Reconciliation and Consolation in Captive Western Gorillas

We studied post-aggression mechanisms in a captive group of western gorillas (Apenheul Primate Park, The Netherlands) and compared them with those of wild mountain gorillas (Gorilla beringei). We found the same trend for reconciliation that wild mountain gorillas show: reconciliation occurred only between adult male-female dyads, while it was absent in the other sex-age class combination. There were both solicited and nonsolicited contacts; the latter finding is in contrast with the result obtained in wild mountain gorillas, in which consolation was absent. Immature females were more likely to offer consolation toward both related and unrelated individuals. Consolation did not reduce the likelihood of further attacks among group members. It may be that, as the α-male plays a fundamental role in preventing the spread of conflicts throughout the entire group, triadic contacts become ineffective for the function. The levels of consolation were higher in absence of reconciliation than in its presence, suggesting that consolation may function as an alternative mechanism in stress reduction of the victim.     

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.     

Digesta Passage,Digestibility and Behavior in Captive Gorillas Under Two Dietary Regimens

Gorilla adaptation has been debated in recent years given the wide variation among diets of gorillas in different habitats. Gorillas are the largest of living primates, have large colons and should be capable of processing tough foods. Preliminary captive studies have suggested that they may well have long average gut retention times relative to smaller hominoids, which should facilitate digestive efficiency in their wild counterparts. Indeed, wild gorillas consume large amounts of fibrous foods as staples or fall-back foods across their range, in response to habitat-related or seasonal changes in fruit availability. Fluctuations in diet might be matched by changes in digesta passage and digestibility, with possible selective retention of harder to digest items. We further studied digestive processes via chemical cobalt and chromium markers to track liquid and solids, as they passed through the guts of gorillas at the San Francisco Zoo (SFZ). In addition, we examined the effects of variation in captive diets on intake, digesta passage, digestion and behavior. The SFZ gorillas exhibited high digestibility coefficients, and gut passage was long relative to those of smaller-bodied hominoids. The results permit us to understand more fully the relationships of digestive processes to adaptation and dietary flexibility in the wild and to inform the development of dietary recommendations to improve the well-being of captive gorillas.     

Habitat Use and Structural Preferences of Captive Western Lowland Gorillas (Gorilla gorilla gorilla(: Effects of Environmental and Social Variables

Exhibit naturalism has become the architectural standard for new zoo exhibits, yet our scientific understanding of how animals interact with naturalistic environments is very limited. Given the expense incurred in building new exhibits, it is essential that more information on animal-environment interactions be obtained and that it be incorporated at the outset into exhibit designs. We documented four years of habitat use and structural preferences of western lowland gorillas at Zoo Atlanta. We found that quality of space rather than quantity of space was important, as subjects spent 50% of their time in <15% of the exhibits. Subjects showed strong preferences for the areas near structures, particularly the holding building, and spent significantly less time away from structures than expected. Temperature interacted with structural preferences in that the subjects spent more time away from structures when temperatures were cold and more time near them when temperatures were hot. There was no difference in habitat use and structural preferences due to age, sex, and rearing history, but social factors appear to play a role. Our results are similar to those found a decade earlier in the same population and to other studies of space use in apes but are the first to include significant temperature effects. Additionally, they suggest that managers and designers need to take into account factors such as quality of space, attractiveness of the holding building, and the interaction of structural preferences with variables such as temperature and social factors when designing exhibits.     

Factors Influencing the Well-Being and Longevity of Captive Female Orangutans

I examined factors affecting the health and mortality of female orangutans in captivity and identified specific aspects of husbandry that appear to be critical to long-term survival in captivity. There are certain high-hazard events in the life of a captive female orangutan, including birth and parturition. Other than the short periods of high hazard, and perhaps of more significance to long-term survival, the major factors that appeared to influence long-term health and mortality included primiparous age, interbirth interval, rearing type, and weight.     

Peripartum Sex Steroid Profiles and Endocrine Correlates of Postpartum Maternal Behavior in Captive Gorillas (Gorilla gorilla gorilla)

An association between pregnancy levels of estrogen and progesterone and maternal behavior has been demonstrated in several taxonomic orders of nonprimate and primate mammals, but has not so far been investigated in the gorilla. In this study we investigated whether prepartum titers of urinary estrone conjugates (E1C) or pregnanediol-3alpha-glucuronide (PdG) were related to postpartum maternal behavior in eight multiparous Western lowland gorilla females (Gorilla gorilla gorilla) housed in four zoological gardens. Urine samples were collected from each study animal for 14 days prepartum and 14 days postpartum, and measures of maternal responsiveness were scored during the first 15 days postpartum. Urine samples were assayed with radioimmunoassay for E1C and PdG. Results for the peripartum profiles of urinary E1C, as well as postpartum profiles of PdG, agree with previous findings for the gorilla, while results for late-pregnancy profiles of urinary PdG were inconclusive in confirming a prepartum increase or decrease. Neither prepartum levels of E1C or PdG, nor the E1C/PdG ratio were found to be related to measures of postpartum maternal behavior. This lack of association between late-pregnancy E1C titers per se and postpartum maternal behavior is contrary to findings in nonprimate and other primate species.     

Female Genito-genital Rubbing in a Group of Captive Chimpanzees

Wild and captive female bonobos exhibit a form of noncopulatory sexual behavior—genito-genital (GG) rubbing—in which 2 individuals rub their genital regions together. GG-rubbing in bonobos occurs in several contexts and may serve several functions, including tension reduction and reconciliation. It has not been reported for chimpanzees. In a study of captive, adolescent chimpanzees at the New Iberia Research Center, I observed that females rubbed genital regions at rates equivalent to those of bonobos, though the form of the behavior differed in significant ways from classic bonobo GG-rubbing. I describe the pattern of GG-rubbing in the chimpanzees and provide preliminary tests of 5 hypotheses for why the behavior occurs. All 4 females in the study group participated in GG-rubbing, though the frequency with which they initiated and participated varied. Females that GG-rubbed the most also groomed each other the most, supporting the hypothesis that GG-rubbing reinforces or at least reflects social bonds. The data do not support the hypothesis that females GG-rub to reconcile conflicts, to reduce tension during feeding, to signal social status, or to attract mates.     

Influence of Female Copulation Calls on Male Sexual Behavior in Captive Macaca fascicularis

We investigated the function of copulation calls—vocalizations by females during mating—in captive groups of long-tailed macaques. We tested predictions of the contest-competition, sperm competition, synchronized orgasms, mate again, alpha-male notification and graded-signal hypotheses. We observed 371 copulations of 36 females wherein the presence or absence of a copulation call was clear. Females call equally often with different males and shortly after ejaculation. Copulation calls occurred equally with copulations with and without ejaculation. Calls did not incite disruptions of the mating. Following calls females mated again, more often than expected, with their mating partner. Both pregnant and fertile females uttered copulation calls. Two females conceived and mated mainly with the alpha male then. We conclude that copulation calls do not incite male contest competition for sexual access to females and that it is unlikely that calls synchronize male and female orgasms. Several hypotheses remain plausible, but not all predictions are borne out unequivocably. This alerts us to the possibility that the calls could have multiple beneficial effects; natural selection might strike a compromise among functions. Investigation of the mate again, sperm competition and alpha-male notification hypotheses, and of hypotheses not tested in our study concerning female breeding overlap and female-female agonism, is required.     

Effects of Manual Disability on Feeding Skills in Gorillas and Chimpanzees

Foraging gorillas and chimpanzees employ skilful bimanual techniques to process food plants, and there are wild populations in which 10–20% of individuals have severe hand injuries. We examined the feeding skills of one injured gorilla and two injured chimpanzees, while they dealt with plants for which intact peers employ complex techniques. Feeding efficiency was only slightly compromised in the disabled subjects, and none of them had acquired novel techniques specific to their remaining capacity. Instead, all three subjects used techniques like those of able-bodied individuals, though when several optional methods were available, their balance of preference was different. They adapted the techniques to their disabilities by the flexible substitution of a range of alternative means to achieve each step of the process, including unusual grips, use of one hand instead of two or vice versa, and use of the mouth or a foot. Compensation was at the level of detailed execution rather than overall technique and depended on transferring motor organization to novel effector organs. Our findings have implications for theories of the acquisition of complex manual skills in great apes and for the flexibility of great ape mental skills.     

Feeding Competition and Agonistic Relationships Among Bwindi <Emphasis Type="Italic">Gorilla beringei</Emphasis>

Testing predictions of socioecological models, specifically that the types of feeding competition and social relationships female primates exhibit are strongly influenced by the distribution, density, and quality of food resources, requires studies of closely related populations of subjects living under different ecological conditions. I examined feeding competition and the resulting female social relationships in mountain gorillas (Gorilla beringei beringei) of Bwindi Impenetrable National Park, Uganda, which has ecological conditions distinctive from those where other gorilla populations live. I observed 1 group of gorillas for 29 mo to examine the proportion of time spent foraging on fruit, the relationship between patch size and occupancy patterns of fruit trees, and agonistic interactions. Patch occupancy time while foraging in fruit trees decreased with increasing number of gorillas in a tree and decreasing tree size, suggesting that fruit trees represent limiting patches and can lead to intragroup scramble competition. Gorillas exhibited higher levels of aggression while feeding on fruit versus other food resources, which indicates intragroup contest competition. I observed a linear dominance hierarchy with no bidirectionality via displacements, and a similar hierarchy via aggression, though a notable proportion of the dyads contained 2-way interactions. However, most aggression was of low intensity (vocalizations) and the recipient typically ignored it. Despite differences in ecological conditions and diet between the Virunga Volcanoes and Bwindi, agonistic relationships among females are largely similar in the 2 populations and are best characterized as dispersal individualistic.     

Competition and succession among coral endosymbionts

Host species often support a genetically diverse guild of symbionts, the identity and performance of which can determine holobiont fitness under particular environmental conditions. These symbiont communities are structured by a complex set of potential interactions, both positive and negative, between the host and symbionts and among symbionts. In reef‐building corals, stable associations with specific symbiont species are common, and we hypothesize that this is partly due to ecological mechanisms, such as succession and competition, which drive patterns of symbiont winnowing in the initial colonization of new generations of coral recruits. We tested this hypothesis using the experimental framework of the de Wit replacement series and found that competitive interactions occurred among symbionts which were characterized by unique ecological strategies. Aposymbiotic octocoral recruits within high‐ and low‐light environments were inoculated with one of three Symbiodiniaceae species as monocultures or with cross‐paired mixtures, and we tracked symbiont uptake using quantitative genetic assays. Priority effects, in which early colonizers excluded competitive dominants, were evidenced under low light, but these early opportunistic species were later succeeded by competitive dominants. Under high light, a more consistent competitive hierarchy was established in which competitive dominants outgrew and limited the abundance of others. These findings provide insight into mechanisms of microbial community organization and symbiosis breakdown and recovery. Furthermore, transitions in competitive outcomes across spatial and temporal environmental variation may improve lifetime host fitness.     

Competition for Resources and Its Behavioral Consequences Among Female Primates

Via the current model on the evolutionary ecology of female social relationships, Sterck et al. (1997) argue that ecological conditions determine how competition over food resources affects female fitness. The relative importance of different modes of competition then affects female social relationships and dispersal patterns. I outline the model and review relevant data. There are 3 modes of feeding competition: within-group scramble (WGS), within-group contest (WGC), and between-group contest (BGC), which occur in various combinations in different populations of nonhuman primates. Ecological measures support predictions that limiting resources lead to WGS and clumped resources induce WGC. The ecological basis of BGC remains elusive, but it is probably linked to resource abundance. Tests of the proxies of feeding competition support the idea that short-term search substrates and increasing group size lead to WGS, while high-quality patches of intermediate size relative to group size lead to WGC. However, when tested across populations, independent measures of aggression rates do not always match the actual or presumed competitive regimes. This mismatch might be explained by confounding factors and the predominately indirect measures of feeding competition. Predicted relationships between feeding competition and female social relationships/dispersal are only partly supported. This might be attributed to the fact that few studies have taken ultimate approaches using mechanistic correlates of fitness (net energy gain) or lifetime reproductive success to measure consequences of feeding competition. But to resolve existing inconsistencies, additional factors need to be taken into account as well, for example, male sexual strategies may affect female feeding competition; constraints on group size may enforce female dispersal; and demography may alter rates of alliances. More explicitly, ultimate approaches are needed to test the consistency of the socioecological model.     

Modeling the Impact of Ebola and Bushmeat Hunting on Western Lowland Gorillas

The 2003 outbreak of Ebola in the Republic of Congo killed 114 people and up to 800 western lowland gorillas. This outbreak and all outbreaks between 2001–2003 began with human handling of infected animal carcasses. Ebola has since spread, putting the entire gorilla population at risk. An epidemiological model is presented to describe the combined effects of Ebola and hunting on persistence of gorillas. The number of infected gorillas also provides a means of assessing the risk of transmission to humans. Under current harvest practices and the estimated annual outbreak rate, the gorilla population is predicted to undergo a 97% decline within 100 years. Controlling bushmeat hunting may not be enough to prevent extinction if frequent outbreaks occur.     

Female competition in chimpanzees

Female chimpanzees exhibit exceptionally slow rates of reproduction and raise their offspring without direct paternal care. Therefore, their reproductive success depends critically on long-term access to high-quality food resources over a long lifespan. Chimpanzee communities contain multiple adult males, multiple adult females and their offspring. Because males are philopatric and jointly defend the community range while most females transfer to new communities before breeding, adult females are typically surrounded by unrelated competitors. Communities are fission–fusion societies in which individuals spend time alone or in fluid subgroups, whose size depends mostly on the abundance and distribution of food. To varying extents in different populations, females avoid direct competition by foraging alone or in small groups in distinct, but overlapping core areas within the community range to which they show high fidelity. Although rates of aggression are low, females compete for space and access to food. High rank correlates with high reproductive success, and high-ranking females win direct contests for food and gain preferential access to resource-rich sites. Females are aggressive to immigrant females and even kill the newborn infants of community members. The intensity of such aggression correlates with population density. These patterns are compared to those in other species, including humans.     

Gorillas of Bwindi-Impenetrable Forest and the Virunga Volcanoes: Taxonomic implications of morphological and ecological differences

Based on their geographic proximity to the Virunga Volcanoes (≈ 25 km), the Bwindi-Impenetrable Forest gorillas have been referred to the subspecies Gorilla gorilla beringei. Differences in anatomy, habitat, ecology, and behavior, however, suggest Bwindi gorillas are distinct from those in the Virungas. Relative to Virunga gorillas, Bwindi gorillas live at lower elevations, in warmer temperatures, are much more arboreal, have longer day ranges and larger home ranges, and eat much more fruit and pith, and less bamboo and leaves. Morphological differences reflect the differences in ecology, habitat, and behavior. Bwindi gorillas measured have smaller bodies, relatively longer limbs, hands, and feet, shorter trunks, thumbs, big toes, and tooth row lengths, and narrower trunks and orbital breadths than Virunga gorillas. These differences indicate Bwindi gorillas do not belong to G.g. beringei and should not be referred to as “mountain gorillas.” How unique the distinguishing features of Bwindi gorillas are, and whether or not they should be assigned to a new taxon, depends on the expression of these features in eastern lowland gorillas (G.g. graueri). © 1996 Wiley-Liss, Inc.     

Sperm Competition in the Acari

Literature on sperm competition in the Acari is reviewed and supplemented with the author's unpublished observations. As in other animals, morphology and physiology of the female reproductive tract are key factors establishing the sperm competition pattern. First-male sperm priority was found in Gamasida and Actinedida, whereas in Acaridida the last males to mate had the highest reproductive success. However, this general pattern may be modified by behavioural factors, and another aspect needs scrutiny: in many groups of mites sperm cells follow routes other than the lumen of the reproductive tract to meet and fertilize the oocyte. The Acari are a very promising group for future research because of the wide range of sperm competition patterns, accompanied by great diversity in reproductive morphology, physiology and mating.     

Rates of Agonism by Diurnal Lemuroids: Implications for Female Social Relationships

Sterck and colleagues (Behaviour 134:749–774, 1997) focused attention on the evolutionary ecology of female social relationships within and between groups and proposed a model that distinguishes 4 categories of female relationships, which correspond to particular types of intra- and intergroup competition. They emphasized literature on haplorhines in their model because of numerous, detailed studies conducted on a range of species in the wild; in contrast, strepsirrhines such as the lemuroids are poorly represented. We evaluate more closely their classification of lemuroids as Dispersal-Egalitarian using a greater number of species of Lemur, Eulemur, Varecia, Hapalemur, Indri, and Propithecus. For the focal species we found that female philopatry occurs rarely, agonistic rates are relatively low, female dominance hierarchies are not stable and do not exist year-round, and intra- and intergroup female-female competition is infrequent. Therefore, our results support the suggestion that a majority of lemuroid taxa we surveyed correspond to the Dispersal-Egalitarian category with 2 probable exceptions: Lemur catta and Propithecus edwardsi. Because female Lemur catta are philopatric, have year-round dominance hierarchies with female matrilines, exhibit the highest rates of agonism in studied lemuroids, and have frequent intra- and intergroup female-female competition, it would seem that they more closely correspond to the category Resident-Nepotistic. However, maternal Lemur catta rarely support their offspring in agonistic contests and matrilineal rank is not inherited, which leads us to state that the species does not fit into any existing category that explains the nature of female social relationships. The relationships of female Propithecus edwardsi are also a challenge to categorize under the current model because some of their characteristics —typical female dispersal and low agonistic rates— fall into the Dispersal-Egalitarian category, yet other behaviors —intense targeted aggression and stable and year-round female dominance hierarchies— do not.     

育幼经历对圈养成年雌性大熊猫尿液皮质醇含量的影响

为研究育幼期不同的育幼方式对圈养成年雌性大熊猫Ailuropoda melanoleuca福利状况的影响情况,了解圈养育幼模式可能引发成年雌性大熊猫的应激问题,完善育幼期大熊猫的饲养管理和兽舍参数设计,提高育幼过程中圈养大熊猫福利状况,选择成都大熊猫繁育研究基地育幼期采取不同育幼方式管理的成年雌性大熊猫为研究对象,利用...     

Female receptivity and male mate-guarding in the jewel spiderGasteracantha minax thorell (Araneidae)

In field populations, several male jewel spidersGasteracantha minax Thorell (Araneidae) may be found at the periphery of the orb web of a female, indicating that males may compete for fertilisation success. Laboratory experiments revealed that virgin femaleG. minax readily remate shortly after their first mating. However, they appear to enter a refractory period between 1 and 24 h postmating and respond aggressively to courting males. Males that have mated with a female initially defend her from rival males but cease to do so after the onset of the refractory period. These data can be interpreted within the context of mate-guarding and sperm competition. There is marked size dimorphism in this species, which may be the result of selection for protandry. In contrast with other orb-weaving spiders, this selection pressure does not seem to be counterbalanced by selection for larger male size through either sexual cannibalism or male-male competition.