Sex differences in the behaviour of immature captive lowland gorillas

Studies of the behaviour of 26 (12 males and 14 females) captive infant and juvenile lowland gorillas showed clear sex differences. Females showed greater interest in young infants and were more active in nest building as well as in solitary and social grooming. Males were more active in locomotive, dominance, and aggressive behaviour and in social play. Hand-rearing further increased aggression. Males were more aggressive when they lived with only one partner, and they rose in rank even above older females, a pattern that has not been observed in naturally reared gorillas.     

Exchange of objects between humans and captive western lowland gorillas

An experiment was carried out to assess the ability of captive western lowland gorillas (Gorilla g. gorilla) to exchange objects with a human partner. Before the exchange itself, individuals had to learn to give an item to the experimenter. Four individuals out of nine performed active giving. After that familiarization with giving behaviour, two kinds of exchanges, of increasing complexity, were tested with gorillas. Simple exchange (give C to obtain D) was the first procedure and six gorillas out of nine succeeded. Double exchange (give B to receive C, then give back C to obtain D) was performed only by three individuals and triple exchange (give A to receive B, give back B to receive C, then give back C to obtain D) was performed only by two individuals. Different types of errors were scored, as well as social constraints on learning. Results showed that (1) naive gorillas are able to get a reward (D) in an experimental exchange situation; and (2) the complex rules of multiple successive exchanges were not easily understood by gorillas.     

Postconflict behavior in captive western lowland gorillas (Gorilla gorilla gorilla)

Postconflict (PC) behaviors, including reconciliation and consolation, have been observed in many primate and several nonprimate species. Using the PC-matched control (MC) method, PC behavior was examined in two groups (n=13) of captive western lowland gorillas, a species for which no conflict resolution data have been published. Analyses of 223 conflicts showed significantly more affiliation between former opponents after a conflict when compared to control periods, indicating reconciliation. Results also showed significantly more affiliation between the victim and a third-party after a conflict, indicating consolation. Both solicited and unsolicited consolation were observed. The majority of the affiliative interactions observed for both reconciliation and consolation were social proximity, which suggests that unlike most nonhuman primates, proximity, rather than physical contact, may be the main mechanism for resolving conflicts in western lowland gorillas. PC behavior was not uniform throughout the groups, but rather varied according to dyad type.     

Adult–adult play in captive lowland gorillas (Gorilla gorilla gorilla)

Primates - Among African great apes, play is virtually absent between adult lowland gorillas (Gorilla gorilla gorilla). Here, we report an extremely rare case of adult–adult play observed in...     

Food transfers in immature wild western lowland gorillas (Gorilla gorilla gorilla)

The transfer of food items between individuals has been described in primates as serving an informative purpose in addition to supplementing the diet of immature individuals. This behaviour has yet to be described in western lowland gorillas (Gorilla gorilla gorilla), and results are presented here of observations of food transfers in immature gorillas at Mbeli Bai, Republic of Congo. The frequency of food transfers decreased with increasing immature age, while the frequency of independent feeding and processing of food increased. Transfers between mothers and infants were the most frequent, with infants attempting to take items from the mother. These attempts were not always successful and the item was relinquished on less than 50% of attempts. Mothers also took items from their offspring. The results point to the functional significance of food transfers in western lowland gorillas being informational. In a bai environment, where one species forms the majority of a visiting gorilla’s diet despite other species being available, the initiation of food transfers by immatures is proposed to serve the purpose of familiarising them with which species, and which parts of those species, may be eaten.     

Using behavior to determine immature life‐stages in captive western gorillas

Ontogenic development is divided into infant, juvenile, adolescent and adult life‐stages. Although the developmental trajectory of an individual is a flexible entity, which differs within species, environment and sex, life‐stage classifications are generally structured, age‐based systems. This invariably leads to rigidity within a dynamic system and consequently hampers our understanding of primate life history strategies. We propose that life‐stage classifications should be quantitative, flexible entities, which use a reliable measurement of development. Here, we provide a methodological example where placement into a life‐stage is based upon behavioral variance between other similar‐aged individuals. Behavioral data were collected from 12 male (3–11 years old) and 9 female (3–8 years old) captive immature western gorillas (Gorilla gorilla gorilla) housed in five family groups, using continuous focal sampling; 900 hr of data were collected over 131 days. Data were applied to four published life‐stage classifications for mountain gorillas (Gorilla beringei beringei), which showed variable ability to determine life‐stage in western gorillas. A new life‐stage classification (Hutchinson & Fletcher) was proposed specifically for western gorillas, whereby multiple co‐varying behavior provided a robust measure of linear development across immaturity. Each life‐stage was found to be a distinct ontogenic phase and the classification discriminated life‐stage with a high level of accuracy. Using the Hutchinson & Fletcher classification we provide evidence for disparity in developmental trajectories between the sexes from the juvenile period onwards. To expand the understanding of primate life histories, we propose that flexible classifications should be used to enable comparison of allometric life history traits within and between species, from birth onwards. Am. J. Primatol. 72:492–501, 2010. © 2010 Wiley‐Liss, Inc.     

Long‐term proximity relationships in a captive social group of western lowland gorillas (Gorilla gorilla gorilla)

To visualize long‐term social relationships among 12 gorillas in a captive breeding group at the San Diego Wild Animal Park, San Diego, CA, the multidimensional scaling (MDS) procedure was applied to proximity (within 5 m) values collected in five different periods over 2.5 years. The resulting two‐dimensional representations clearly show that in samples taken during this 2.5‐year period, the silverback male was surrounded by five adult females, while three infant or young juvenile females, gathered in a body, remained between the silverback male and adult females. Some adult females maintained proximity to the silverback male more frequently than other adult females throughout the five periods. Unlike mountain gorillas in the wild, females with dependent offspring did not tend to stay near the silverback male more frequently than other females, and related females did not tend to spend more time near each other than non‐related females. Three older juvenile or young adult males were plotted the furthest from the silverback male, with gradual changes with increasing age. The usefulness of the MDS procedure, which can be used to visualize easily and clearly the social relationships among individuals, is discussed from the viewpoint of the management of breeding groups of captive gorillas. Zoo Biol 20:197–209, 2001. © 2001 Wiley‐Liss, Inc.     

Spatial memory and foraging competition in captive western lowland gorillas (Gorilla gorilla gorilla)

Spatial memory and foraging competition were investigated in three mother/offspring pairs of western lowland gorillas,Gorilla gorilla gorilla, using a naturalistic foraging task at the Toronto Zoo. Sixteen permanent food sites were placed throughout the animals’ enclosures. All of the sites were baited and a pair of animals was free to visit the sites and collect the food. Five of the subjects collected the food with accuracy better than chance. Most of the subjects visited the sites using a pattern, and for half the subjects this was one of adjacency. The high accuracy of five of the subjects and the lack of a consistent adjacency pattern suggest that the animals did in fact use spatial memory. Furthermore, the gorillas tended to avoid visiting food sites that had been previously depleted by their partner. They also appeared to split their search of the enclosures, each visiting only a proportion of the food sites. This indicated that the animals were competing and altering their foraging behaviour based on the behaviour of their partner. Therefore, accuracy was recalculated to take this into account. When the sites depleted by either animal in a pair during a given trial were worked into the accuracy calculations for individual animals, three of the animals still maintained accuracy above chance. This suggests that the animals were not only able to remember which sites they had depleted, but those sites depleted by their foraging partner as well.     

The development of infant play in a captive group of lowland gorillas (Gorilla gorilla gorilla)

Three infant gorillas born at the Yerkes Regional Primate Research Center of Emory University were studied during their first year and one-half of life for various aspects of behavior, including solitary and social play, using both focal animal and one-zero sampling. The appearance of play in these infants followed a developmental trend, beginning with mother-infant play, and followed by solitary play, and then social play among peers. Mothers were initially very vigilant, repeatedly interrupting infant peer play bouts, this relaxing toward the end of the first year. Evidence for a possible sex difference in active social play is presented.     

Effects of combination birth control on estrous behavior in captive western lowland gorillas, Gorilla gorilla gorilla

Combination birth control pills (CBC) are one of the most common birth control methods used for western lowland gorillas (Gorilla gorilla gorilla) housed in zoos. Since zoos are interested in maintaining as many natural behaviors as possible, it is important to know how contraception may affect social and sexual interactions among group members. Although some data are available regarding the influence of the pill on sexual behavior in human females, no data are available on its effects on gorilla estrous behavior. We examined temporal trends of estrous, aggressive, affiliative, and activity budget data in four females on CBC at the Lincoln Park Zoo in Chicago, IL. Behavioral data were collected using point sampling, all-occurrence records, and one-zero sampling. Estrous behavior occurred in less than 1% of observations. Using all-occurrence and one-zero sampling, estrous behavior occurred more frequently in week one of the cycle than any other week. The focal females exhibited affiliative, aggressive, and activity budget data evenly across their cycles. There were also no temporal trends in proximity to the silverback. Females varied by the types of estrous behavior they exhibited. We give a hormonal explanation for the prevalence of estrous behaviors in week one, and recommendations for effective behavioral sampling of gorilla estrous behavior.     

A training program for noninvasive semen collection in captive western lowland gorillas (Gorilla gorilla gorilla)

Staff at Omaha's Henry Doorly Zoo devised a training program for noninvasive semen collection from three male gorillas. Trained behaviors, shaped using food rewards and verbal praise, enabled the trainer to perform a cursory physical examination and collect >270 semen samples during the study period. Ejaculate volume, sperm motility, and progressive forward motion were recorded. Two of the gorillas regularly produced semen samples that were cryopreserved. Frozen semen from one gorilla was successfully used in an assisted reproductive procedure. The training program may be used as a guideline for other institutions to create similar programs. Zoo Biol 17:143–151, 1998. © 1998 Wiley-Liss, Inc.     

Paternity determination in captive lowland gorillas and orangutans and wild mountain gorillas by microsatellite analysis

Paternity exclusion studies provide useful information for testing certain theories of behavioral ecology and for the management and conservation of both wild and captive populations of endangered species. This study used eight human nuclear microsatellite loci, in the absence of species-specific PCR primers, to genetically identify the sires of 12 captive lowland gorillas (Gorilla gorilla gorilla) and 2 captive orangutans (Pongo pygmaeus pygmaeus andPongo p. abelii). Parentage assignments were confirmed by excluding all except a single potential sire for each offspring with the least two loci. Sire-offspring relationships were verified in 12 of the 14 cases, and reassigned in the case of two gorilla offspring. The orangutan paternity typing was supplemented by DNA fingerprinting. Additionally, five of the eight microsatellite loci, in conjunction with behavioral data, were used for a non-exhaustive set of paternity exclusions for five wild mountain gorillas (Gorilla g. beringei). The eight loci described in this study should be useful additions to the tools available for the study of genetics in the great apes.     

Behavioral effects of changing group membership among captive lowland gorillas

Group membership was modified among 4 gorilla groups at Zoo Atlanta in an attempt to increase compatibility and sexual behavior. The modifications ultimately moved 7 adult females among the groups. The moves were made by individually introducing the new group members to the old members, resulting in 18 total introductions. Several individual and social behaviors differed in the two conditions. The most obvious change was a temporary increase in aggressive display behaviors, as well as an increase in contact and social examination, in the postmovement condition. Additionally, the individual behaviors of travelling and eating increased after the introductions, while resting decreased. Other self-maintenance behaviors did not change. Finally, there was an increase in time mothers and offspring spent closer to one another following the moves. Previous research and anecdotal evidence suggest that gorillas respond to changes in group membership with aggression; these data support and extend those data to different age/sex groups, postmovement trends in behavior, and other aspects of gorilla behavior. © 1996 Wiley-Liss, Inc.     

Baby-transfer and other interactions between its mother and grandmother in a captive social group of lowland gorillas

This report describes the responses of an experienced gorilla mother to inappropriate maternal behavior displayed by her young adult daughter toward a newborn baby and repeated acts of baby-transfer between these two females in a captive social group of lowland gorillas (Gorilla g. gorilla). The quality of infant care by the young adult daughter clearly improved during the first 4 days after birth, and this improvement was at least partly based on her mothers encouragement. Thus, the mothers activities can be considered scaffolding for her daughter with regard to maternal infant care.     

Body mass in lowland gorillas: a quantitative analysis

Body proportions and tissue composition (e.g., relative contributions of muscle, skin, bone, and adipose to total body mass) were determined through dissection of four adult captive lowland gorillas. The relative contribution of bone varies little among the four animals (10.2-13.4%) despite considerable range in body weights (99.5-211 kg). In tissue composition, three animals have on average 37.3% muscle relative to body mass. Maximum estimates of body fat range between 19.4-44%. Differences in age, sex, and life history events partially explain the observed variation in body proportions and tissue composition among the four animals. Although gorillas are considered extremely sexually dimorphic in body weight and canine size, differences in tissue are not as dramatic as body mass differences suggest. This study found sex differences mostly in the upper body; males have relatively heavier forelimbs, including heavier deltoid, trunk-binding, and deep back muscles compared to the younger female. The old, obese female had one half the muscle tissue of the other three animals (16% vs. 37.3%), and twice the body fat (44%); forelimbs and upper body musculature were relatively well-developed to compensate for the restricted hip-joint movement due to arthritis. Data on the variation in tissue composition and body proportions in gorillas provide a basis for comparison with other hominoids, including humans. For example, compared to highly dimorphic orangutans, gorillas have more muscle, less adipose tissue, lighter forelimbs and heavier hindlimbs. Such analyses complement studies of the skeleton and contribute to our understanding of human evolution and adaptation.     

Deception and hiding in captive lowland gorillas (Gorilla gorilla gorilla)

Deception and hiding are described for a group of captive lowland gorillas comprising two adult males, one female, and her infant. The males repeatedly interacted with the infant, and their interest was repeatedly thwarted by the mother. As in other social situations in which one animal controls access to an object desired by another, deceit was used to thwart the other's control. Either one or both males at times concealed their actions toward the infant, inhibited their attention to the infant, created a neutral image to get the infant, or interacted with the infant to tease the mother.     

Ecological correlates of space use patterns in wild western lowland gorillas

The distribution of resources is a crucial determinant of animals' space use (e.g., daily travel distance, monthly home range size, and revisitation patterns). We examined how variation in ecological parameters affected variability in space use patterns of western lowland gorillas, Gorilla gorilla gorilla. They are an interesting species for investigating this topic because key components of their diet are nonfruit items (herbaceous vegetation and tree leaves) that occur at low density and are sparsely distributed, and fruits, which show high spatiotemporal variation in availability. We estimated how availability of nonfruit foods and fruit, frugivory (proportion of feeding time consuming fruit), and swamps in areas used by the gorillas influenced daily travel distance, monthly home range size, and revisit frequency to grid cells in the home range of one habituated gorilla group in Loango National Park, Gabon. Using location data from 2015 to 2018, we found that the gorillas decreased their daily travel distance as both the density of nonfruit foods and the proportion of swamps in areas used increased. Daily travel distances were shorter when both frugivory and availability of fruit were higher, yet, daily travel distances were longer when availability of fruit was low but frugivory was still high. Furthermore, monthly home range size increased as frugivory increased and monthly revisit frequencies to an area increased as fruit availability of an area increased. In conclusion, the availability of both nonfruit and fruit influenced the gorillas' space use patterns. Gorillas decreased foraging effort when food availability was high but were willing to incur increasing foraging costs to feed on fruit when availability was low. This study highlights how animals have to adjust their space use with changing resource availability and it emphasizes the value of examining multiple parameters of space use.     

Plant selection for nest building by western lowland gorillas in Cameroon

We examined 834 nests built by western lowland gorillas in Cameroon between July 2008 and July 2011 to identify the plant species used in their construction. Preference for each plant species for nesting was assessed using a ‘preference index’ calculated by combining information on the occurrence of each species in the forest and in the nests. Forty-six plant species representing about 15 % of the total number of species in the forest and 26 % of species used for nest building were frequently used by gorillas. Preference levels significantly varied among these species. Nests were mostly built with herbs of the families Marantaceae and Zingiberaceae and woody species such as Manniophyton fulvum (liana) and Alchornea floribunda (shrub). As observed in other gorilla populations, suitability for nest building and availability of gorilla food in stems were the likely determinants of plant selection. The total number of species used per nest ranged from 1 to 11, with an average of 4.9. This is high compared to other sites, emphasizing variability in the availability of nest building materials and habitat differences across the range of the western gorilla. Seasonal changes in the use of different habitat types for nesting did not appear to influence plant use for nest building as there was little variation in plant selection across seasons or the composition of nests. Our findings suggest that gorillas non-randomly select plant species to build nests, and use a particular set of species combined at varying proportions, with no clear seasonal or spatial patterns.