Wild Gorillas Research Articles

Differences in gorilla nettle-feeding between captivity and the wild: local traditions, species typical behaviors or merely the result of nutritional deficiencies?

Behavioral and cognitive studies on captive apes often pay little attention to the specific environmental conditions of their study subjects. A recent report form Byrne et al. (Anim Cogn doi: 10.1007/s10071-011-0403-8, 2011), comparing nettle-feeding techniques between captive and wild gorillas, claimed to document "the strongest evidence yet to come from any great ape that observational learning of a skilled conspecific" can allow social learning and culture in gorillas. An earlier study with similar findings placed emphasis instead on the many similarities and claims for species typical behavior, thus a genetic hypothesis instead of a cultural hypothesis. This commentary aims at formulating a third environmental hypothesis based on path-dependent behavioral differences owing to different diet and availability of nutritional resources of wild and captive gorillas. Captive diet provides gorillas with a much lower concentration of fibers. Gorillas are hindgut fermenters, and this deficit of natural fermentation of fibers may impact their health and their behavior in zoos. Results of Byrne et al.'s study will be discussed comparing feeding choice and availability of nutritional resources of wild and captive gorillas, showing that in captivity gorilla, motivation to consume certain food or certain plant parts may differ drastically from that of wild gorillas. This view does not intend to deny that social learning and culture may exist in gorillas, but to guide and encourage future works investigating social learning in great apes to take more accurately into account the living conditions and, when comparing populations, the possible environmental differences.

Unusual feeding behavior in wild great apes, a window to understand origins of self-medication in humans: Role of sociality and physiology on learning process

Certain toxic plants are beneficial for health if small amounts are ingested infrequently and in a specific context of illness. Among our closest living relatives, chimpanzees are found to consume plants with pharmacological properties. Providing insight on the origins of human self-medication, this study investigates the role social systems and physiology (namely gut specialization) play on learning mechanisms involved in the consumption of unusual and potentially bioactive foods by two great ape species. We collected data from a community of 41–44 wild chimpanzees in Uganda (11months, 2008), and a group of 11–13 wild western gorillas in Central African Republic (10months, 2008–2009). During feeding, we recorded food consumed, its availability, and social interactions (including observers watching conspecifics and the observers' subsequent activity). Unusual food consumption in chimpanzees was twice higher than in gorillas. Additionally chimpanzees relied more on social information with vertical knowledge transmission on unusual foods by continually acquiring information during their life through mostly observing the fittest (pre-senescent) adults. In contrast, in gorillas observational learning primarily occurred between related immatures, showing instead the importance of horizontal knowledge transmission. As chimpanzees' guts are physiologically less specialized than gorillas (more capable of detoxifying harmful compounds), unusual-food consumption may be more risky for chimpanzees and linked to reasons other than nutrition (like self-medication). Our results show that differences in sociality and physiology between the two species may influence mechanisms that discriminate between plants for nutrition and plants with potential therapeutic dietary components. We conclude that self-medication may have appeared in our ancestors in association with high social tolerance and lack of herbivorous gut specialization.

Local traditions in gorilla manual skill: evidence for observational learning of behavioral organization

Elaborate manual skills of food processing are known in several species of great ape; but their manner of acquisition is controversial. Local, "cultural" traditions show the influence of social learning, but it is uncertain whether this includes the ability to imitate the organization of behavior. Dispute has centered on whether program-level imitation contributes to the acquisition of feeding techniques in gorillas. Here, we show that captive western gorillas at Port Lympne, Kent, have developed a group-wide habit of feeding on nettles, using two techniques. We compare their nettle processing behavior with that of wild mountain gorillas in Rwanda. Both populations are similar in their repertoires of action elements, and in developing multi-step techniques for food processing, with coordinated asymmetric actions of the hands and iteration of parts of a process as "subroutines". Crucially, however, the two populations deal in different ways with the special challenges presented by nettle stings, with consistently different organizations of action elements. We conclude that, while an elaborate repertoire of manual actions and the ability to develop complex manual skills are natural characteristics of gorillas, the inter-site differences in nettle-eating technique are best explained as a consequence of social transmission. According to this explanation, gorillas can copy aspects of program organization from the behavior of others and they use this ability when learning how to eat nettles, resulting in consistent styles of processing by most individuals at each different site; like other great apes, gorillas have the precursor abilities for developing culture.

Human Metapneumovirus Infection in Wild Mountain Gorillas, Rwanda

The genetic relatedness of mountain gorillas and humans has led to concerns about interspecies transmission of infectious agents. Human-to-gorilla transmission may explain human metapneumovirus in 2 wild mountain gorillas that died during a respiratory disease outbreak in Rwanda in 2009. Surveillance is needed to ensure survival of these critically endangered animals.

A novel adenovirus of Western lowland gorillas (Gorilla gorilla gorilla)

Adenoviruses (AdV) broadly infect vertebrate hosts including a variety of primates. We identified a novel AdV in the feces of captive gorillas by isolation in cell culture, electron microscopy and PCR. From the supernatants of infected cultures we amplified DNA polymerase (DPOL), preterminal protein (pTP) and hexon gene sequences with generic pan primate AdV PCR assays. The sequences in-between were amplified by long-distance PCRs of 2 - 10 kb length, resulting in a final sequence of 15.6 kb. Phylogenetic analysis placed the novel gorilla AdV into a cluster of primate AdVs belonging to the species Human adenovirus B (HAdV-B). Depending on the analyzed gene, its position within the cluster was variable. To further elucidate its origin, feces samples of wild gorillas were analyzed. AdV hexon sequences were detected which are indicative for three distinct and novel gorilla HAdV-B viruses, among them a virus nearly identical to the novel AdV isolated from captive gorillas. This shows that the discovered virus is a member of a group of HAdV-B viruses that naturally infect gorillas. The mixed phylogenetic clusters of gorilla, chimpanzee, bonobo and human AdVs within the HAdV-B species indicate that host switches may have been a component of the evolution of human and non-human primate HAdV-B viruses.

Behavioral responses of one western lowland gorilla (Gorilla gorilla gorilla) group at Bai Hokou, Central African Republic, to tourists, researchers and trackers

Gorilla tourism, widely perceived as a lucrative industry, is propelled by strong market demand with programs in five countries and for three of four gorilla subspecies. Human presence may negatively affect wild gorillas, potentially lowering immunity and increasing the likelihood of acquiring human-borne disease. Yet, behavioral impacts of humans on wild gorilla behavior remain largely unexplored, particularly for western lowland gorillas. We evaluate the impact of tourist presence, human observer numbers (tourists, trackers, and researchers), and human observer distance on the behavior of one habituated gorilla group at Bai Hokou, Central African Republic. Behavioral data were collected for more than 12 months from January 2007. Of silverback aggressive events, 39% (N=229) were human directed, but 65% were low-level soft barks. Adult females, and one in particular, were responsible for the highest number of aggressive events toward humans. Humans maintained closer proximity to the silverback when tourists were present, although tourist numbers had no significant impact on overall group activity budgets or rates of human-directed aggression. However, as research team size increased, group feeding rates decreased. Close observer-silverback distance correlated with a decrease in his feeding rates and an increase in human monitoring. He directed less aggression toward observers at distances >10 m, although observers spent 48.5% of time between 6 and 10 m of the silverback. We discuss gorilla personality as a factor in human-directed aggression. We explore whether the current 7 m distance limit governing gorilla tourism, based on disease transmission risks, is sufficient considering the potential behavioral stressor of close human presence. We recommend increasing minimum observation distance to >10 m where possible, decreasing observer group sizes, particularly after a visit consisting of maximum numbers and restricting tourist access to 1 visit/day.

Validation of Multiple Diagnostic Techniques to Detect Cryptosporidium sp. and Giardia sp. in Free-Ranging Western Lowland Gorillas (Gorilla gorilla gorilla) and Observations on the Prevalence of these Protozoan Infections in Two Populations in Gabon

Anthropozoonotic diseases threaten the survival of western lowland gorillas (Gorilla gorilla gorilla). Use of accurate diagnostic techniques in gorilla health monitoring contributes to the conservation of gorillas by providing robust information for appropriate management decisions. To identify suitable protozoa diagnostic techniques for wild gorillas, 95 fecal specimens were collected in Lopé National Park and east of Moukalaba-Doudou National Park in Gabon, areas with high and low levels of human activity, respectively. The samples were examined for Cryptosporidium sp. and Giardia sp. by using the following diagnostic techniques: a commercially available immunofluorescent antibody test kit, Merifluor, and a rapid immune-assay, ImmunoCard STAT!, to detect Cryptosporidium sp. and Giardia sp., and a modified Ziehl-Neelsen stain to detect Cryptosporidium sp. oocysts. The results obtained from the Merifluor test, considered the "gold standard" in human studies, were used to estimate the prevalence of Cryptosporidium sp. and Giardia sp. infections in Lopé National Park (19.0% and 22.6%, respectively) and east of Moukalaba-Doudou National Park (0% and 9.1%, respectively). The difference in prevalence in both areas may be associated with differing levels of anthropogenic disturbance. The sensitivity and specificity of the latter two diagnostic techniques were calculated by using the Merifluor test as a control. The ImmunoCard STAT! was found suitable for Giardia sp. antigen detection (specific but not sensitive) and inappropriate for Cryptosporidium sp. antigen detection (not specific or sensitive). The modified Ziehl-Neelsen stain was found to be highly specific but not sensitive in the detection of Cryptosporidium sp. oocysts. These results underline the necessity of using ancillary tests and concentration methods to correctly identify positive samples. This is the first report of Cryptosporidium sp. and Giardia sp. infections in free-ranging western lowland gorillas and highlights the importance of verifying the accuracy of diagnostic techniques developed for human use before applying these to non-human primates.

Effective non-invasive genetic monitoring of multiple wild western gorilla groups

Obtaining reliable population size or abundance estimates of endangered species is critical for their conservation and management. Genotyping non-invasively collected samples is an effective way to gain insights into the number and grouping patterns of rare or elusive animals. In this study we used genetic capture–recapture estimators to obtain a precise estimate of the size of a western gorilla population inhabiting an intensely sampled 101 km 2 area in Loango National Park, Gabon. Using 394 putative gorilla samples collected opportunistically over a 3 year period, we identified 83 unique genotypes. We used a rarefaction curve, Bayesian estimator and two maximum-likelihood methods to estimate that between 87 and 107 individuals used the study area between February 2005 and September 2007. The confidence interval surrounding the genetic estimate was smaller than that obtained using traditional ape survey methods. In addition, genetic analysis showed that gorilla and chimpanzee faeces were identified with 98% and 95% accuracy in the field, respectively. Patterns of co-occurrence of individual gorillas suggest that at least 11 gorilla social groups and five lone silverback males lived in the study area and that several individuals transferred between groups during the 3-year study period. When properly designed and implemented as part of a long-term biomonitoring program, genetic capture-recapture should prove an invaluable tool for evaluating, even on a large-scale, the population size and dynamics of apes and other elusive species.

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.

Molecular Epidemiology of Simian Immunodeficiency Virus Infection in Wild-Living Gorillas

Chimpanzees and gorillas are the only nonhuman primates known to harbor viruses closely related to HIV-1. Phylogenetic analyses showed that gorillas acquired the simian immunodeficiency virus SIVgor from chimpanzees, and viruses from the SIVcpz/SIVgor lineage have been transmitted to humans on at least four occasions, leading to HIV-1 groups M, N, O, and P. To determine the geographic distribution, prevalence, and species association of SIVgor, we conducted a comprehensive molecular epidemiological survey of wild gorillas in Central Africa. Gorilla fecal samples were collected in the range of western lowland gorillas (n = 2,367) and eastern Grauer gorillas (n = 183) and tested for SIVgor antibodies and nucleic acids. SIVgor antibody-positive samples were identified at 2 sites in Cameroon, with no evidence of infection at 19 other sites, including 3 in the range of the Eastern gorillas. In Cameroon, based on DNA and microsatellite analyses of a subset of samples, we estimated the prevalence of SIVgor to be 1.6% (range, 0% to 4.6%), which is significantly lower than the prevalence of SIVcpzPtt in chimpanzees (5.9%; range, 0% to 32%). All newly identified SIVgor strains formed a monophyletic lineage within the SIVcpz radiation, closely related to HIV-1 groups O and P, and clustered according to their field site of origin. At one site, there was evidence for intergroup transmission and a high intragroup prevalence. These isolated hot spots of SIVgor-infected gorilla communities could serve as a source for human infection. The overall low prevalence and sporadic distribution of SIVgor could suggest a decline of SIVgor in wild populations, but it cannot be excluded that SIVgor is still more prevalent in other parts of the geographical range of gorillas.

The strategic use of sex in wild female western gorillas

Human females, unlike most mammals, are sexually active outside of fertile periods. This decoupling of sexual behavior from its conceptive function has had an enormous impact on human social relationships, and yet we know little about why there was selection for nonconceptive mating. Here we examine one form of nonconceptive mating, the mating that occurs during pregnancy or post-conceptive (PC) mating, in wild western gorillas (Gorilla gorilla). Using a near complete mating record for five females during gestation, we show that pregnant females varied in the timing and frequency of mating, and used PC mating conditionally, synchronizing copulations to occur on days when other females mated, and refraining from mating for lengthy periods when no other females mated. As pregnant females mated exclusively with the same male before and after conception, and mated in response to group female (and not male) behavior, we conclude that western gorillas used PC mating as a form of female competition, and not to confuse paternity or to obtain immediate benefits from the male, as suggested earlier. The male initiated copulations preferentially with females of high rank, rather than distinguishing between pregnant and cycling females. Therefore, PC mating appears to be a strategy by which high-ranking pregnant females attempt to minimize male interest in other females, while reinforcing their own status and potentially delaying conception in others. These findings indicate that female-mating competition is more important than considered earlier, and may be a factor in the evolution of nonconceptive mating in humans.

Interactions between zoo‐housed great apes and local wildlife

Although there are published reports of wild chimpanzees, bonobos, and orangutans hunting and consuming vertebrate prey, data pertaining to captive apes remain sparse. In this survey-based study, we evaluate the prevalence and nature of interactions between captive great apes and various indigenous wildlife species that range into their enclosures in North America. Our hypotheses were threefold: (a) facilities housing chimpanzees will report the most frequent and most aggressive interactions with local wildlife; (b) facilities housing orangutans and bonobos will report intermediate frequencies of these interactions with low levels of aggression and killing; and (c) facilities housing gorillas will report the lowest frequency of interactions and no reports of killing local wildlife. Chimpanzees and bonobos demonstrated the most aggressive behavior toward wildlife, which matched our predictions for chimpanzees, but not bonobos. This fits well with expectations for chimpanzees based on their natural history of hunting and consuming prey in wild settings, and also supports new field data on bonobos. Captive gorillas and orangutans were reported to be much less likely to chase, catch and kill wildlife than chimpanzees and bonobos. Gorillas were the least likely to engage in aggressive interactions with local wildlife, matching our predictions based on natural history. However unlike wild gorillas, captive gorillas were reported to kill (and in one case, eat) local wildlife. These results suggest that some behavioral patterns seen in captive groups of apes may be useful for modeling corresponding activities in the wild that may not be as easily observed and quantified. Furthermore, the data highlight the potential for disease transmission in some captive settings, and we outline the associated implications for ape health and safety.

Hand-clapping as a communicative gesture by wild female swamp gorillas

Hand-clapping is a form of gestural communication commonly observed in captive great apes yet only isolated instances of this behaviour have been documented in the wild. Nearly 20 years ago Fay recorded the first observations of hand-clapping in western lowland gorillas (Gorilla gorilla gorilla) in the Central African Republic. Here we present observations of Likouala swamp gorillas using hand-clapping as a form of gestural communication in previously undocumented contexts in the wild. We observed hand-clapping on four different occasions in four different groups. The hand-clap was always exhibited by an adult female and always consisted of two consecutive claps conducted in front of the body. We suggest the functional significance of the behaviour was to maintain and enforce group cohesiveness during instances of alarm. These observations suggest western lowland gorillas have a means of communicating that is thus far absent in their eastern counterparts (Gorilla beringei ssp.). This could be a gestural culture found only in western lowland gorillas which should be investigated further to shed light on the evolution of communication among hominoids.

The 60th anniversary of Japanese primatology

Japanese primatology is considered to have begun in December 1948, when Kinji Imanishi (then 46 years old), Shunzo Kawamura (24 years old), and Junichiro Itani (22 years old) visited Koshima Island to look into the possibility of conducting a field study of Japanese monkeys there. Kawamura and Itani were already impressed by the well-organized procession of monkeys they had observed at Cape Toimisaki the previous month when they were observing semi-wild horses as Imanishi’s assistants (Itani 1991; Kawai 2007). As Imanishi had already read through Carpenter’s pioneering studies, his change of study targets from horses to macaques was not according to the mood of the moment. This was a case in which Fortune knocks on the door of the person who is prepared for it (Nishida 2007). Since this epoch-making survey of Japanese monkeys in the natural habitat of southern Japan, 60 years have passed, and we celebrated the 60th anniversary of Japanese primatology last year. During this period, Japanese primatology gradually took shape as it passed several milestones, such as the establishment of the Japan Monkey Centre in 1956, the Laboratory of Physical Anthropology at Kyoto University in 1962, the Kyoto University Primate Research Institute in 1966, the Tsukuba Primate Research Center in 1978, and the Primate Society of Japan in 1985. The first issue of Primates appeared in 1957, 5 years earlier than the second primate journal, Folia Primatologica. However, owing to its irregular publication in the earliest period, producing only one volume in 2 years, vol. 50 appears this year, in 2009. Consequently, vol. 50 of Primates is publishing special contributions in the first and second issues. Primates has published many momentous papers, including those on the proposal to study primate culture, the initiation of wild gorilla research, and primate politics (vol. 1); the importance of kin relationships in organizing primate societies and the first detailed study of primate vocalizations (vol. 4); the first documentation of primate cultural transmission and the discovery of infanticide (vol. 6); discoveries of chimpanzee (vol. 9) and bonobo (vols. 20 and 23) social units; and a contribution to the social brain hypothesis (vol. 31). As one of the events to commemorate the 60th anniversary, the associate editors based in Japan gathered at Inuyama in the summer of 2007 to discuss the special publication of Primates. Because the special issue on Japanese macaque studies had been published not long before (vol. 39), we decided to collect reviews on recently developed studies based on the editors’ recommendations. Regrettably, some candidates could not comply with our request because of the tight publishing schedule. However, 11 authors eventually accepted our invitations, and we are pleased to announce that we will feature the first three papers in this issue. Eight papers will appear in the next issue, to be published in April. In 2002, the editorial board of Primates was reorganized into associate editors and an advisory board, whose members have increased both in number and in international representation. In 2003, Springer began publishing Primates with the financial support of the Japan Monkey Centre and the Primate Society of Japan. These two changes strengthened the reviewing power of the journal (three referees ordinarily review original articles and one referee reviews short communications, in T. Nishida (&) Japan Monkey Centre, Inuyama, Japan e-mail: nishida@jinrui.zool.kyoto-u.ac.jp

Host-Parasite Ecology of the Helminths in Mountain Gorillas

To understand patterns of intestinal parasitism in healthy, undisturbed endangered mountain gorillas (Gorilla beringei), we regularly collected fecal samples from a group of 14 wild gorillas residing in Bwindi Impenetrable National Park (BINP), Uganda, for about 1 yr. The objectives of the study were to collect baseline data in order to document the helminth parasites infecting this group of gorillas and to examine the effects of season and host age-sex class on patterns of parasite infection. In addition to weekly surveys of feces from all group members, fecal samples from 4 identified individuals were examined almost daily. We identified the diagnostic stages of the following parasites: strongylids (Strongylida), Anoplocephala gorillae, Probstmayria sp., Strongyloides fuelleborni, and a trematode. Monthly and daily fluctuations in strongylid egg counts were observed. Infants had lower strongylid egg counts compared with other group members. Both of the silverbacks had higher mean egg counts in the wet season than in the dry season. Examination of fecal samples from identifiable gorillas revealed high day-to-day variation in strongylid egg counts. No evidence of anthropozoonotic transmission of intestinal helminths was found.

Genetic identification of elusive animals: re‐evaluating tracking and nesting data for wild western gorillas

AbstractWestern gorillas Gorilla gorilla have been exceedingly difficult to habituate to the presence of human observers. Nevertheless, researchers have amassed a wealth of information on population densities and group structure for this ape species by locating and counting the sleeping nests of wild individuals. Such nest‐count studies have suggested that western gorilla groups often have multiple silverbacks and these multimale groups occasionally divide into smaller subgroups. However, observational data from forest clearing sites and from a few recently habituated western gorilla groups show no evidence of multimale family groups or of subgrouping. This discrepancy underscores a long‐standing question in ape research: How accurately do nesting sites reflect true group compositions? We evaluated these indirect measures of group composition by using DNA from faeces and hair to genetically identify individual gorillas at nesting sites. Samples were collected from unhabituated wild western gorillas ranging near Mondika Research Center in the Central African Republic and Republic of Congo. DNA extracted from these samples was genotyped at up to 10 microsatellite loci and one X–Y homologous locus for sex identification. Individuals were then identified at nesting sites by their unique multilocus genotypes, thus providing a ‘molecular census’ of individual gorillas. Results confirm that western gorillas often build more than one nest at a nesting site and, thus, nest counts can be highly inaccurate indicators of group size and composition. Indeed, we found that nest counts can overestimate group size by as much as 40%, indicating that true gorilla population numbers are probably lower than those reported from census surveys. This study demonstrates how genetic analysis can be a valuable tool for studying and conserving elusive, endangered animals.

Food preferences of wild mountain gorillas

Determining the nutritional and phenolic basis of food preference is important for understanding the nutritional requirements of animals. Preference is a measure of which foods would be consumed by an animal if there was no variation in availability among food items. From September 2004 to August 2005, we measured the food preferences of four wild mountain gorilla groups that consume foliage and fruit in Bwindi Impenetrable National Park, Uganda, to determine what nutrients and phenols are preferred and/or avoided. To do so, we asked the following questions: (1) Which plant species do the gorillas prefer? (2) Considering the different plant parts consumed of these preferred species, what nutrients and/or phenols characterize them? (3) Do the nutritional and phenolic characteristics of preferred foods differ among gorilla groups? We found that although some species were preferred and others were not, of those species found in common among the different group home ranges, the same ones were generally preferred by all groups. Second, all groups preferred leaves with relatively high protein content and relatively low fiber content. Third, three out of four groups preferred leaves with relatively high sugar amounts. Fourth, all groups preferred pith with relatively high sugar content. Finally, of the two groups tested, we found that the preferred fruits of one group had relatively high condensed tannin and fiber/sugar contents, whereas the other group's preferred fruits were not characterized by any particular nutrient/phenol. Overall, there were no differences among gorilla groups in nutritional and phenolic preferences. Our results indicate that protein and sugar are important in the diets of gorillas, and that the gorillas fulfil these nutritional requirements through a combination of different plant parts, shedding new light on how gorillas balance their diets in a variable environment.

An experimental study of nettle feeding in captive gorillas

Mountain gorillas (Gorilla beringei beringei) in Karisoke, Rwanda, feed on the stinging nettle Laportea alatipes by means of elaborate processing skills. Byrne [e.g. Philosophical Transactions of the Royal Society of London, Series B, Biological Sciences 358:529-536, 2003] has claimed that individuals acquire these skills by means of the so-called program-level imitation, in which the overall sequence of problem-solving steps (not the precise actions) is reproduced. In this study we present western lowland gorillas (Gorilla gorilla gorilla) with highly similar nettles. Twelve gorillas in three different groups (including also one nettle-naïve gorilla) used the same program-level technique as wild mountain gorillas (with differences mainly on the action level). Chimpanzees, orangutans, and bonobos did not show these program-level patterns, nor did the gorillas when presented with a plant similar in structural design but lacking stinging defenses. We conclude that although certain aspects (i.e. single actions) of this complex skill may be owing to social learning, at the program level gorilla nettle feeding derives mostly from genetic predispositions and individual learning of plant affordances.

The development of feeding behaviour in wild western lowland gorillas (Gorilla gorilla gorilla)

Before being weaned, young primates must learn which plant species are suitable to eat, and how to process and combine them to meet their nutritional needs. This study provides a detailed investigation of the development of feeding behaviour in a wild population of western lowland gorillas observed in a bai habitat. The behaviour of 56 known immature individuals aged between 0 and 8 years was recorded at Mbeli Bai, Republic of Congo, using focal, scan and all-occurrence sampling over a period of 16 months. Results indicated that independent feeding skills were acquired by the end of infancy and did not undergo significant change beyond this point. However, suckling continued into the juvenile period, suggesting that some continued investment by the mother was necessary until the strength and skills of immatures developed, perhaps to enable self-sufficiency through periods of succulent fruit (assumed to be the most suitable weaning food) scarcity. This was supported by observations of an increase in feeding-related behaviour with age, particularly during infancy. Additionally, age-related differences in the species and plant parts selected by different age classes, as well as in levels of food processing, were apparent. Play feeding, suggested to be a precursor to independent feeding, was most prevalent in infancy, decreasing as feeding skills developed. Thus, it would appear, given that food transfer also occurs (Nowell & Fletcher, 2006), that feeding skills in western lowland gorillas are learned by a combination of exposure to and manipulation of food items, and probably also through the observation of conspecifics. Observations in a forest habitat are needed to improve our understanding of feeding development in this subspecies.

Patterns of paternity and group fission in wild multimale mountain gorilla groups

To understand variation among social systems, it is essential to know the relative reproductive success of individuals in group-living species. Particularly interesting for such studies are taxa such as mountain gorillas in which both one-male and multimale groups are common, because of the opportunity to estimate the costs and benefits to males of pursuing different reproductive strategies. We genotyped 68 individuals from two groups of multimale mountain gorilla groups in Bwindi Impenetrable National Park, Uganda to determine the distribution of paternity among the males. In both groups, the dominant male sired the majority of offspring. One group underwent a fission, and we found that the eight offspring assigned to the dominant silverback (and their mothers) remained with their father, while the two offspring of unknown paternity ended up in the small group headed by the formerly subordinate silverback. This is consistent with the proposal that the outcome of group fission in primates is not only influenced by maternal relationships among individuals, but also by patrilineal relationships. Results of this study show that subordinate males may gain reproductive benefits even while queuing for dominance status. Despite ecological differences between Bwindi and the Virunga Volcanoes, male mountain gorillas living in both populations benefit from remaining in multimale groups.