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.     

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.     

Behavioral and hormonal responses to the availability of forage material in Western lowland gorillas (Gorilla gorilla gorilla)

We investigated how forage material affects indicators of welfare in three male Western lowland gorillas (Gorilla gorilla gorilla) at the Detroit Zoo. In addition to their maintenance diet and enrichment foods, the gorillas generally received forage material four times a week. From this baseline, we systematically manipulated how much forage material the group received on a weekly basis, with either daily or bi (twice)‐weekly presentation of browse (mulberry, Morus sp.) or alfalfa hay. We collected behavioral data (60 hr per gorilla) and measured fecal glucocorticoid metabolites (FGM). Mixed models indicated that the presence of forage material significantly increased time feeding (F_2,351 = 9.58, p < 0.001), and decreased rates of noncontact aggression (F_2,351 = 3.69, p = 0.03), and regurgitation and reingestion (F_2,353 = 4.70, p = 0.01). Regurgitation and reingestion were never observed during the condition when forage material was provided daily. When forage material was provided, time spent feeding was similar across gorillas, compared to a disproportionately greater amount of time spent feeding by the dominant individual when forage material was absent. Providing forage material in addition to the regular diet likely created more opportunities for equitable feeding for the subordinate gorillas. FGM concentrations did not vary based on the presence or type of forage material available and, instead, likely reflected group social dynamics. In general, alfalfa and mulberry had similar impacts on behavior, indicating that alfalfa can be an adequate behavioral substitute during times when browse is less readily available for gorillas housed in seasonally variable climates.     

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.     

Non‐invasive fecal hormone analysis and behavioral observations for monitoring stress responses in captive western lowland gorillas (Gorilla gorilla gorilla)

Non‐invasive techniques for monitoring the stress response in captive western lowland gorillas (Gorilla gorilla gorilla) were investigated. Fecal samples for cortisol measurement and concurrent behavioral data were collected from six individuals in a socially housed gorilla group (one adult male, three adult females and their three offspring) over a 7‐month period. Despite inter‐individual variation in the dynamics of fecal cortisol concentrations over time, several major secretory peaks coincided across individuals. High cortisol concentrations in feces were correlated with induced stressors or behavioral observations indicating high social tension, with a 1–2 day lag period. Entry progression order of the gorillas into a den complex and a supplant‐based dominance index were suitable indicators of overall dominance hierarchies, and fluctuations over time reflected periods of instability. Diurnal variation in fecal cortisol was not apparent when comparing afternoon and morning samples, however the sample collection interval was relatively short (3–5 hr). These results demonstrate the feasibility of monitoring stress responses based on the dynamics of both fecal cortisol excretion and behavior. This non‐invasive approach may be used for gauging responses to changes in husbandry, environment and group structure of captive gorillas. Zoo Biol 0:1–15, 2005. © 2005 Wiley‐Liss, Inc.     

Insectivory by Gorilla gorilla gorilla in Southeast Cameroon

Our study extends quantitative analyses of insect-eating by gorillas (Gorilla gorilla gorilla) to Cameroon. During a 2-mo period (May–June 2001), we recorded and analyzed feeding traces on plants and insects and in gorilla feces. We found 180 feeding traces, 17% of which involved insectivory. Seventy-eight percent of the fecal samples (n = 36) contained insects. Ants were found in 61% of the samples, termites in 39%, while 56% of the samples contained remains of other insects. We added 14 new species to the known insect diet of western lowland gorillas. Overall, social insects are predominant. The choice of prey by the Ntonga gorillas gives new clues for the existence of cross-cultural differences among gorilla populations. A comparison of the overall frequency of insectivory with those at other sites in Central Africa indicate a possible effect of forest disturbance on the insectivorous behavior of gorillas. The study suggests the existence of temporal variation in ant- and termite-eating by gorillas.     

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.     

Relative growth,ontogeny, and sexual dimorphism in gorilla(Gorilla gorilla gorilla and G. g. beringei): Evolutionary and ecological considerations

Gorillas are the largest and among the most sexually dimorphic of all extant primates. While gorillas have been incorporated in broad-level comparisons among large-bodied hominoids or in studies of the African apes, comparisons between gorilla subspecies have been rare. During the past decade, however, behavioral, morphological, and molecular data from a number of studies have indicated that the western lowland (Gorilla gorilla gorilla) and eastern mountain (Gorilla gorilla beringei) subspecies differ to a greater extent than has been previously believed. In this study I compare patterns of relative growth of the postcranial skeleton to evaluate whether differences between subspecies result from the differential extension of common patterns of relative growth. In addition, patterns of ontogeny and sexual dimorphism are also examined. Linear skeletal dimensions and skeletal weight were obtained for ontogenetic series of male and female G.g. gorilla (n = 315) and G.g. beringei (n = 38). Bivariate and multivariate methods of analysis were used to test for differences in patterns of relative growth, ontogeny, and sexual dimorphism between sexes of each subspecies and in same-sex comparisons between subspecies. Results indicate males and females of both subspecies are ontogenetically scaled for postcranial proportions and that females undergo an earlier skeletal growth spurt compared to males. However, results also indicate that the onset of the female growth spurt occurs at different dental stages in lowland and mountain gorillas and that mountain gorillas may be characterized by higher rates of growth. Finally, data demonstrate lowland and mountain gorilla females do not differ significantly in adult body size, but mountain gorilla males are significantly larger than lowland gorilla males, suggesting mountain gorillas are characterized by a higher degree of sexual dimorphism in body size. Thus, although lowland and mountain gorillas do not appear to have evolved novel adaptations of the postcranium which correlate with differences in locomotor behavior, the present investigation establishes subspecies differences in ontogeny and sexual dimorphism which may be linked with ecological variation. Specifically, these findings are evaluated in the context of risk aversion models which predict higher growth rates and increased levels of sexual dimorphism in extreme folivores. Am. J. Primatol. 43:1–31, 1997. © 1997 Wiley-Liss, Inc.     

The effect of visitors on the behavior of zoo-housed western lowland gorillas (Gorilla gorilla gorilla)

Primates, especially apes, are popular with the public, often attracting large crowds. These crowds could cause behavioral change in captive primates, whether positive, neutral, or negative. We examined the impact of visitors on the behavior of six western lowland gorillas (Gorilla gorilla gorilla), observing the troop over 6 weeks during high season (4.5 hr/day, 35 days, May–July 2016). We used focal scan sampling to determine activity budget and enclosure usage, and focal continuous sampling to identify bouts of anxiety-related behavior (visitor-directed vigilance, self-scratching, and aggression). Both daily zoo-entry numbers (V_GATE) and instantaneous crowds at the exhibit (V_DENSITY) were measured. Overall, V_GATE had little effect across behaviors. However, consistent with the more acute time frame of measurement, V_DENSITY was a better predictor of behavior; at high crowd volumes, we observed significant group-level changes in activity budget (increased inactivity, increased locomotion, and decreased environment-related behaviors), increase in some anxiety-related behaviors, and decreased enclosure usage. Although contributing similar effects, it could not be determined if crowd numbers, composition, or noise most affected the troop, nor any chronic effects of exposure to large crowds. Nevertheless, our findings suggest that measures to minimize the impacts of large crowds at the exhibit would be beneficial. Furthermore, we highlight potential discrepancies between common methods for measuring visitor numbers: V_GATE is less sensitive to detecting visitor effects on behavioral indices than V_DENSITY. Future studies should appropriately match the biological time frame of welfare indicators and visitor measures used to ensure the reliability of findings.     

Sexual behavior and hormonal estrus cycles in captive aged lowland gorillas (Gorilla gorilla)

To evaluate whether observed cycles in proceptive behavior in aging lowland gorilla females (age 40+) at Brookfield Zoo were driven by ovarian activity, we compared monthly behavioral data to estradiol and progestogen cycles based on fecal hormone assessments. Progestogen peaks showed regularity and close coincidence with monthly sexual behaviors. Estradiol was more variable. Progestogen peaks varied between 22+/-5 days for the control female (29 years old), to 24+/-2.5 and 29+/-8 for the two aged subjects. In the first aged female, which was housed with other females and a silverback, the high degree of cyclicity in sexual behavior, regularity of progestogen cycles, and close concordance between hormonal cycling and sexual behavior strongly compared to patterns found (in this and other studies) in gorilla females <35 years old. Cyclical progestogen peaks were longer and more variable in the second aged female-perhaps because she lacked the social mediation of other females or a male. For husbandry reasons she is not housed with the gorilla group, behavioral data were not collected from her. The value of our longitudinal study is in obtaining reproductive profiles of primate females that are approaching maximum lifespan. This pilot study is part of a larger research project on reproductive senescence that will include other captive females >35 years old, a population that is rapidly increasing in North American zoos as gorillas continue to age.     

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.     

Sexual and Hormonal Cycles in Geriatric Gorilla gorilla gorilla

As our closest living relatives, great apes likely experience behavioral and physiological patterns associated with reproductive aging and menopause that are similar to human patterns. We present results from a nationwide zoo-based study on behavioral and hormonal changes in female western gorillas. We evaluated progestogen concentrations via daily fecal sampling in 30 gorillas, 22 of which were geriatric (≥30). We collected concurrent behavioral data 1–3 times weekly on 16 of the females. While control females cycled regularly, ca. 23% of geriatric females are acyclic (menopausal), and another 32% show variable hormonal patterns suggesting perimenopause. Patterns included increased variability in cycle length and peak progestogen values, and frequent insufficient increases in progestogen levels during the luteal phase. Acyclic females have significantly lower overall progestogen concentrations than the self cycling females, though differences are not significant when cycle phase is incorporated. We detected behavioral estrus in 9 of 10 cycling females for which data were available. In all but 1 case, proceptive behavior occurred during the follicular phase, preceding ovulation on average by 6.6 d. Females spent more time in proximity to the silverback male while in behavioral estrus than during other periods. To date, maximum longevity in captive female gorillas is 52 yr, with poor reproductive prognosis beginning from the age of 37. We demonstrate that both perimenopause and menopause characterize aged female gorillas, which may experience a postreproductive lifespan of >25% of their lives. Continued study of aging apes is warranted, and apes may serve as models for age-related reproductive changes in humans.     

A case study of primiparous maternal and infant gorilla (Gorilla gorilla gorilla) behavior

A primiparous mother and her infant were subjects of a longitudinal behavioral study at Lincoln Park Zoo in Chicago, Illinois. From November 1998 to November 1999, we collected a total of 100 hr of focal nonhuman animal instantaneous point sampling and all occurrence data on the mother and her infant. After 8 months, we introduced the 4-member focal group to an 8-member gorilla group, thus providing an opportunity to study the effect of the introduction on mother-infant behavior. Overall, time the pair spent physically apart was notably high with contact and proximate scores steadily decreasing over time. The infant was largely responsible for maintaining social proximity, rarely leaving the mother and frequently approaching her. In contrast, the mother left the infant frequently and approached the infant minimally. Maternal affiliative and nonaffiliative behaviors fluctuated throughout the study. Over time, both decreased. Despite a precarious maternal relationship, infant developmental trends were typical for captive gorillas. The results of this study suggest that allowing flexibility in judging maternal conduct can be of benefit to successful gorilla husbandry.     

Nest Building in Captive Gorilla gorilla gorilla

Although various aspects of gorilla nest building have been described in wild populations, nest-building behavior of captive gorillas has not been subject to much scientific review. We observed nest building in 17 gorillas during three periods: summer baseline, winter baseline, and winter treatment, in which the amount of available nesting material was doubled. We conducted observations exclusively in the indoor holding area in the hour following evening departure of animal care staff. During baseline, gorillas engaged in nest-building on 3.1% of scans and were on a constructed nest on 27.9% of scans. Overall, gorillas spent significantly more time on elevated nests than on floor nests. There were no statistically significant sex, age class, or rearing history differences in nest building, time spent on a nest, or nest location preference. Nest building consisted of both gathering and manipulating materials. The gorillas spent significantly more time building nests in winter than in summer. Additionally, they were more often on elevated nests during winter than summer. Doubling the amount of nesting material did not increase nest building, but it increased the number of floor nests. Our results suggest that providing adequate materials to captive gorillas for nest building may facilitate performance of species-typical nest-building, minimize competition among individuals for nesting sites and materials, and permit individual thermoregulation.     

Urinary estrogen excretion during pregnancy in the gorilla (Gorilla gorilla), orangutan (Pongo pygmaeus) and the human (Homo sapiens)

Urinary estrogen components were separated, identified and quantified throughout the pregnancy of the gorilla (Gorilla gorilla) and orangutan (Pongo pygmaeus) and compared to estrogen levels in normal human pregnancies. Fetal and neonatal adrenals from each species were also compared in terms of weight and relative amounts of fetal zone. The results demonstrate that gorillas and chimpanzees excrete 4- to 5-fold less estrogen during pregnancy than the human and orangutan which are similar to each other. The lower estrogen excretion appears to be related to a smaller fetal adrenal in both the gorilla and chimpanzee which reveal both a reduced adrenal weight and increased definitive to fetal zone ratio when compared to either the human or orangutan.     

Socialization processes in a female lowland gorilla

A low-interactive, captive, female lowland gorilla, Molly, was studied following the introduction into her enclosure of three gorillas, two males and a female, raised from birth in captive gorilla groups. Observations were made 6 mo after the new gorillas were introduced. Throughout the period of observation, Molly interacted in an affiliative manner with one of the males, playing or sitting quietly with him in a tree (where Molly spent most of her time) and occasionally on the ground. Agonistic displays between Molly and the new female decreased after they were released in the enclosure without the males for a series of days. Molly, however, continued to react to the other male, the most dominant, in an agonistic manner, and usually retreated from his reach, climbed the tree, and/or grimaced and piloerected whenever he approached. Although Molly's continued avoidance of the dominant male impeded her complete socialization, we propose that the interventions employed in this study—introduction of new younger gorillas into and an enclosure, and a series of dyadic separations between the noninteractive gorilla and each of the new group members—are possible strategies that can be used to facilitate socialization of captive, noninteractive gorillas.     

Mitochondrial COII Introgression into the Nuclear Genome of Gorilla gorilla

Numts are nonfunctional mitochondrial sequences that have translocated into nuclear DNA, where they evolve independently from the original mitochondrial DNA (mtDNA) sequence. Numts can be unintentionally amplified in addition to authentic mtDNA, complicating both the analysis and interpretation of mtDNA-based studies. Amplification of numts creates particular issues for studies on the noncoding, hypervariable 1 mtDNA region of gorillas. We provide data on putative numt sequences of the coding mitochondrial gene cytochrome oxidase subunit II (COII). Via polymerase chain reaction (PCR) and cloning, we obtained COII sequences for gorilla, orangutan, and human high-quality DNA and also from a gorilla fecal DNA sample. Both gorilla and orangutan samples yielded putative numt sequences. Phylogenetically more anciently transferred numts were amplified with a greater incidence from the gorilla fecal DNA sample than from the high-quality gorilla sample. Data on phylogenetically more recently transferred numts are equivocal. We further demonstrate the need for additional investigations into the use of mtDNA markers for noninvasively collected samples from gorillas and other primates.     

The Intestinal Entodiniomorph Ciliates of Wild Lowland Gorillas (Gorilla gorilla gorilla) in Gabon,West Africa

Examination of 109 fecal samples from wild lowland gorillas revealed the presence of five species of entodiniomorph ciliates: Troglodytella ahrassarti, Troglodytella gorillae, and three unclassified species. These latter three species were also found in the feces of a captive gorilla in Gabon and are considered to be intestinal parasites or commensals.     

A Case Study of Primiparous Maternal and Infant Gorilla (Gorilla gorilla gorilla) Behavior

A primiparous mother and her infant were subjects of a longitudinal behavioral study at Lincoln Park Zoo in Chicago, Illinois. From November 1998 to November 1999, we collected a total of 100 hr of focal nonhuman animal instantaneous point sampling and all occurrence data on the mother and her infant. After 8 months, we introduced the 4-member focal group to an 8-member gorilla group, thus providing an opportunity to study the effect of the introduction on mother-infant behavior. Overall, time the pair spent physically apart was notably high with contact and proximate scores steadily decreasing over time. The infant was largely responsible for maintaining social proximity, rarely leaving the mother and frequently approaching her. In contrast, the mother left the infant frequently and approached the infant minimally. Maternal affiliative and nonaffiliative behaviors fluctuated throughout the study. Over time, both decreased. Despite a precarious maternal relationship, infant developmental trends were typical for captive gorillas. The results of this study suggest that allowing flexibility in judging maternal conduct can be of benefit to successful gorilla husbandry.     

Monitoring the reproductive status of female gorillas (Gorilla gorilla gorilla) by measuring the steroid hormones in fecal samples

The reproductive status of female gorillas (Gorilla gorilla gorilla) was estimated by measuring the sex steroid hormones in fecal samples instead of in blood samples. Fecal samples from female gorillas were used to examine the reliability of this non-invasive assay system, which included the extraction method for estradiol-17β (E₂) and progesterone (P) from fecal samples. Fecal samples from three female gorillas were collected daily for about 55 days, and fecal E₂ and P were assayed to clarify the fluctuation patterns of these steroids in the feces. Fecal sampling from one female was repeated for another 50-day period (starting 75 days after the end of the first observation period) and assayed to confirm if the menstrual cycle of this subject was ovulatory. Although fecal E₂ concentration measurements were quantitative by using this assay system, fecal P concentration measurements were semi-quantitative. Relative amounts of fecal P in fecal samples were estimated by using the values of B/B₀ (bound/total binding in the radioimmunoassay system). Two of the four fluctuation patterns of fecal hormones observed throughout the menstrual cycle for the three female gorillas were typical for normal ovulatory cycles. In the subject observed for two periods, one pattern was typical and the other atypical. The results show that this non-invasive method is simple and practical for monitoring the reproductive status of great apes as well as Old World monkeys.