Preference for structural environmental features in captive lowland gorillas (Gorilla gorilla gorilla)

Increased interest in the behavioral effects of captive environments and advances in zoo design have resulted in a proliferation of “naturalistic” animal habitats. Although scientists have demonstrated that such exhibits have some positive effects on behavior, very little is known about the complex relationships between these specialized environments and behavior. This study sought to determine preferences for structural environmental features, such as slope and trees, exhibited by captive lowland gorillas (Gorilla gorilla gorilla). An average of 496 location scans were collected on each of 23 gorillas, housed in 6 groups. The results indicate that enclosure usage was significantly associated with the features of slope, nearness to holding facility, and presence of environmental components additional to substrate (e.g., tree or rock). In particular, areas with flat ground that were near the holding building and that included at least one component additional to substrate were preferred by all animals. © 1993 Wiley-Liss, Inc.     

Gorilla Behavior Index revisited: Age,housing and behavior

While personality and temperament assessments are becoming more common in animal science as a means for predicting behavior patterns, they are virtually unstudied in exotic animals outside of chimpanzees. We assessed behavioral profiles of 119 male gorillas using the Gorilla Behavior Index (GBI) and paired this assessment with behavior data on 25 male gorillas from our ongoing analysis of multi-male gorilla groups in North America. This comparison was made to determine if the GBI could be predictive of behavior patterns, with the hope that this could eventually be utilized to aid in the management of multi-male gorilla groups. With few exceptions, GBI factor scores were not related to age of subjects or their current housing condition. Individuals scoring high on the Extroverted factor were more likely to initiate and receive affiliative behaviors, but they were also more likely to initiate contact aggression. Those gorillas who scored high on the Dominant factor were more likely to initiate displacements and less likely to receive them. Gorillas rated as more Fearful were less likely to initiate displacements and those rated as less Understanding were more likely to initiate non-contact aggression. Two facts are evident from these data: (1) further work needs to be done to verify that the GBI is indeed assessing traits and not current behavioral states and (2) any relationships with behavior frequency are relatively weak. In addition to further exploration of the GBI we recommend other assessments to compliment the GBI to establish stronger links between subjective assessments and objective behavior patterns. If established, these short-term assessments may be combined with other historical information on these animals, such as age, rearing history, the number of animals in the group, and the design of the exhibit, to aid in forming and maintaining multi-male gorilla groups in captivity.     

The cryptic genetic structure of the North American captive gorilla population

Western lowland gorillas (Gorilla gorilla gorilla) were imported from across their geographical range to North American zoos from the late 1800s through 1974. The majority of these gorillas were imported with little or no information regarding their original provenance and no information on their genetic relatedness. Here, we analyze 32 microsatellite loci in 144 individuals using a Bayesian clustering method to delineate clusters of individuals among a sample of founders of the captive North American zoo gorilla collection. We infer that the majority of North American zoo founders sampled are distributed into two distinct clusters, and that some individuals are of admixed ancestry. This new information regarding the existence of ancestral genetic population structure in the North American zoo population lays the groundwork for enhanced efforts to conserve the evolutionary units of the western lowland gorilla gene pool. Our data also show that the genetic diversity estimates in the founder population were comparable to those in wild gorilla populations (Mondika and Cross River), and that pairwise relatedness among the founders is no different from that expected for a random mating population. However, the relatively high level of relatedness (R = 0.54) we discovered in a pair of known breeding pairs reveals the need for incorporating genetic relatedness estimates in the captive management of western lowland gorillas.     

The birth of a lowland gorilla in captivity

A 7 year-old captive-born mother-reared gorilla gave birth to a healthy infant after being in labor for 27 min. The young female was a wholly competent mother although she had never observed gorilla mothers or interacted with gorilla infants. This reinforces previous evidence, still often ignored by zoo officials, that gorillas do not have to learn maternal behavior.     

Factors influencing the formation and maintenance of all‐male gorilla groups in captivity

The social housing of males is of particular importance for captive managers of polygynous species. In this work, we highlight nine areas we believe are important to the successful formation and maintenance of all‐male gorilla groups. Although more data are still needed to complete our understanding of these groups, the general picture that emerges is that all‐male groups can be a functional social unit for male gorillas, the longevity of which might be improved by careful planning. It appears that the best time to form all‐male groups is when the animals are still immature, as males in this age class have been observed to form some of the most stable groups. Efforts should be made to diversify the groups in terms of rearing history, particularly with respect to hand‐reared males, and to limit the amount of hand rearing experienced by males. Groups probably should contain no more than three or four adult males. Exhibits should be designed to provide visual barriers, refuges for subordinate animals, the means to separate individuals, and possibly the means to isolate all‐male groups from mixed‐sex groups. Additionally, because it is likely that some males will have to be removed from all‐male groups, zoos need to design facilities that provide state‐of‐the‐art housing for solitary individuals. It is hoped that by identifying bachelor males early in life and establishing cooperative management plans across institutions, zoos will be able to provide beneficial, long‐term social situations for all male gorillas in captivity. Zoo Biol 23:189–203, 2004. © 2004 Wiley‐Liss, Inc.     

Group differences in captive gorillas’ reaction to large crowds

The impact of visitor crowd size on captive animal behaviour can be difficult to interpret in many institutions due to the lack of variation in crowd size over short periods of time. In attempts to compare greater variation in crowd size, animal behaviour is often compared over conditions that create additional confounds, such as day of week or season. Fluctuations in attendance over the holiday season at Disney's Animal Kingdom^® Theme Park provided an opportunity to examine the impact of variation in crowd size on gorilla behaviour without the confounds found in many other studies. Ten western lowland gorillas (Gorilla g. gorilla) in a bachelor group (n = 4) and a family group (n = 6) were observed over a period of 8 weeks in late 2005. Observations were classified into Large Crowd and Small Crowd days and a repeated-measures ANOVA procedure was conducted to determine the effects of crowd size and social group on gorilla behaviour patterns. Few overall differences were found in behaviour during the Large Crowd and Small Crowd conditions. However, similar to previous research, gorillas were less visible during the Large Crowd condition (F_1,8 = 14.15, P = 0.01). There was also an interaction of crowd size and group (F_1,8 = 7.58, P = 0.01), indicating the bachelor group of gorillas was more aggressive during the Large Crowd condition, whereas the family group showed no increase in aggression with large crowds. These results indicate the importance of providing complex enclosures with visual barriers to allow animals to move away from large crowds if they choose. Future research should focus on the individual characteristics of animals and enclosures that may contribute to behavioural differences in visitor–animal interaction research, as well as the proximate cues associated with behaviour change when exposed to large crowds.     

Behavioral responses of silverback gorillas (Gorilla gorilla gorilla) to videos

This study examined the impact of video presentations on the behavior of 4 silverback, western lowland gorillas (Gorilla gorilla gorilla). On each of 5 occasions, gorillas viewed 6 types of videos (blue screen, humans, an all-male or mixed-sex group engaged in low activity, and an all-male or mixed-sex group engaged in agonistic behavior). The study recorded behavioral responses and watching rates. All gorillas preferred dynamic over static videos; 3 watched videos depicting gorillas significantly more than those depicting humans. Among the gorilla videos, the gorillas clearly preferred watching the mixed-sex group engaged in agonistic behavior; yet, this did not lead to an increase in aggression or behavior indicating agitation. Further, habituation to videos depicting gorillas did not occur. This supports the effectiveness of this form of enrichment, particularly for a nonhuman animal needing to be separated temporarily due to illness, shipment quarantine, social restructuring, or exhibit modification.     

Group size and structure in western lowland gorillas (Gorilla gorilla gorilla) at Mbeli Bai, Republic of Congo

This paper describes the size and structure of western lowland gorilla groups visiting Mbeli Bai, a swampy forest clearing in northern Congo. An observation platform at the edge of the clearing was used over a 3-yr period, with gorillas visible for 1,681 hr. Data are presented on 14 groups and seven solitary males. Mean group size (excluding solitary males) was 8.4 +/- SD 4.3, and did not differ significantly from most other gorilla studies. All groups at Mbeli contained only one fully mature male and did not show evidence of fission-fusion or regular subgrouping. All emigrating males that remained in the population became solitary, and "bachelor" groups were not observed. Methodological issues are raised, including the classification of multi-male groups, and the demographic profiles of other populations are discussed in the light of results from Mbeli.     

‘Separating the wheat from the chaff’: A novel food processing technique in captive gorillas (Gorilla g. gorilla)

Observation of a novel food processing technique is reported for captive zoo gorillas (Gorilla g. gorilla). It is similar in function to that of Japanese macaques' wheat placer mining behaviour and consists of puffing/blowing air with the mouth onto a mixture of oat grains and chaff in order to separate out the oat grains. Three females in two of four groups regularly use this behaviour. Other individuals in these groups or individuals of the two other groups in the same zoo do not use it. However, a very similar behaviour has been observed in three other individuals in a gorilla group of another zoo. The existence of this technique in spatially separated groups implies that multiple individuals have invented it for themselves. The possible role of social transmission is still to be investigated.     

Behavioral Responses of Silverback Gorillas (Gorilla gorilla gorilla) to Videos

This study examined the impact of video presentations on the behavior of 4 silverback, western lowland gorillas (Gorilla gorilla gorilla). On each of 5 occasions, gorillas viewed 6 types of videos (blue screen, humans, an all-male or mixed-sex group engaged in low activity, and an all-male or mixed-sex group engaged in agonistic behavior). The study recorded behavioral responses and watching rates. All gorillas preferred dynamic over static videos; 3 watched videos depicting gorillas significantly more than those depicting humans. Among the gorilla videos, the gorillas clearly preferred watching the mixed-sex group engaged in agonistic behavior; yet, this did not lead to an increase in aggression or behavior indicating agitation. Further, habituation to videos depicting gorillas did not occur. This supports the effectiveness of this form of enrichment, particularly for a nonhuman animal needing to be separated temporarily due to illness, shipment quarantine, social restructuring, or exhibit modification.     

A preliminary behavioral comparison of two captive all-male gorilla groups

Gorillas live in polygamous harem groups, generally composed of one male, several adult females, and their offspring. With an equal numbers of male and female gorillas born in captivity, however, housing gorillas in social breeding units inevitably means that some males will not have access to female social partners. Thus, the future of the captive gorilla population depends on the collective ability of zoos to house equal numbers of males and females. This study examined the behavioral profiles of two all-male groups of captive lowland gorillas (Gorilla gorilla gorilla) to provide information on this type of housing situation. One group consisted of three subadult individuals, while the other consisted of two subadults and a silverback. Data were collected during two 6-month intervals, for a total of 284 hr. The behavioral profiles of the animals were stable over the course of the study but proximity patterns changed. Differences in feeding, solitary play, and object-directed behavior were found between groups, while no significant differences were observed in affiliative or agonistic social behavior. At both institutions, group cohesion appeared to be high, particularly between subadults; these individuals spent approximately 10% of their time engaging in social behavior and 25-50% of their time in close proximity (within 5 m). However, the Zoo Atlanta males spent significantly more time within 1 m and 5 m of each other than the Santa Barbara males, which may reflect a higher level of cohesiveness among members of the Zoo Atlanta group. The behavioral profiles of the animals in this study were similar to those found in bachelor groups of wild mountain gorillas. One notable exception was the absence of homosexual behavior between the silverback and subadults in Santa Barbara and the low frequency of this behavior between subadults in both groups. Although more longitudinal data are needed, these data suggest that all-male groups can be a feasible housing strategy for males at certain periods of their life span. Zoo Biol 20:27-40, 2001. Copyright 2001 Wiley-Liss, Inc.     

The Occurrence and Ape-to-Ape Transmission of the Entodiniomorphid Ciliate Troglodytella abrassarti in Captive Gorillas

ABSTRACT. Entodiniomorphid ciliates are often present in the colons of wild apes. In captive apes the infection tends to gradually disappear, with the exception of Troglodytella abrassarti . We used fecal examinations to screen the gorillas ( Gorilla gorilla gorilla ) in European (Czech Republic, UK) and Australian Zoos to explore the ape-to-ape transmission pattern of T. abrassarti . Gorillas from two out of three European Zoos were positive for T. abrassarti , while gorillas from the Australian Zoo were negative. We documented a horizontal transmission of T. abrassarti to a non-infected adult gorilla introduced into a Troglodytella -positive group in the Prague Zoo and traced the origin of the ciliate infection to the Paignton Zoo (UK) using serial fecal examinations. During this study, two infant gorillas born in the Prague Zoo (CZ) first became positive for T. abrassarti at the age of 9 mo. Ciliate morphology and the sequencing of the small subunit rRNA gene and the internal transcribed spacer rDNA spacer region revealed that T. abrassarti affects both captive gorillas and chimpanzees. We conclude that zoo transport plays a major role in the distribution of T. abrassarti among captive gorillas.     

Use of buckets as tools by Western lowland gorillas

While all great apes have been documented to use tools, gorillas are arguably the least proficient tool users. In 2009, a Western lowland gorilla (Gorilla gorilla gorilla) at the Buffalo Zoo was observed using a bucket, which had been provided as part of normal enrichment, as a tool to collect water. We conducted a brief, ad libitum investigation to confirm the validity of the initial observation. We then carried out a systematic investigation of the behavior in 2010. We collected 72 hr of videotaped data and tested the null hypothesis that the gorillas did not differ in their prevalence of engaging in bucket-use behaviors. We documented that all four adult gorillas in the group used buckets as drinking tools; however, there was significant individual variation in frequency and type of use of buckets. Four of the eight behaviors showed significant variation among individuals. The silverback male and the youngest adult female contacted and held the bucket significantly more than the remaining two adult females. The young female carried and drank from the bucket significantly more than any other individual. Furthermore, she was observed to fill the bucket with water four of the six times during which this behavior was observed. These data provide evidence of the ability of gorillas to utilize tools, given the appropriate environmental conditions. We continue to explore the abilities of gorillas to recognize the functionality of buckets as tools.     

Nuclear integrations of mitochondrial DNA in gorillas

Great ape systematics, particularly at the species level and below, is currently under debate, due in part to the recent influx of molecular data. The phylogenies of previously published mitochondrial control region (or D-loop) DNA sequences in gorillas show deep splits within West African gorillas (Gorilla gorilla gorilla), and very high levels of nucleotide diversity in this subspecies. Here we demonstrate that several previously reported D-loop haplotypes from West African gorillas are in all likelihood nuclear integrations of mitochondrial DNA. Revised estimates of the amount and pattern of mitochondrial DNA diversity in gorillas are provided, revealing two reciprocally monophyletic and highly divergent groups of gorillas, concurrent with their geographic distribution.     

Patterns of mandibular variation in Pan and Gorilla and implications for African ape taxonomy

Pan and Gorilla taxonomy is currently in a state of flux, with the number of existing species and subspecies of common chimpanzee and gorilla having been recently challenged. While Pan and Gorilla systematics have been evaluated on the basis of craniometric and odontometric data, only a handful of studies have evaluated multivariate craniometric variation within P. troglodytes, and none have evaluated in detail mandibular variation in either P. troglodytes or Gorilla gorilla. In this paper, we examine ontogenetic and adult mandibular variation in Pan and Gorilla. We test the hypothesis that patterns and degrees of mandibular variation in Pan and Gorilla closely correspond to those derived from previous analyses of craniometric variation. We then use these data to address some current issues surrounding Pan and Gorilla taxonomy. Specifically, we evaluate the purported distinctiveness of P.t. verus from the other two subspecies of Pan troglodytes, and the recent proposals to recognize Nigerian gorillas as a distinct subspecies, Gorilla gorilla diehli, and to acknowledge mountain and lowland gorillas as two separate species. Overall, patterns and degrees of multivariate mandibular differentiation parallel those obtained previously for the cranium and dentition. Thus, differences among the three conventionally recognized gorilla subspecies are somewhat greater than among subspecies of common chimpanzees, but differences between P. paniscus and P. troglodytes are greater than those observed between any gorilla subspecies. In this regard, the mandible does not appear to be more variable, or of less taxonomic value, than the face and other parts of the cranium. There are, however, some finer differences in the pattern and degree of morphological differentiation in Pan and Gorilla, both with respect to cranial and dental morphology, and in terms of the application and manner of size adjustment. Mandibular differentiation supports the conventional separation of bonobos from chimpanzees regardless of size adjustment, but size correction alters the relative alignment of taxa. Following size correction, intergroup distances are greatest between P.t. verus and all other groups, but there is considerable overlap amongst chimpanzee subspecies. Amongst gorillas, the greatest separation is between eastern and western gorillas, but adjustment relative to palatal vs. basicranial length results in a greater accuracy of group classification for G.g. gorilla and G.g. graueri, and more equivalent intergroup distances amongst all gorilla groups. We find no multivariate differentiation of the Nigerian gorillas based on mandibular morphology, suggesting that the primary difference between Nigerian and other western lowland gorillas lies in the nuchal region. Though intergroup distances are greatest between P.t. verus and other chimpanzee subspecies, the degree of overlap amongst all three groups does not indicate a markedly greater degree of distinction in mandibular, as opposed to other morphologies. Finally, mandibular differentiation corroborates previous craniodental studies indicating the greatest distinction amongst gorillas is between eastern and western groups. Thus, patterns and degrees of mandibular variation are in agreement with other kinds of data that have been used to diagnose eastern and western gorillas as separate species.     

Visitor effects on the behavior of captive Western lowland gorillas: the importance of individual differences in examining welfare

Given the conflicting and somewhat limited findings available on the effect of zoo visitors on primate behavior, the primary purpose of this study was to provide additional data on gorillas' response to variations in crowd size and to look at what other factors—both intrinsic (e.g. personality, sex, and rearing history) and extrinsic (e.g. group)—might influence such responses. Subjects included four groups (three mixed‐sex and one bachelor) of captive western lowland gorillas living at Zoo Atlanta. Overall activity budgets and rates of social behaviors in high and low crowd conditions were compared. Behaviors suggestive of general welfare did not vary as a function of crowd size alone, although we did observe one significant interaction and several trends. Specifically, two groups spent more time engaged in undesirable behaviors in the high crowd condition, whereas the other two groups showed the opposite response pattern. Additionally, males, but not females, showed a trend toward greater rates of contact aggression in the high crowd condition, although wounding rates did not vary with crowd size. We also found trends towards variation in response to crowd size as a function of individual personality ratings. These results suggest that although the visitor effect may constitute an impediment to optimal animal welfare, this may only apply to some individuals or groups. We emphasize the importance of continuing to explore individual differences and the limitations of a one‐size‐fits‐all approach when describing influences on animal welfare. Zoo Biol 31:586‐599, 2012. © 2011 Wiley Periodicals, Inc.     

Nutritional Aspects of Western Lowland Gorilla (Gorilla gorilla gorilla) Diet During Seasons of Fruit Scarcity at Bai Hokou,Central African Republic

Traditionally, gorillas were classified as folivores, yet 15 years of data on western lowland gorillas (Gorilla gorilla gorilla) show their diet to contain large quantities of foliage and fruit, and to vary both seasonally and annually. The consumption of fruit by gorillas at Bai Hokou, Central African Republic, is correlated with rainfall and ripe fruit availability (Remis, 1997a). We investigated the nutritional and chemical content of gorilla foods consumed at Bai Hokou during two seasons of fruit scarcity as measured by phenological observations and compared our findings with the nutrient content of gorilla foods at other African sites. We conclude that during lean times, Bai Hokou gorillas consumed fruits with higher levels of fiber and secondary compounds than those of other populations of western lowland or mountain gorillas. Conversely, leaves consumed by Bai Hokou gorillas were relatively low in fiber and tannins. Bai Hokou gorillas appeared to meet their nutritional needs by eating a combination of fruit and foliage. They ate fruits comparatively high in secondary compounds and fiber when necessary. While gorillas are selective feeders, wherever and whenever preferred foods are scarce, their large body size and digestive anatomy enable them to consume and process a broader repertoire of foods than smaller bodied-apes.     

Social structure and life-history patterns in western gorillas (Gorilla gorilla gorilla)

Life-history traits and ecological conditions have an important influence on primate social systems. Most of what we know about the life-history patterns and social structure of gorillas comes from studies of eastern gorillas (Gorilla beringei sp.), which live under dramatically different ecological conditions compared to western gorillas (Gorilla gorilla sp.). In this paper we present new data on western gorilla social structure and life histories from four study sites, and make comparisons with eastern gorilla populations. Data were obtained from two study sites with gorilla groups undergoing the habituation process (Lossi, Democratic Republic of Congo and Bai Hokou, Central African Republic) and two "bai" studies (Maya Nord and Mbeli Bai, Republic of Congo). The size and structure of these groups were similar to those seen in eastern gorillas. However, differences in the occurrence of various group transitions (group formations, changes between one-male and multimale composition, and group disintegrations) exist, and western gorillas notably exhibit much higher rates of male emigration and correspondingly fewer multimale groups compared to mountain gorillas. Certain phenomena have been observed only rarely, including predation by leopards. The preliminary data show no significant differences in birth rates between western gorillas and mountain gorillas. The ecological variability across gorilla habitats likely explains the flexibility in the social system of gorillas, but we need more information on the social relationships and ecology of western gorillas to elucidate the causes for the similarities and differences between western and eastern gorillas on the levels of individuals, social groups, and population dynamics.     

Ape behavior in two alternating environments: comparing exhibit and short‐term holding areas

In many facilities, primates are voluntarily transferred between different enclosures on a daily basis to facilitate animal husbandry and exhibit maintenance. This procedure is particularly relevant in the management of great apes living in zoos, where the requirements of functional management must be balanced with the desire to maintain enriching and naturalistic exhibit enclosures that benefit ape residents and attract the visiting public. In these settings, examinations of ape behavior and welfare typically focus exclusively on activity in the primary exhibit area. However, physical, social and sensory experiences unique to each area may shape different patterns of behavior. In the current study, zoo‐living chimpanzees and gorillas were moved each day from exhibit areas to off‐exhibit holding areas for a short duration as a part of regular management procedures. Behavioral data indicated species‐specific reactions to the holding area, including increased aggression and self‐directed behavior by chimpanzees and increased activity and prosocial behavior among gorilla subjects. Both species showed more feeding‐foraging behavior while in the exhibit enclosure. Results suggest that holding areas may not meet all behavior needs of captive great apes and demonstrate the importance of including all components of the captive enclosure in comprehensive analyses of great ape behavior and welfare. Am. J. Primatol. 72:951–959, 2010. © 2010 Wiley‐Liss, Inc.     

Inter-group encounters and female transfer in mountain gorillas: Influence of group composition on male behavior

This paper examines the behaviors used by male gorillas (G. gorilla beringei) to influence female choice during inter-group encounters. Encounters are related to the acquisition of females rather than to the defense of a group's range. Data on 58 independent encounters confirms that encounters are generally aggressive, but that contact aggression is less frequent than previously reported. A high number of potential migrants in one of the groups involved, rather than the newness of either group, predicts the likelihood of contact aggression between males. Encounters with a newly formed unit last longer than encounters involving established groups. Male herding serves to prevent female transfer. Females without dependent offspring are more likely than others to be herded. Proceptive females are the target of herding more often than non-proceptive cycling females. Groups where herding has been observed are more likely to be newly formed, and they include significantly more males. This last result confirms that there are advantages to male cooperation in gorillas. It also raises the question of why such cooperation does not occur more often. © 1993 Wiley-Liss, Inc.