A novel attention-getting vocalization in zoo-housed western gorillas.

As a critical aspect of language, vocal learning is extremely rare in animals, having only been described in a few distantly related species. New evidence, however, extends vocal learning/innovation to the primate order, with zoo-housed chimpanzees and orangutans producing novel vocal signals to attract the attention of familiar human caregivers. If the ability to produce novel vocalizations as a means of navigating evolutionarily novel circumstances spans the Hominidae family, then we can expect to find evidence for it in the family's third genus, Gorilla. To explore this possibility, we conduct an experiment with eight gorillas from Zoo Atlanta to examine whether they use species-atypical vocalizations to get the attention of humans across three different conditions: just a human, just food, or a human holding food. Additionally, we survey gorilla keepers from other AZA-member zoos to compile a list of common attention-getting signals used by the gorillas in their care. Our experiment results indicated that Zoo Atlanta gorillas vocalized most often during the human-food condition, with the most frequently used vocal signal being a species-atypical sound somewhere between a sneeze and a cough (n = 28). This previously undescribed sound is acoustically different from other calls commonly produced during feeding (i.e., single grunts and food-associated calls). Our survey and analyses of recordings from other zoos confirmed that this novel attention-getting sound is not unique to Zoo Atlanta, although further work should be done to better determine the extent and patterns of transmission and/or potential independent innovation of this sound across captive gorilla populations. These findings represent one of the few pieces of evidence of spontaneous novel vocal production in non-enculturated individuals of this species, supporting the inclusion of great apes as moderate vocal learners and perhaps demonstrating an evolutionary function to a flexible vocal repertoire.

Survey of current group demographics and management practices of bachelor groups of western lowland gorillas (Gorilla gorilla gorilla) across North America.

Recommendations for the successful management and maintenance of bachelor groups of western lowland gorillas (Gorilla gorilla gorilla) in zoological settings have been an increasingly prevalent focus within the zoological community. Over the past two decades, studies have examined the impact of various environmental factors on the stability of bachelor groups, proposed management strategies for bachelor groups, and compared behavioral trends between bachelor and mixed-sex groups. These studies have clearly demonstrated that bachelor groups are complex social units that require specialized management approaches. In this study, we aimed to assess the extent to which bachelor group management across North American zoos accredited by the Association of Zoos and Aquariums aligns with established recommendations. We distributed a comprehensive survey broadly encompassing habitat and housing, aggression and wounding, group demographics, feeding, and training to the 22 zoos housing bachelor groups as of 2019. We received completed surveys from 19 zoos, representing a total of 21 social units and 59 individual gorillas. We used descriptive statistics to represent the range of current management strategies across the surveyed population and ANOVAs to assess significant variation in key demographic variables. Our results demonstrate that a majority of zoos have adopted the best practices for the formation of social groups established by Stoinski et al. in 2004. However, there is much less standardization across zoos in protocols surrounding training and feeding. Additionally, important variables in the assessment of wounding, such as time of day and location, are often unknown or not observed by animal care professionals. We highlight these two areas as being of particular focus in developing and adhering to consistent protocols across institutions.

Who is there? Captive western gorillas distinguish human voices based on familiarity and nature of previous interactions.

The ability to recognize conspecifics by their acoustic signals is of crucial importance to social animals, especially where visibility is limited, because it allows for discrimination between familiar and unfamiliar individuals and facilitates associations with and the avoidance of particular conspecifics. Animals may also benefit from an ability to recognize and use the information coded into the auditory signals of other species. Companion species such as dogs, cats, and horses are able to discriminate between familiar and unfamiliar human voices; however, whether this ability is widespread across vertebrates is still unknown. Using playback experiments, we tested whether western gorillas living at Zoo Atlanta were able to discriminate between the voices of subgroups of people: i.e., unfamiliar individuals, familiar individuals with whom the gorillas had positive interactions, and familiar individuals with whom they had negative interactions. Gorillas responded significantly more often (longer gazing duration, higher gazing frequency, shorter latency, and larger number of distress behaviors) to the voices of unfamiliar and familiar-negative individuals than to those of familiar-positive individuals, indicating that they recognized the voices of subgroup of people based on familiarity and possibly the nature of the relationship with them. Future studies should determine whether this is also the case in the wild, where interspecific associations with humans are less intense than they are in captive settings.

Western lowland gorillas (Gorilla gorilla gorilla) do not show an aversion to inequity in a token exchange task.

Although individuals in some species refuse foods they normally accept if their partner receives a more preferred one, this is not true across all species. The cooperation hypothesis proposes that this species-level variability evolved because inequity aversion is a mechanism to identify situations in which cooperation is not paying off, and that species regularly observed cooperating should be more likely to be averse to inequity. To rule out other potential explanations of inequity aversion, we need to test the converse as well: species rarely observed cooperating, especially those phylogenetically close to more cooperative species, should be less likely to be inequity averse. To this end, we tested eight zoo-housed Western lowland gorillas (Gorilla gorilla gorilla) on a token exchange task in which subjects received either the same food reward or a less-preferred reward for the same or more effort than their partner, recording both refusals to participate in the exchange and refusals to accept the reward. Supporting the cooperation hypothesis, even with procedural differences across sessions, gorillas were significantly more likely to refuse in all conditions in which they received a low-value food reward after completing an exchange, regardless of what their partner received, suggesting that gorillas were not inequity averse, but instead would not work for a low-value reward. Additionally, gorillas were more likely to refuse later in the session; while the pattern of refusals remained unchanged after accounting for this, this suggests that species should be tested on as many trials as is practical.

Comparative performance of orangutans (Pongo spp.), gorillas (Gorilla gorilla gorilla), and drills (Mandrillus leucophaeus), in an ephemeral foraging task.

A goal of the comparative approach is to test a variety of species on the same task. Here, we examined whether the factors that helped capuchin monkeys improve their performance in a dichotomous choice task would generalize to three other primate species: orangutans, gorillas, and drill monkeys. In this task, subjects have access to two options, each resulting in an identical food, but one (the ephemeral option) is only available if it is chosen first, whereas the other one (the permanent option) is always available. Therefore, the food-maximizing solution is to choose the ephemeral option first, followed by the permanent option for an additional reward. On the original version (plate task), the options were discriminated by the color and pattern of the plates holding the food, while on two subsequent versions we used altered cues that we predicted would improve performance: (1) the color of the foods themselves (color task), which we hypothesized was relevant to primates, who choose foods rather than substrates on which foods are found when foraging, and (2) patterned cups covering the foods (cup task), which we hypothesized would help primates avoid the prepotent response associated with visible food. Like capuchins, all three species initially failed to solve the plate task. However, while orangutans improved their performance from the plate to the color task, they did not for the cup task, and only a few gorillas and no drills succeeded in either task. Unfortunately, our ability to interpret these data was obscured by differences in the subjects' level of experience with cognitive testing and practical constraints that precluded the use of completely identical procedures across species. Nonetheless, we consider what these results can tell us, and discuss the value of conducting studies across multiple sites despite unavoidable differences.

Endowment effects in gorillas (Gorilla gorilla).

Reports of endowment effects in nonhuman primates have received considerable attention in the comparative literature in recent years. However, little is known about the mechanisms underlying these effects. Continuing to explore endowment effects across different species of primate may reveal subtle differences in behavior that can help formulate specific hypotheses about the relevant mechanisms and the social and ecological factors that have shaped them. In this study, we use a paradigm that has previously been used to test chimpanzees (Pan troglodytes) and orangutans (Pongo spp.) to explore whether western lowland gorillas (Gorilla gorilla) exhibit comparable endowment effects. We find that gorillas exhibit endowment effects when in possession of food, but not nonfood, items, and that they show a statistically stronger effect than chimpanzees but not orangutans. These findings are consistent with the hypothesis that mechanisms for endowment effects in primates may be related to inhibitory control or risk aversion.