Flexibility in the social structure of male chimpanzees (Pan troglodytes schweinfurthii) in the Budongo Forest, Uganda

ABSTRACTIndividuals of social species face a trade-off between the competitive costs and social benefits of group living. Species show a range of social strategies to deal with this trade-off, for example atomistic fission-fusion dynamics in which temporary social groups of varying size and membership form and re-form; or molecular fission-fusion dynamics which contain stable sets of multilevel nested subgroups. Chimpanzees are considered an archetypical atomistic fission-fusion species, using dynamic changes in day-to-day association to moderate the costs of within-group competition. It has been argued that humans’ highly flexible social organisation allows us to live in extremely large groups. Using four years of association data from two neighbouring communities of East African chimpanzees (Pan troglodytes schweinfurthii), we describe new levels of flexibility in chimpanzee social organisation and confirm the presence of subgrouping in a second, large community of chimpanzees. We show that males from the larger Waibira community (N males 24-31) exhibited additional levels of semi-stable subgrouping, while males from the smaller Sonso community (N males 10-13) did not. Subgroup membership showed stability across some years, but flexibility across others. Our data support the hypothesis that chimpanzees can incorporate strategies other than fission-fusion to overcome costs of social living, and that their social organisation may be closer to that of modern humans than previously described.SIGNIFICANCE STATEMENTSocial living offers benefits and costs; groups can more easily locate and defend resources, but experience increased individual competition. Many species, or individuals, flexibly adjust their social organization when faced with different competitive pressures. It is argued that humans are unique among primates in combining multigroup social organisation with fission-fusion dynamics flexibly within and across groups, and that doing so allows us to live in extremely large groups. Using four-years of association data we show new levels of flexibility in chimpanzee social organization. Males from a typically-sized community employed atomistic fission-fusion dynamics, but males in an unusually large community incorporated additional semi-stable levels of subgrouping. Our data support the hypothesis that chimpanzee males combine social organization strategies, and that doing so may allow them, like humans, to mitigate individual costs at larger community sizes.

Agreement between oscillometric and direct blood pressure measurements in anesthetized captive chimpanzees (Pan troglodytes)

OBJECTIVE To evaluate the level of agreement (LOA) between direct and oscillometric blood pressure (BP) measurements and the ability of oscillometric measurements to accurately detect hypotension in anesthetized chimpanzees (Pan troglodytes). ANIMALS 8 captive, adult chimpanzees. PROCEDURES During prescheduled annual examinations, each chimpanzee underwent general anesthesia and patient monitoring for their examination, echocardiography for a concurrent study, and measurement of direct BP with the use of tibial artery catheterization and oscillometry with the use of a cuff placed around a brachium and a cuff placed around the second digit of the contralateral forelimb for the present study. Bland-Altman plots were generated to compare results for direct and oscillometric BP measurements. Mean bias and 95% LOAs were calculated for oscillometric measurements of systolic arterial pressure (SAP), diastolic arterial pressure (DAP), and mean arterial pressure (MAP) for each cuff site. Sensitivity and specificity in detecting hypotension were also determined for each cuff site. RESULTS There were 74 paired direct and brachial oscillometric measurements of each, SAP, MAP, and DAP and 66 paired direct and digit oscillometric measurements of each, SAP, MAP, and DAP. Only brachial oscillometric measurements of MAP had adequate sensitivity (78%) and specificity (95%) to accurately detect hypotension, and this technique also had the least mean bias (0.8 mm Hg; 95% LOA, –29 to 31 mm Hg). CONCLUSIONS AND CLINICAL RELEVANCE Results indicated that brachial oscillometric measurement of MAP provided reasonable agreement with tibial arterial direct MAP measurement and performed well in diagnosing hypotension in anesthetized chimpanzees.

Behavioural diversity as a potential welfare indicator for professionally managed chimpanzees (Pan troglodytes): Exploring variations in calculating diversity using species-specific behaviours

Behavioural diversity may serve as a positive indicator of animal welfare that can be applied in long-term monitoring schemes in managed settings (eg zoos, laboratories, farms). Behavioural diversity is often higher when animals live in stimulating environments and experience positive events. Unfortunately, welfare researchers have not adopted consistent, standardised approaches to measuring behavioural diversity. The goal of this exploratory study was to utilise data from 41 adult chimpanzees (Pan troglodytes) housed across 16 zoological institutions to examine various models of Shannon's Diversity Index. Specifically, we investigated the impact of: combining versus splitting behaviours, including only positive behaviours, including human interaction, and considering recipient behaviours. We evaluate how the inclusion or exclusion of different behaviours impacts the relationship of behavioural diversity with: (i) concentrations of faecal glucocorticoid metabolites (GCM), a common indicator of adrenal activity; (ii) concentrations of immunoglobulin-A (IgA), an indicator of immune function and potential indicator of positive welfare; and (iii) stereotypic behaviour, a validated indicator of poor welfare. Most indices had significant negative relationships with faecal GCM. Animals that express a variety of behaviours from their species-typical repertoire have lower average faecal GCM concentrations and are likely experiencing better welfare. We did not find significant relationships between the behavioural diversity indices and IgA concentrations. Two indices were inversely associated with stereotypic behaviour. Our findings provide additional support for using Shannon's Diversity Index to calculate behavioural diversity as a robust, valid measure of positive welfare. However, future publications must justify the process for including or excluding behaviours from calculations.

Search asymmetries for threatening faces in chimpanzees (Pan troglodytes)

For primates, the ability to efficiently detect threatening faces is highly adaptive, however, it is not clear exactly how faces are detected. This study investigated whether chimpanzees show search asymmetries for conspecific threatening faces featuring scream and bared teeth expressions. Five adult female chimpanzees participated in a series of touchscreen matching-to-sample visual search tasks. In Experiment 1, search advantages for scream versus neutral targets, and scream versus bared teeth targets were found. A serial search strategy indicated greater difficulty in disengaging attention from scream versus neutral distractors. In Experiments 2a and 2b, search advantages for scream versus neutral targets remained when the mouth was darkened, suggesting that the brightness contrast of the mouth was not critical for the efficient detection of scream targets. In Experiments 3a and 3b, search advantages for inverted scream versus neutral targets disappeared, indicating configural processing. Together, exclusion of the brightness contrast of the mouth as a low-level perceptual confound, and evidence of configural processing, suggested the scream faces may have been perceived as threatening. However, the search advantage for scream faces is most likely explained by the presence of teeth, independently of threat. The study provides further support that an attentional bias towards threatening faces is a homologous trait, which can be traced back to at least the last common ancestor of Old-World monkeys and apes.

Overlooked evidence for semantic compositionality and signal reduction in wild chimpanzees (Pan troglodytes)

Recent discoveries of semantic compositionality in Japanese tits have enlivened the discussions on the presence of this phenomenon in wild animal communication. Data on semantic compositionality in wild apes are lacking, even though language experiments with captive apes have demonstrated they are capable of semantic compositionality. In this paper, I revisit the study by Boesch (Hum. Evol. 6:81–89, 1991) who investigated drumming sequences by an alpha male in a chimpanzee (Pan troglodytes) community in the Taï National Park, Côte d’Ivoire. A reanalysis of the data reveals that the alpha male produced semantically compositional combined messages of travel direction change and resting period initiation. Unlike the Japanese tits, the elements of the compositional expression were not simply juxtaposed but displayed structural reduction, while one of the two elements in the expression coded the meanings of both elements. These processes show relative resemblance to blending and fusion in human languages. Also unlike the tits, the elements of the compositional expression did not have a fixed order, although there was a fixed distribution of drumming events across the trees used for drumming. Because the elements of the expression appear to carry verb-like meanings, the compositional expression also resembles simple verb-verb constructions and short paratactic combinations of two clauses found across languages. In conclusion, the reanalysis suggests that semantic compositionality and phenomena resembling paratactic combinations of two clauses might have been present in the communication of the last common ancestor of chimpanzees and humans, not necessarily in the vocal modality.