour of resident individuals. Staged encounters were carried out to induce interactions within dyads. The home cage of each responding individual was connected by a corridor to an unfamiliar “arrival cage” to measure the latency to leave their own home cage after each encounter. Our results showed that the time that pairs spent in close proximity was longer when a dispersing individual was introduced in the home cage. The latency to leave the home cage was longer after the introduction of a dispersing individual. These response variables were not influenced by the relative body sizes of contestants nor by the level of aggression towards each other. In contrast, the aggressive response was significantly influenced by the residency asymmetry established experimentally (“owner” of the home cage vs introduced individual). Our results suggest that the space use by resident individuals is influenced by the dispersal status of conspecifics. The potential ultimate causes driving this effect are discussed.     

Rank differences in energy intake rates in white-faced capuchin monkeys, Cebus capucinus: the effects of contest competition

The effect of aggressive competition over food resources on energy intake rate is analyzed for individuals of three groups of 25–35 white-faced capuchin monkeys, Cebus capucinus, living in and near Lomas Barbudal Biological Reserve, Costa Rica. An individuals energy intake rate on a given food species was affected by its rank and the number of agonistic interactions within the feeding tree. Dominant group members had higher energy intake rates relative to subordinate group members whether or not there was agonism within the feeding tree. Low- and mid-ranked individuals had lower energy intake rates in trees with higher amounts of aggression, while energy intake rate of high-ranked individuals was not affected by the amount of aggression in the feeding tree. Energy intake was not influenced by the sex of the individual when rank was held constant statistically. Energy intake was positively correlated with total crown energy (measured in kilojoules) within the feeding tree for two of three study groups. This difference may be explained by the quality of each groups territory. Finally, high-ranked individuals are responsible for the majority of agonism within feeding trees and target middle- and low-ranked individuals equally. These findings fit the predictions of current socioecological models for within-group contest competition over food resources. The results of this study suggest that within-group competition affects energy intake rate in white-faced capuchin monkeys.     

Relative importance of multiple plumage ornaments as status signals in golden whistlers (<Emphasis Type="Italic">Pachycephala pectoralis</Emphasis>)

Status signals are traits that advertise an individual’s competitive abilities to conspecifics during aggressive disputes. Most studies of status signals in birds have focussed on melanin-based plumage signals, but recent research shows that carotenoid-based signals may also play a role in aggressive signaling. We assessed the relative importance of melanin- and carotenoid-based plumage patches as agonistic signals in a small passerine, the golden whistler (Pachycephala pectoralis). Display signals in male golden whistlers include an unpigmented white throat patch, a carotenoid-based yellow breast and nape band, and a melanin-based black chin-stripe. We found that only the white throat patch was correlated with contest-related attributes. Males possessing large throat patches defended larger territories and commenced breeding earlier. When caged males with either experimentally reduced, or unmanipulated throat patches were presented to conspecifics, those with experimentally reduced patches attracted less aggression from male subjects. Focal males also responded faster to caged males with throat patches similar in size to their own, suggesting that they may assess relative throat patch size before engaging in aggressive encounters. Females did not discriminate between “reduced” or “control” treatments. Our data strongly suggest that only the unpigmented throat patch functions as a status signal. As this signal is unlikely to have significant development costs, honesty may be maintained through social costs.     

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Cooperation in animal social groups may be limited by the threat of free riding, the potential for individuals to reap the benefits of other individuals actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other groups territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.Communicated by P. Kappeler     

Predation, scramble competition, and the vigilance group size effect in dark-eyed juncos (Junco hyemalis)

In socially feeding birds and mammals, as group size increases, individuals devote less time to scanning their environment and more time to feeding. This vigilance “group size effect” has long been attributed to the anti-predatory benefits of group living, but many investigators have suggested that this effect may be driven by scramble competition for limited food. We addressed this issue of causation by focusing on the way in which the scan durations of free-living dark-eyed juncos (Junco hyemalis) decrease with group size. We were particularly interested in vigilance scanning concomitant with the handling of food items, since a decrease in food handling times (i.e. scan durations) with increasing group size could theoretically be driven by scramble competition for limited food resources. However, we showed that food-handling scan durations decrease with group size in an environment with an effectively unlimited food supply. Furthermore, this food-handling effect was qualitatively similar to that observed in the duration of standard vigilance scans (scanning exclusive of food ingestion), and both responded to changes in the risk of predation (proximity of a refuge) as one might expect based upon anti-predator considerations. The group size effects in both food-handling and standard scan durations may reflect a lesser need for personal information about risk as group size increases. Scramble competition may influence vigilance in some circumstances, but demonstrating an effect of competition beyond that of predation may prove challenging. Received: 22 September 1998 / Received in revised form: 1 February 1999 / Accepted: 14 February 1999     

Proximate mechanisms of contest competition among female Bwindi mountain gorillas (Gorilla beringei beringei)

Socioecological models provide a framework for predicting how animals respond competitively to the abundance and distribution of food resources. Testing predictions of socioecological models requires analysis of relationships among food resource characteristics, competitive behaviors, and measures of rank-related skew in energy balance or reproductive success. A positive relationship between dominance rank and energy balance has been observed among female mountain gorillas in Bwindi Impenetrable National Park, Uganda. This study examines the proximate mechanisms underlying feeding competition among those females. To assess the contestability of food resources, we measured the time a female spent feeding at a food site (food site residence time). We also examined the relationship between dominance rank and the access to resources, as well as the rate, context, and direction of aggression, and the number of neighbors in close proximity. As predicted, females had longer food site residence times and higher aggression rates with fruit and decaying wood than with herbaceous vegetation, suggesting that those resources may be contestable. Aggression was predominantly directed down the dominance hierarchy, although against expectation, rank was not significantly correlated with aggression rates or the time spent feeding on contestable foods. Higher-ranking females had significantly fewer neighbors, suggesting that lower-ranking females avoid higher-ranking ones. This study provides additional support for the claim that there is variability in how primates respond to the quality and distribution of food resources and that avoidance as a strategy to cope with feeding competition may result in similar skew in energy balance as rank-related aggression.     

Risk-prone hunting by chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>) increases during periods of high diet quality

Most studies suggest that during times of nutritional stress, an animal faced with two foraging choices should follow a risk-prone strategy, choosing the option with highest payoff variance. This “scarcity/risk” hypothesis was developed to account for the foraging patterns of small animals with high metabolic rates susceptible to the threat of starvation. In this paper, we propose that animals should also be risk-prone when their diet quality is particularly high, far exceeding that which is needed to survive. Under these circumstances, the costs of experiencing a low or negative payoff can easily be recouped. We suggest that large-bodied omnivores are most likely to adopt this “abundance/risk” strategy. We investigate this question among wild chimpanzees (Pan troglodytes) that choose between a risk-averse strategy of feeding on plant material and a risk-prone strategy of hunting red colobus monkeys. Using 14 years of data on the Kanyawara chimpanzees of Kibale National Park, Uganda, we find strong evidence that chimpanzees follow the “abundance/risk” strategy. Both hunting rate (hunts/100 observation hours) and the probability of hunting upon encountering red colobus monkeys were positively correlated with seasonal consumption of ripe drupe fruits, a class of preferred food associated with elevated reproductive performance by females. Critically, these results remained statistically significant after controlling for the potentially confounding effects of male chimpanzee party size and the presence of sexually receptive females. These findings suggest that the relationship between risk-sensitive foraging and diet quality depends upon the daily probability of starvation, the number of alternative foraging strategies, and the degree to which diet quality satisfies an animal’s nutritional requirements.     

Within-group spatial position in ring-tailed coatis: balancing predation, feeding competition, and social competition

A variety of factors can influence an individual’s choice of within-group spatial position. For terrestrial social animals, predation, feeding success, and social competition are thought to be three of the most important variables. The relative importance of these three factors was investigated in groups of ring-tailed coatis (Nasua nasua) in Iguazú, Argentina. Different age/sex classes responded differently to these three variables. Coatis were found in close proximity to their own age/sex class more often than random, and three out of four age/sex classes were found to exhibit within-group spatial position preferences which differed from random. Juveniles were located more often at the front edge and were rarely found at the back of the group. Juveniles appeared to choose spatial locations based on feeding success and not predation avoidance. Since juveniles are the most susceptible to predation and presumably have less prior knowledge of food source location, these results have important implications in relation to predator-sensitive foraging and models of democratic group leadership. Subadults were subordinate to adult females, and their relationships were characterized by high levels of aggression. This aggression was especially common during the first half of the coati year (Nov–April), and subadults were more peripheralized during this time period. Subadults likely chose spatial positions to avoid aggression and were actively excluded from the center of the group by adult females. In the Iguazú coati groups, it appeared that food acquisition and social agonism were the major determinants driving spatial choice, while predation played little or no role. This paper demonstrates that within-group spatial structure can be a complex process shaped by differences in body size and nutritional requirements, food patch size and depletion rate, and social dominance status. How and why these factors interact is important to understanding the costs and benefits of sociality and emergent properties of animal group formation.     

The lemur syndrome unresolved: extreme male reproductive skew in sifakas (<Emphasis Type="Italic">Propithecus verreauxi</Emphasis>), a sexually monomorphic primate with female dominance

The primates of Madagascar (Lemuriformes) are unusual among mammals in that polygynous species lack sexual dimorphism, and females dominate males socially in most species. Moreover, lemur groups are relatively small and characterized by even adult sex ratios despite the fact that one male should be able to exclude other males from the group. One hypothesis to explain this combination of behavioral, morphological, and demographic traits (the “lemur syndrome”) postulates that male–male competition is relaxed and, hence, variance in male reproductive success is low. Reproductive skew theory provides a framework for testing this and several related predictions about lemur social evolution. Specifically, low reproductive skew is also predicted if dominant males or adult females make reproductive concessions to subordinates or if the latter group successfully pursues alternative reproductive tactics. However, suitable data on paternity, demography, and behavior for a conclusive test of these predictions have not been available in the past. In this paper, we show that male reproductive success in ten groups of Verreaux’s sifakas (Propithecus verreauxi) was extremely skewed in favor of the dominant male over 9 years. Our genetic analyses also revealed that more than a third of all groups are effectively harem groups because only one male was unrelated to the resident female(s). In groups with two or more non-natal males, the dominant sired 91% of 33 infants. Together, males pursuing one of several alternative reproductive tactics, such as roaming among several groups or immigrating peacefully, sired only 11% of infants. Thus, female sifakas do not control group composition by offering reproductive opportunities to subordinate males as staying incentives, intrasexual selection is not relaxed, and dominant males prevail in a tug-of-war over subordinate males. Because male reproductive skew in sifakas is even more pronounced than in harem-living anthropoids studied to date, intrasexual selection is clearly not relaxed, and the lemur syndrome is more puzzling than ever.     

Darwin’s “tug-of-war” vs. starlings’ “horse-racing”: how adaptations for sequential encounters drive simultaneous choice

Charles Darwin aided his private decision making by an explicit deliberation, famously deciding whether or not to marry by creating a list of points in a table with two columns: “Marry” and “Not Marry”. One hundred seventy-two years after Darwin’s wedding, we reconsider whether this process of choice, under which individuals assign values to their options and compare their relative merits at the time of choosing (the tug-of-war model), applies to our experimental animal, the European Starling, Sturnus vulgaris. We contrast this with the sequential choice model that postulates that decision-makers make no comparison between options at the time of choice. According to the latter, behaviour in simultaneous choices reflects adaptations to contexts with sequential encounters, in which the choice is whether to take an opportunity or let it pass. We postulate that, in sequential encounters, the decision-maker assigns (by learning) a subjective value to each option, reflecting its payoff relative to background opportunities. This value is expressed as latency and/or probability to accept each opportunity as opposed to keep searching. In simultaneous encounters, choice occurs through each option being processed independently, by a race between the mechanisms that generate option-specific latencies. We describe these alternative models and review data supporting the predictions of the sequential choice model.