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.     

Are gorillas right-handed or not?

Comparative data on laterality of function in primates are useful for elucidating its evolution, including its link to asymmetry of cerebral structure. Recently the gorilla (Gorilla gorilla, Pongidae) has been nominated as the other hominoid species closest toHomo sapiens in terms of handedness. This study aims to scrutinise the 22 accounts of gorillas' hand preferences. Numerous shortcomings exist in the published literature on the topic, so that a firm conclusion is not yet possible. It is not yet clear whether gorillas are right-preferent, left-preferent, or ambi-preferent, and additional data are needed.     

A case study of mother-son transfer in mountain gorillas

In mountain gorillas (Gorilla g. beringei), male immigration in bi-sexual units is rare. This paper presents the case of a nearly weaned male infant gorilla who followed his mother in her transfer. This case was recorded in the study population at the Kariske Research Center in 1988. The data come from observation in Group B (on 12 days just prior to the transfer and on 54 days after the transfer over a period of 6 months). The situation of the infant did match the conditions in which infanticide occurs in gorillas, but he was not killed, despite receiving male aggression and being wounded twice. In fact, both the mother and the infant received male aggression more frequently than the long term residents in the group. The aggression received by the mother decreased after she mated with the males and after she weaned the infant. The aggression received by the infant, however, did not decrease after his mother mated with the males, and increased in intensity. The infant reacted fearfully to male aggression, in marked contrast to his mother, who reacted either with indifference, or by simply avoiding the males. The aggression eventually stopped, and the infant became a blackback in Group B. Evolutionarily, the death of the infant would not have markedly accelerated the mother's return to estrus, but the death of the infant could still have benefitted the males, by decreasing the reproductive output of a competitor. Adult male gorillas are also presumably selected to resist male immigration. Proximately, the aggression directed towards the infant was not related to mating access to the mother. The sex of the infant may contribute to explain the post-transfer male aggression, but data on the integration of old female infants in a new group is needed to test whether the sex of the infant has an effect on their vulnerability to infanticide. Also, the intense fear displayed by the infant may have played a role in prompting male aggression.     

The function of male aggressive displays towards females in mountain gorillas

In groups ofGorilla g. beringei, male aggression towards females regularly takes the form of male display. This paper examines male display towards females in two Karisoke study groups (Group 5 and Group BM) in 1989, a period when none of the females were new immigrants. Results are based on 259 hr of focal observations and 121 hr ofad libitum observations on male behaviors towards females. The goal is to see if the data are compatible with four non-mutually exclusive hypotheses to explain male displays towards females: (1) demonstration of male fighting abilities to influence female long term residence decision; (2) decrease potential competitive inequities between females; (3) provision to females of an occasion to confirm their subordinance to a male; and (4) short term influence on mating. First, male-female proximity was tested against proportion of male displays, to rule out the possibility that males display towards females simply because they happen to be close by. There was no association between proximity and male display. Dominant males were responsible for a higher proportion of displays than subordinate males. This is consistent with the idea that males display to demonstrate their fighting abilities, or their qualities as protector, since dominant males are the ones offering long term protection against infanticide and predators. Females that were in a position to transfer did not receive a higher proportion of male display, however. Long term resident dominant females received a higher proportion of displays from the dominant males, which is consistent with the idea that males attempt to decrease potential competitive inequities between females. There was an association between female appeasement reactions and male displays, which suggests that males display to create occasions for the females to confirm their subordinance to them. Estrous females did not receive a higher proportion of male displays, and there was no association between male display and copulation, suggesting that male displays are not a form of courtship aggression aimed at influencing mating in the short term.     

The subspecies ofHomo erectus

ClassifyingHomo erectus into subspecies can be based on either temporal or geographical differences, but there is no accepted system for using both. This can be done with subspecies names consisting of two elements — a prefix ofneo, meso, orpaleo to indicate grade, followed by a geographical term ofeuropus, africus, sinicus, orindicus to indicate line. Thus Rhodesian isHomo erectus neoafricus, Ngandong isHomo erectus neoindicus, Peking isHomo erectus mesosinicus, ER 3733 isHomo erectus paleoafricus, etc.     

‘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.     

Spontaneous use of sticks as tools by captive gorillas (Gorilla gorilla gorilla)

The present report describes the spontaneous use of sticks, as tools by young adult gorillas (Gorilla gorilla gorilla) in a social group at the San Diego Wild Animal Park, CA, USA. Three 8-year-old gorillas (one female and two males) threw sticks into the foliage of trees, which the gorillas could not climb due to electric wire, to knock down leaves and seeds. Two of the three gorillas selected sticks that were more suitable (i.e. longer or thicker sticks) for throwing. Moreover, they looked up at the target (i.e. the foliage of the tree) before throwing and grasped the stick at a position appropriate, for throwing (i. e. the end of the stick). They were more likely to throw sticks when particular adult group members were not nearby. These two gorillas also pulled branches of trees toward themselves by using sticks to capture them (branch-pulling), and one of these two beat a branch with a stick to knock down leaves and seeds (branch-beating). One of these two gorillas used much longer sticks for branch-pulling than for stick-throwing, indicating that he was capable of task-dependent selection of sticks. Analyzing the spontaneous use of sticks as tools by gorillas in captivity can lead to a better understanding of not only their cognitive ability but also of their social relationships which may otherwise be concealed.     

Intrapopulation differences in ant eating in the mountain gorillas of Bwindi Impenetrable National Park, Uganda

Variability in ant eating has been observed in several populations of eastern and western gorillas. We investigated the occurrence of ant (Dorylus sp.) eating in two groups of mountain gorillas (Gorilla beringei beringei) with overlapping home ranges within Bwindi Impenetrable National Park, Uganda from September 2001 to August 2002. We calculated the frequency of ant eating by an indirect method of analyzing fecal samples from silverbacks, adult females, and juveniles. One group consumed ants significantly more often than the other (3.3 vs 17.6% of days sampled). Furthermore, the group that consumed ants more often also consumed them on a seasonal basis (September–February monthly range: 0–8%; March–August monthly range: 30–42.9%). Finally, females and juveniles of this group consumed ants significantly more often than did the silverback (total samples containing ants: silverback, 2.1%; adult female, 13.2%; juvenile, 11.2%). Differences in ant eating between groups are likely due to variability in use of habitats where ants occur (particularly secondary forests). Surveys of ant densities in differing habitats, nutritional analysis of ants, and quantification of the amount of ants in their diets are necessary to understand if ant consumption is due to availability, nutritional value, group traditions, or taste preference.     

Intra-specific variation in social organization of gorillas: implications for their social evolution

We analysed intra-specific variation in the social organization of gorillas and ecological and social factors influencing them, based on recent data on diet, day journey length, home range size, group size and proportion of multi-male groups in three subspecies [western lowland gorillas (WLG); eastern lowland gorillas (ELG); mountain gorillas (MG)]. Median group size was similar across subspecies and across habitats, but the extraordinarily large group including >30 gorillas was only found in habitat with dense terrestrial herbaceous vegetation. Within-group competition may determine the upper limit of group size in frugivorous WLGs and ELGs in lowland habitats with scarce undergrowth. A frugivorous diet may be a causal factor of subgrouping in multi-male groups of WLGs and ELGs, while a folivorous diet may prevent subgrouping in multi-male groups of MGs. Social factors, rather than ecological factors, may play an important role in the formation of multi-male groups and their cohesiveness in MGs. High gregariousness of female gorillas and their prolonged association with a protector male are explained by their vulnerability to both infanticide (MGs) and predators (ELGs). Comparison of long-term changes in group composition and individual movements between ELGs in Kahuzi and MGs in the Virungas suggest that the occurrence of infanticide may promote kin-male association within a group. Threat of infanticide may stimulate MG females to transfer into multi-male groups to seek reliable protection and maturing MG males to stay in their natal groups after maturity. By contrast, the absence of infanticide may facilitate ELG females to associate with infants and other females at transfer and ELG males to establish large groups in a short period by taking females from their natal groups, by luring females from neighbouring groups, or by takeover of a widow group after the death of its leading male. These conditions may prevent ELG and WLG maturing males from remaining to reproduce in their natal groups and possibly result in a rare occurrence of multi-male groups in their habitats. Similar reproductive features of MG and ELG females suggest both female strategies have been adaptive in their evolutionary history.     

Potential for female kin associations in wild western gorillas despite female dispersal

Female philopatry and male dispersal are the norm for most mammals, and females that remain in their natal region often derive foraging or social benefits from proximity to female kin. However, other factors, such as constraints on group size or a shortage of potential mates, may promote female dispersal even when female kin associations would be beneficial. In these cases, female kin associations might develop, not through female philopatry, but through female emigration to the same group. To date, little attention has been focused on the potential for kin-biased behaviour between females in female-dispersing species. Here we investigate the genetic relationships among adults in eight wild groups of unhabituated western gorillas (Gorilla gorilla) at the Mondika Research Center using microsatellite genotyping of DNA collected from hair and faeces. We found that almost half (40%) of adult females had an adult female relative in the same group and average within-group relatedness among females was significantly higher than that expected under a model of random dispersal. This provides the first genetic evidence that females can maintain social associations with female relatives in spite of routine natal and secondary dispersal. In addition, we show that females appear to avoid related silverback males when making dispersal decisions, suggesting that a strategy of non-random female dispersal may also function to avoid inbreeding.     

Intra- and inter-group interactions of an all-male group of virunga mountain gorillas (Gorilla gorilla beringei)

Six unrelated male gorillas formed an all-male group within the Virunga mountain gorilla population. Frequent homosexual interactions characterized the high cohesiveness of this group. Such homosexual behavior reduced the inter-individual distances and increased the social tension between the two silverbacks in the group. The silverbacks retained “ownership” of the homosexual partners, but competed and fought with each other violently when the partners avoided or ignored their courtship. Neither submissive nor reassurance behavior was noted between the silverbacks. Thus, their relationships may not be explained in terms of dominance and subordinancy. However, the loser-support and mediating behavior observed in the group prevented them from engaging in severe fights. Aggression was always directed from the elder and dominant males to the younger and subordinate males, while supporting interactions occurred in the opposite direction. The blackbacks frequently supported the subadult aggressees by attacking the silverbacks, and the younger males displayed mediating behavior in violent fights between the silverbacks. The group's ranging was influenced by encounters with neighboring social units. The members avoided contact with other units and shifted their range after several encounters. On the other hand, when and after a subadult male had immigrated into their group, they frequently encountered other units and did not move away from the encounter site. An all-male group may not be a favorable unit for females to transfer to, but may be profitable for maturing males to associate with. Its formation is probably related to recent social change in the Virunga gorilla population.     

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.     

The role of taste in food selection by African apes: implications for niche separation and overlap in tropical forests

Ripe fruit eating shapes the behavior of most of the apes. Gorillas (Gorilla gorilla) and chimpanzees (Pan troglodytes) are very different sizes and, consequently, have been traditionally viewed as ecologically distinct, but few studies have explored the behavioral and physiological foundations of their diets. Debate continues on the extent that large-bodied gorillas may be less selective and more opportunistic fruit eaters than chimpanzees. Taste responses have been predicted to relate to body size and digestive strategies. This study employs laboratory research on taste perception and discrimination among captive zoo-housed chimpanzees and relates it to previous work on gorillas to better characterize diets and niche separation among these apes. During the captive trials, differences were recorded in consumption patterns of water and varying concentrations of dilute aqueous fructose (sweet) and tannic acid solutions (astringent), compounds commonly found in wild foods. The chimpanzees exhibited similar preference thresholds for fructose (50 mM) to other primates studied. They exhibited slightly lower inhibition thresholds for tannic acid solutions than gorillas, but higher than smaller primates studied to date. These preliminary findings suggest that tannin tolerance may well be mediated by body size, though possible species differences in salivary proteins or other sensory differences remain to be explored. This research furthers our efforts to understand the roles of body size and physiological adaptations in shaping diet and niche separation of chimpanzees and gorillas.     

Buccal dental microwear variability in extant African Hominoidea: taxonomy versus ecology

Buccal microwear patterns on teeth are good indicators of the abrasiveness of foodstuffs and have been used to trace the dietary habits of fossil species, including primates and hominids. However, few studies have addressed the variability of this microwear. The abrasiveness of dietary components depends not only on the hardness of the particles ingested, but also on the presence of dust and other exogenous elements introduced during food processing. These elements are responsible for the microwear typology observed on the enamel surfaces of primate teeth. Here we analyzed the variability of buccal microwear patterns in African Great Apes (Gorilla gorilla and Pan troglodytes), using tooth molds obtained from the original specimens held in several osteological collections. Our results suggest that ecological adaptations at subspecies or population level account for differences in microwear patterns, which are attributed to habitat and ecological conditions within populations rather than differences between species. The findings from studies on the variability of buccal dental microwear in extant species will contribute to a better understanding of extinct hominids’ diet and ecology.     

Gorillas (Gorilla gorilla gorilla) in the Likouala swamp forests of north central Congo: Preliminary data on populations and ecology

Until recently it was generally believed that gorillas avoid water. In two recent, independent faunal surveys in the Likouala swamp of north central Congo, we have found that gorillas occur at high densities, extending the known range of Gorilla gorilla gorillavery close to the Oubangui River. The gorillas in this area fed on the abundant ground vegetation of herbaceous monocotyledons in the swamp forest. In adjacent islands of terra firmathere was little evidence of gorilla nesting or feeding, indicating that they spend at least part of the year almost exclusively in the swamp forests. Recent reports from several sites in West Africa indicate that swamp forests may generally provide abundant food resources that are used regularly by gorillas.     

Revision of the subspecies ofAustralopithecus africanus (primates: Hominidae), including a new subspecies from the late pliocene of Ethiopia

The subspecies ofAustralopithecus africanus Dart, 1925 have been revised in a morphological and statistical analysis. Four subspecific names were previously proposed, but only one was found to be valid. The subspeciesA. africanus transvaalensis (Broom, 1936), from the Plio/Pleistocene of South Africa, cannot be sustained due to an insufficient sample, and is combined with the nominate race,A. a. africanus. The type ofA. africanus afarensis Tobias, 1980 is a mistake in identification and notA. africanus, but a pongid. The population ofA. africanus from the late Pliocene of Ethiopia does indeed represent a relatively small-toothed geographical race for which the nameA. africanus aethiopicus was conditionally proposed; and the lectotype for it, A.L. 288-1, is notA. africanus, but the type ofHomo antiquus Ferguson, 1984. The trinominalaethiopicus is thus unavailable for the Ethiopian race, which is redescribed as a new subspecies,A. africanus miodentatus n. ssp., and the mandible A.L. 266-1 is designated as the holotype.     

Newly Discovered Gorilla Population in the Ebo Forest,Littoral Province,Cameroon

We report on a new population of gorillas discovered in November 2002 in the Ebo Forest, Littoral Province, Cameroon. We observed A group of q7 gorillas directly for 83 min, and they were in auditory range for 155 min. Further evidence of gorilla presence included 8 nest groups totaling 38 nests, distinctive feeding signs accompanied by footprints, and a gorilla cranium collected from the nearby village of Iboti. This newly discovered gorilla population is geographically intermediate between the 2 extant populations of western gorillas: Gorilla gorilla gorilla, the most populous gorilla subspecies living in Gabon, Equatorial Guinea, Congo-Brazzaville, Central African Republic and Cameroon to the south of the Sanaga River, and G. g. diehli or the Cross River gorilla, a small population of ca. 250 individuals on the Cameroon-Nigeria border. It is not possible to assign the new gorilla population to either subspecies on the basis of measurements of the single male cranium. Genetic analyses of freshly shed hairs, collected from gorilla nests, may help to resolve the taxonomic status of the Ebo gorillas.     

Foods consumed by sympatric populations ofGorilla g. gorilla andPan t. troglodytes in Gabon: Some preliminary data

Data on foods consumed by gorillas and chimpanzees living in primary forest in Gabon were collected, mainly by examination of the contents of feces. Gorillas ate fruit very regularly (some fruit remains were present in 97.6% of 246 fecal samples examined), in addition to leaves, stems, pith, and bark. Some fruit remains were present in all chimpanzee fecal samples examined. Mean numbers of fruit species per fecal sample were 2.5 for gorillas and 2.1 for chimpanzees. Sixty percent of all identified foods recorded for gorillas were recorded for chimpanzees as well. Our results indicate that important differences in diet exist between western lowland gorillas and the eastern gorilla populations of Kahuzi-Biega and the Virunga Volcanoes. It is now clear that western gorillas cannot be accurately classed as folivores.