Regurgitation/reingestion and coprophagy in captive gorillas

The behavior of wild animals can be significantly affected by captivity. Among the least documented adaptations are (1) the regurgitation and reingestion of food and (2) coprophagy. This paper reviews the occurrence of the two phenomena in captive lowland gorilla populations. In addition, data and speculations based on wild populations are discussed. Data from a preliminary study at the Stone Zoo are reported, and several causal relationships are explored.     

Complex leaf-gathering skills of mountain gorillas (Gorilla g. beringei): Variability and standardization

The skills that mountain gorillas use to deal with the stings, tiny hooks, and spines protecting common plant leaves in their diet were examined for variation within and between animals. Many elements of uni- and bimanual performance were identified, often involving delicate precision and coordination, and varying idiosyncratically, erch individual having a different set of preferred elements. Many of these elements are functionally equivalent, and all but one weaned animals showed full processing capability; the history of the one exception suggests that early experience with the task may be important. Gorillas' idiosyncrasy in manual skill elements is entirely consistent with trial-and-error learning at this level. By contrast, each individual uses very few techniques (structured sequences of elements) for most processing, and these techniques are the same across the population. Where animals deviate from this generalization, they largely employ the simpler technique normally used for undefended leaves. Lateralization increases from start to finish, consistent with a logical structure in which each stage has a laterality bias and each stage is sequentially dependent on the last. Variations from their commonest, techniques occur in all animals (on average, about nine variant techniques were recorded from each animal). The repertoire of techniques increases significantly with age, whereas the repertoire of elements does not. This points to an initial reliance on a single logical structuring that is well established by weaning (about 3.5 years), with subsequent development of the ability to vary the technique used so as to take advantage of variations in the environment. Standardization of logical organization, despite variability between different animals in individual elements and behavioral laterality, suggests that the logical ordering of elements and the interrelationships of processing stages is copied by program-level imitation. © 1993 Wiley-Liss, Inc.     

Observations on the ontogeny of feeding behavior in mountain gorillas (Gorilla gorilla beringei)

This report describes observations on the ontogeny of food choice in mountain gorillas (Gorilla gorilla beringei), made during a 17-month field study of mountain gorilla feeding ecology in the Parc National des Volcans, Rwanda. Data are presented on the feeding behavior of two infants observed from birth to the age of 8 months and on older infants and juveniles. This information is compared with data on the composition and diversity of the diets of young adults and adults in the same social group. Initial feeding by infants is usually synchronized with the mother's behavior: infants ingest the same food, or a different part of the same plant species, currently being eaten by the mother or just eaten by her. This suggests that observational learning is largely responsible for the transmission of food preferences. Most feeding by young infants, whether or not synchronized with the mother's, is on those foods eaten most frequently by adults. Infants also independently sample potential foods, some of which are apparently not consumed by adults. The frequency of sampling declines with age, although even adults occasionally ingest foods not observed to be eaten by other adults. By the age of 3 years, young mountain gorillas have developed the basic dietary patterns of adults, in terms of the number of foods eaten, the proportions in which specific foods are consumed, and diet diversity and equitability. There is a strong possibility that chemical cues influence food choice, but their role remains unclear.     

Herbivore feeding preferences in captive and wild populations

Investigations into the mechanisms underpinning plant selection by herbivores have often yielded conflicting results. Such inconsistency might stem from whether selection experiments are conducted with captive or wild populations, and upon the different measures of plant selection used to determine herbivore preference. Here we compared the feeding preferences of captive and wild kangaroos (Macropus fuliginosus) using a standard set of plant species (14 Hakea spp., Proteaceae) and several measures of herbivore selection to examine how environment influenced relative consumption. Three indices of herbivore consumption were measured: number of plants (NP), total shoot volume per plant (TV) and percentage available shoot volume (PV) consumed. NP and TV were closely correlated in the wild and captive populations and consistently correlated with six morphological and chemical plant attributes examined, the most notable being a strong negative correlation with shoot phenolic content. This uniformity suggests that plant selection by captive kangaroos is broadly consistent with that observed in field trials, and consequently that for macropods, at least, captive trials offer a valid way to determine the relative acceptability of different plant species. However, the fact that our third measure of herbivore selection PV was weakly correlated in captive and wild populations and showed no relationship with shoot phenolic content highlights the importance of which measure of plant selection is applied. We suggest that, while NP and TV are potentially confounded by plant size and availability, they offer the clearest insight into plant selection from the point of view of the herbivore, while PV is more suitable for plant‐centred studies.     

A preliminary study of selective visual attention in female mountain gorillas (Gorilla gorilla beringei)

Visually attending to conspecifics can give group-living primates important ecological information, help them to anticipate the behavior of others and to regulate interactions with them, and provide other valuable social information. Variation in the importance and quality of social relationships should influence the way individuals selectively attend to fellow group members. Preliminary data on visual monitoring of conspecifics by wild female mountain gorillas (Gorilla gorilla beringei) show that selective attention mirrors variation in social relationships. Social bonds between males and females are central to gorilla society; correspondingly, females are more likely to stop feeding, and focus their attention on males who walk into view than on females, especially when males give dislays. Females are more likely to focus on other females with whom they have antagonistic relationships than those (mostly close relatives) with whom they have affiliative, cooperative ones. Further research on the context and consequences of visual monitoring could help to address questions about the regulation of social relationships and about social cognition in gorillas.     

Grooming,kinship, and co‐feeding in captive spider monkeys (Ateles geoffroyi)

Allo‐grooming is perhaps the most powerful affiliative behavior observed in non‐human primates. However, the functional significance of grooming in New World monkeys has not yet been fully understood, perhaps because grooming is less frequently observed in platyrrhines. To differentiate the potential role of affiliative investment and/or kinship on sharing access to food (co‐feeding) in spider monkeys, behavioral data on grooming, embracing, and feeding were collected from two different groups of captive study subjects: a familiar/kin group and a non‐familiar/non‐kin group. The results of this study suggest that family‐related spider monkeys that engage in grooming tend to share access to food resources more than unfamiliar conspecifics that do not groom. One explanation for this difference is that the unfamiliar study subjects had not yet invested in the affiliative social network, were not reciprocating their affiliative investments and hence, had a higher tendency toward single animal monopolizing resources. Degree of relatedness alone was not found to be a determinant for sharing the access to food, suggesting that familiarity in spider monkeys is based on the extent to which animals invest in affiliative relationships. In this study, only animals that had engaged in long‐term grooming and recognized each other as familiar shared the access to food. Therefore, it might be likely that in spider monkeys, long‐term grooming of high intensity has to be developed for co‐feeding to occur. Zoo Biol 20:293–303, 2001. © 2001 Wiley‐Liss, Inc.     

Effect of feeding level and feeding frequency on specific dynamic action in Silurus meridionalis

The effect of feeding level ( F _L; 0·5 to 4% dry diet mass per wet fish body mass) and feeding frequency (once every 4 days to twice per day) on postprandial metabolic response was investigated in southern catfish Silurus meridionalis at 27·5° C. The results showed that there was no significant difference in the specific dynamic action (SDA) coefficient among the groups of different feeding levels ( P  > 0·05). The duration increased from 26·0 to 40·0 h and the peak metabolic rate increased from 207·8 to 378·8 mg O₂ kg⁻¹ h⁻¹ when the feeding level was increased from 0·5 to 4%. The relationship between the peak metabolic rate ( R _P, mg O₂ kg⁻¹ h⁻¹) and F _L could be described as: R _P = 175·4 + 47·3 F _L( r ² = 0·943, n  = 40, P  < 0·001). The relationship between the SDA duration ( D , h) and F _L could be described as D =30·97 F _L^0·248 ( r ²=0·729, n =40, P  < 0·001).     

Effects of hand‐rearing on the reproductive success of western lowland gorillas in North America

This study sought to assess the potential effects of hand‐rearing by evaluating the relationships among rearing type and reproductive success in the American Zoo and Aquarium Association's Species Survival Plan^® for western lowland gorillas. Our study included data on 697 gorillas: 257 wild‐born (WB) and 440 born at zoos or related facilities in North America. We found no significant differences in the number of reproductive zoo‐born (ZB) and WB females, but more WB males sired infants than their ZB counterparts. This was influenced by a skew in the number of reproductive years for WB males in the studbook. ZB males showed no difference in infants produced per reproductive year, as compared to WB males, while ZB females produced more infants per reproductive year than did WB females. Mother‐reared (MR), ZB females produced more offspring and used more reproductive opportunity than hand‐reared (HR) females, whereas rearing had no effect on the reproductive success of ZB males. Moreover, MR and partially hand‐reared (PHR) females were more likely to become nurturing mothers themselves. Zoo Biol 21:389–401, 2002. © 2002 Wiley‐Liss, Inc.     

Preliminary investigation of social interactions and feeding behavior in captive group-housed Tasmanian devils (Sarcophilus Harrisii)

As the number of Tasmanian devils (Sarcophilus harrisii) in captivity increases, an understanding of captive social dynamics and behavior is becoming increasingly important. In the wild, devils are solitary, although sometimes, they congregate to feed on a large carcass. However, it is common to house devils in groups as a form of social enrichment. This study investigated how behavior at feeding time of captive Tasmanian devils varied in groups of different sizes. Observations were made of individually housed devils and devils in groups of two, three, five, and six, when presented with a carcass on which to feed. Total feeding duration ranged from 6.5 to 47.4 minutes per observation period (70 minutes). There was no significant interaction between feeding duration and group size during the experiment. Feeding duration varied daily and depended on carcass size. Social housing of Tasmanian devils enabled them to display dyadic and agonistic behaviors during feeding. Observing behaviors and learning from the outcomes of these interactions can improve husbandry techniques. Creating a captive environment that encourages natural behaviors may enhance survival in the wild following translocation.     

The effect of feeding hierarchy on individual variability in daily feeding of rainbow trout, Oncorhynchus mykiss (Walbaum)

Three groups of rainbow trout, Oncorhynchus mykiss , (initial weight 40 g) were fed one of three rations (low, medium or high) for 73 days. Consumption by individual fish within the three ration groups was measured on four occasions (days 27, 55, 64 and 72) using radiography. Food intake by individual fish varied between days and this variability was expressed using the coefficient of variation (CV). Dominant fish within each ration group were defined as individuals with the greater share of the group meal and these fish had low CVs for food intake indicating relatively little variation in daily consumption. By contrast, the fish which had consumed a low mean proportion of the group meal displayed high CVs for food intake, indicating considerable variability in the sizes of individual meals consumed by these fish. As group ration increased, the range of meal sizes and individual CVs in daily feeding decreased, suggesting that the strength of the feeding hierarchy and the variability in individual consumption decreased as food availability increased. It is suggested that radiography can be used to assess social relationships within groups of fish allowing the assessment of feeding hierarchies in larger groups of fish than would be possible by observational techniques.     

Effect of reward level on individual variability in demand feeding activity and growth rate in Arctic charr and rainbow trout

Groups of Arctic charr and rainbow trout were fed by using demand feeders and their individual trigger actuations registered with a PIT-tag (Passive Integrated Transponder) system. Food was supplied at two reward levels, low and high, to five replicate groups of each species for 21 to 27 days. The reward level was defined as the amount of food obtained in response to a single trigger actuation. The effects of reward level on individual demand feeding activity and growth rale were assessed.
As a result of high demand feeding activity, the daily food rations for trout were in excess of their needs at both reward levels. This can be ascribed to the fact that they compensated a low reward level by increasing their bite activity. In contrast, demand feeding activity in charr did not differ significantly between reward levels. Instead, resulting food rations were limiting and excessive, at low and high reward levels, respectively. The variation in bite activity between individuals (measured as their proportional contribution to the total number of trigger actuations within a group) for charr was significantly higher in the low-reward treatment than in the high-reward level. For trout, the variation in bite activity did not differ significantly between treatments. Differences in response to reward level are suggested to be due to the fact that the social hierarchy is weaker in trout than in charr; i.e. the differences in bite activity between dominant and non-dominant individuals are smaller in trout. At both reward levels, the benefit of being dominant, measured in terms of growth rate was significant for charr but non-significant for trout.     

An experimental study of phytoplankton feeding in three tilapiine cichlids

Three species of tilapia ( Oreochromis lidole, O. karongae and O. niloticus ) were fed on four planktonic algal diets in the laboratory. The algal concentrations were measured over a 24-h period and the ingestion rates calculated. In monocultures of large algae (average filament length=477 μm) O. niloticus had the highest ingestion rate, but on small algae (particle size <8 μm) O. lidole was the fastest, while O. karongae enhanced nanoplankton production. With algae of mixed sizes, O. niloticus was able to remove small algae quicker than the other fish species. Thus, the presence of larger algae can influence the impact of a planktivorous species on nanoplankton. O. lidole is likely to have the greatest impact on plankton communities heavily dominated by smaller algae.     

Laboratory study of the grass shrimp feeding preferences

Grass shrimps are very common in various aquatic habitats, and are generally considered omnivorous. But, despite the numerous studies on interspecific trophic interactions, few studies have evaluated their feeding strategy, especially by experimental approaches.In this study, we determined, in the absence of both intra and interspecific competitive interactions, the feeding preferences of the freshwater grass shrimp Palaemonetes antennarius for the most common potential resources in its habitat. Plant and animal items were offered as food to each adult shrimp in multiple-choice laboratory experiments. Resource selection was determined under conditions of low and high food availability and related to the shrimp body size. The role of microbial colonisation in favouring detritivory was also investigated.Results of these experiments showed that P. antennarius consumed only animal items, mostly preferring the isopod Proasellus coxalis. In addition, palatability of the unselected plant remains and non-living prey was not improved by the microbial enrichment. Shrimps also resorted to cannibalism at low ration. By contrast, at high ration they specialised on a narrower range of resources, directing more their feeding efforts towards isopods. The preference ranking of the selected food types was inversely related to the average mass of the prey and varied with the shrimp body size. In particular, the number of live isopods in the diet increased exponentially with the shrimp mass at food high ration, which resulted not to be limiting. In conclusion, food availability influences the food niche of P. antennarius whose feeding strategy in the laboratory is facultative specialisation on dead and live animals rather than omnivory. Ease of handling and/or capture seems to play a role in the food preference ranking.     

Variations in feeding and social behaviour in a captive breeding group of harbour seals (Phoca vitulina)

Most Phocids are thought to reduce their food intake during the breeding and moulting seasons, even though fish are not in short supply. The present study of a captive breeding colony of harbour seals explored the relationship between changes in amount eaten, social behaviour and general activity levels. Even though the animals were given free access to food, over the year the daily amount they ate varied in three cycles. Food eaten fell below average between pupping and mating in June‐July, and fell again in November until early December, for a third time in mid February, and again in early May. Levels surpassed average during and shortly after moulting, and again in late December. Observations made during the summer and early fall revealed that the adult males became more active, and engaged in more social interaction when they were eating less. The female showed no such activity change as a function of appetite. Play occurred primarily when food levels were high. These results are discussed in the context of systems interaction theories of motivation.     

Brief communication: Hand preference for bimanual and unimanual feeding in captive gorillas: extension in a second colony of apes

Right-hand dominance is widely considered to be a uniquely human trait. Whether nonhuman primates exhibit similar population-level hand preferences remains a topic of considerable debate. Despite extensive research focusing on laterality in nonhuman primates, our interpretation of these studies is limited due to methodological issues including the lack of a common measure of hand preference and the use of tasks that may not be reliable indicators of handedness. The use of consistent methods between studies is necessary to enable comparisons within and between species and allow for more general conclusions to be drawn from these results. The present study replicates methods used in recent research reporting population-level right-handedness in captive gorillas (Meguerditchian et al.,2010). Observational data were collected on hand preference for unimanual and bimanual feeding in 14 captive western lowland gorillas (Gorilla gorilla gorilla). Individual-level preferences were found, primarily for bimanual feeding; however, the data reveal no group-level directional bias (contra Meguerditchian et al.). Like the study by Meguerditchian et al. (2010), though, bimanual feeding revealed significantly stronger hand preferences than unimanual reaching, and age, sex, group membership, or rearing history had no effect on hand preference. Finally, variations in diet and corresponding grip type between studies suggest that hand preferences may vary across bimanual tasks depending on grip morphology. This study aims to contribute to our existing knowledge of primate laterality by increasing the number of individuals investigated using methods that allow for comparisons with similar research.