Look before you leap - individual variation in social vigilance shapes socio-spatial group properties in an agent-based model

Next to predator detection, primate vigilance also serves to keep track of relevant conspecifics. The degree of vigilance towards group members often reflects the dominance rank of an individual: subordinates pay attention to dominants. Although it has been suggested that subordinates' vigilance may result in spatial centrality of dominants, this has not been addressed in either empirical or modeling studies. Using agent-based models, we determined how social vigilance affects socio-spatial properties of primate groups. A basic model without social vigilance, where individuals avoid potential aggressors (avoidance model), was contrasted with two models that each additionally included a different type of social vigilance: a) monitoring a specific potential aggressor to remain informed on its whereabouts (monitoring model) or b) scanning the whole group to detect potential aggressors (scanning model). Adding monitoring or scanning behavior to the avoidance model reinforced spatial centrality of dominants, a pattern often observed in primates, and resulted in more spread out groups. Moreover, variation in scanning tendency alone was already sufficient to generate spatial centrality of dominants: frequently scanning subordinates could move further away from the group center than dominants, before losing sight of group members. In the monitoring model, two mechanisms caused decreased encounter frequencies among subordinates: a) increased inter-individual distances, and b) frequent monitoring of central dominants. In the scanning model, encounters among subordinates decreased due to increased inter-individual distances. This agent-based model study provides a clear indication that individual variation in social vigilance may be an important structuring feature of primate social groups.     

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     

Controls on coarse wood decay in temperate tree species: birth of the LOGLIFE experiment

Dead wood provides a huge terrestrial carbon stock and a habitat to wide-ranging organisms during its decay. Our brief review highlights that, in order to understand environmental change impacts on these functions, we need to quantify the contributions of different interacting biotic and abiotic drivers to wood decomposition. LOGLIFE is a new long-term 'common-garden' experiment to disentangle the effects of species' wood traits and site-related environmental drivers on wood decomposition dynamics and its associated diversity of microbial and invertebrate communities. This experiment is firmly rooted in pioneering experiments under the directorship of Terry Callaghan at Abisko Research Station, Sweden. LOGLIFE features two contrasting forest sites in the Netherlands, each hosting a similar set of coarse logs and branches of 10 tree species. LOGLIFE welcomes other researchers to test further questions concerning coarse wood decay that will also help to optimise forest management in view of carbon sequestration and biodiversity conservation.     

Controls on Coarse Wood Decay in Temperate Tree Species: Birth of the LOGLIFE Experiment

Dead wood provides a huge terrestrial carbon stock and a habitat to wide-ranging organisms during its decay. Our brief review highlights that, in order to understand environmental change impacts on these functions, we need to quantify the contributions of different interacting biotic and abiotic drivers to wood decomposition. LOGLIFE is a new long-term ‘common-garden’ experiment to disentangle the effects of species’ wood traits and site-related environmental drivers on wood decomposition dynamics and its associated diversity of microbial and invertebrate communities. This experiment is firmly rooted in pioneering experiments under the directorship of Terry Callaghan at Abisko Research Station, Sweden. LOGLIFE features two contrasting forest sites in the Netherlands, each hosting a similar set of coarse logs and branches of 10 tree species. LOGLIFE welcomes other researchers to test further questions concerning coarse wood decay that will also help to optimise forest management in view of carbon sequestration and biodiversity conservation.     

Predator guild does not influence orangutan alarm call rates and combinations

Monkey alarm calls have shown that in the primate clade, combinatorial rules in acoustic communication are not exclusive to humans. A recent hypothesis suggests that the number of different call combinations in monkeys increases with increased number of predator species. However, the existence of combinatorial rules in great ape alarm calls remains largely unstudied, despite its obvious relevance to ideas about the evolution of human speech. In this paper, we examine the potential use of combinatorial rules in the alarm calls of the only Asian great ape: the orangutan. Alarm calls in orangutans are composed of syllables (with either one or two distinct elements), which in turn are organized into sequences. Tigers and clouded leopards are predators for Sumatran orangutans, but in Borneo, tigers are extinct. Thus, orangutans make a suitable great ape model to assess alarm call composition in relation to the size of the predator guild. We exposed orangutans on both islands to a tiger and control model. Response compositionality was analyzed at two levels (i.e., syllable and syllable sequences) between models and populations. Results were corroborated using information theory algorithms. We made specific, directed predictions for the variation expected if orangutans used combinatorial rules. None of these predictions were met, indicating that monkey alarm call combinatorial rules do not have direct homologues in orangutans. If these results are replicated in other great apes, this indicates that predation did not drive selection towards ever more combinatorial rules in the human lineage.     

Sneak copulations in long-tailed macaques (Macaca fascicularis): no evidence for tactical deception

Sexual competition is highly prevalent within multi-male multi-female primate groups and may lead to copulations in absence of potentially interfering bystanders. Such avoidance of bystanders may result from tactical deception or from simpler mechanisms such as taking advantage of encountered situations without bystanders, operant conditioning or a peripheral positioning of non-alpha males. We investigated which individuals are avoided as bystanders, how individuals react to the presence of bystanders and whether copulation partners separate themselves from the group in a tactical way. Our observations of a group of 15 female and seven male long-tailed macaques housed in three interconnected, but visually separated compartments revealed that both males and females can interrupt sexual behaviour and that bystanders of both sexes were avoided during copulations (n?=?256). The strength of the effect of bystanders tended to decrease with the dominance rank of male bystanders, but did not depend on the dominance rank of female bystanders. The audience effects of non-alpha individuals did not depend on the strong audience effect of the alpha male in combination with proximity with the alpha male. The effects that we found for separate bystanders suggest that sexual competition concerns rank dependent male–male competition and rank independent female–female competition. Additionally, both male and female copulation partners paid attention to the presence of bystanders and conducted fewer copulation solicitations in their presence. The timing of a male and female’s separation from the group suggests that exploitation of the peripheral position of non-alpha males, and not tactical deception, may cause these audience effects.