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.     

Nutritional Aspects of Western Lowland Gorilla (Gorilla gorilla gorilla) Diet During Seasons of Fruit Scarcity at Bai Hokou,Central African Republic

Traditionally, gorillas were classified as folivores, yet 15 years of data on western lowland gorillas (Gorilla gorilla gorilla) show their diet to contain large quantities of foliage and fruit, and to vary both seasonally and annually. The consumption of fruit by gorillas at Bai Hokou, Central African Republic, is correlated with rainfall and ripe fruit availability (Remis, 1997a). We investigated the nutritional and chemical content of gorilla foods consumed at Bai Hokou during two seasons of fruit scarcity as measured by phenological observations and compared our findings with the nutrient content of gorilla foods at other African sites. We conclude that during lean times, Bai Hokou gorillas consumed fruits with higher levels of fiber and secondary compounds than those of other populations of western lowland or mountain gorillas. Conversely, leaves consumed by Bai Hokou gorillas were relatively low in fiber and tannins. Bai Hokou gorillas appeared to meet their nutritional needs by eating a combination of fruit and foliage. They ate fruits comparatively high in secondary compounds and fiber when necessary. While gorillas are selective feeders, wherever and whenever preferred foods are scarce, their large body size and digestive anatomy enable them to consume and process a broader repertoire of foods than smaller bodied-apes.     

Winter browsing on Alaska feltleaf willow twigs improves leaf nutritional value for snowshoe hares in summer

In boreal forests, browsing by mammals on winter-dormant twigs increases leaf nitrogen, leaf greenness, and leaf size. This suggests browsing reduces competition among meristems for mineral nutrients, and in particular, competition for nitrogen. Winter browsing also reduces the shoot carbohydrate reserves used by leaves to produce condensed tannin. These effects of winter browsing are predicted to improve the nutritional value of leaves for mammals because they increase the mass of digestible nitrogen in leaves. This hypothesis was tested using Alaska feltleaf willow and the snowshoe hare as the experimental system. Six in vivo indicators of leaf nutritional quality were used to compare leaves from winter-browsed plants with leaves from unbrowsed plants. The indicators used were dry matter intake, nitrogen intake, condensed tannin intake, dry matter digestibility, apparent digestibility of nitrogen and nitrogen retention. The results obtained were in agreement with the above hypothesis. In early summer, at the time snowshoe hares and other northern herbivores reproduce, hares fed leaves from browsed plants consumed more nitrogen, digested more of the nitrogen they consumed, and retained more of the nitrogen they digested than did hares fed leaves from unbrowsed plants. The high nitrogen content and low tannin content of leaves from browsed plants may explain this browsing caused increase in leaf nutritional value. How these positive effects of winter browsing on snowshoe hare nutrition at the time of reproduction might affect hare population dynamics are briefly discussed.     

Galagos and Gummivory

An experimental program investigating the role of gum in the diet of captiveGalago senegalensis braccatus is reviewed. One set of studies investigated the role of gum chemistry in dietary preferences. The animals showed a preference forAcacia gums over gums from other plants. If condensed or hydrolized tannins were added to theAcacia gums, such gums were consumed at a lower rate than plainAcacia gum. This effect was stronger with the condensed (quebracho) tannin. The effect of such secondary compounds on gum consumption in wild galagos needs investigation. The second set of studies examined digestion of gums, which are complex polysaccarides probably requiring fermentation in the gut. Substitution of gum for simple carbohydrate sources in the diet lead to increased gut transit time. On the gum diet, animals maintained weight without an increase in intake or a decrease in activity levels. Apparent dry matter digestibility of the two diets was equivalent. Indicators of gut bacterial action increased on the gum diet. These observations are strong indirect evidence for an increase of fermentation in the gut on the gum diet.     

Feeding Ecology of Cross River Gorillas (Gorilla gorilla diehli) at Mawambi Hills: The Influence of Resource Seasonality

Determining the composition of primate diet and identifying factors that affect food choice are important in understanding habitat requirements of primates and designing conservation plans. We studied the diet of Cross River gorillas (Gorilla gorilla diehli) in relation to availability of food resources, in a semideciduous lowland forest site (Mawambi Hills) in Cameroon, from November 2009 to September 2011. Based on 109 d of feeding trail data, 203 fecal samples, and 22 mo of phenological monitoring, we determined that gorillas consumed a total of 242 food items, including 240 plant items from 186 species and 55 taxonomic families. Mawambi gorillas diversified fruit consumption when fruit availability increased, and consumed more fibrous foods (pith, leaf, bark) during times of fruit scarcity, consistent with results of other gorilla studies. However, fruit availability was not related to rainfall, and the period of fruit scarcity was more pronounced at Mawambi than at other gorilla study sites, due to a single long dry season and extreme rainfall at the end of the rainy season that delayed fruit production and ripening. We found no relationship between the daily path length of the gorillas and fruit consumption. We found feeding habits of Mawambi gorillas to be notably similar to those of a population of Cross River gorillas at Afi Mountain, Nigeria, although subtle differences existed, possibly due to site-specific differences in forest composition and altitude. At both sites the liana Landolphia spp. was the single most important food species: the leaves are a staple and the fruits are consumed during periods of fruit scarcity. Snails and maggots were consumed but we observed no further faunivory. We suggest that tree leaves and lianas are important fallback food sources in the gorilla diet in seasonally dry forests.     

Western lowland gorillas (Gorilla gorilla gorilla) change their activity patterns in response to frugivory

The most important environmental factor explaining interspecies variation in ecology and sociality of the great apes is likely to be variation in resource availability. Relatively little is known about the activity patterns of western lowland gorillas (Gorilla gorilla gorilla), which inhabit a dramatically different environment from the well‐studied mountain gorillas (G. beringei beringei). This study aims to provide a detailed quantification of western lowland gorillas' activity budgets using direct observations on one habituated group in Bai Hokou, Central African Republic. We examined how activity patterns of both sexes are shaped by seasonal frugivory. Activity was recorded with 5‐min instantaneous sampling between December 2004 and December 2005. During the high‐frugivory period the gorillas spent less time feeding and more time traveling than during the low‐frugivory period. The silverback spent less time feeding but more time resting than both females and immatures, which likely results from a combination of social and physiological factors. When compared with mountain gorillas, western lowland gorillas spend more time feeding (67 vs. 55%) and traveling (12 vs. 6.5%), but less time resting (21 vs. 34%) and engaging in social/other activities (0.5 vs. 3.6%). This disparity in activity budgets of western lowland gorillas and mountain gorillas may be explained by the more frugivorous diet and the greater dispersion of food resources experienced by western lowland gorillas. Like other apes, western lowland gorillas change their activity patterns in response to changes in the diet. Am. J. Primatol. 71:91–100, 2009. © 2008 Wiley‐Liss, Inc.     

Feeding ecology of western lowland gorillas in the Nouabalé-Ndoki National Park,Congo

The feeding ecology of western lowland gorillas (Gorilla gorilla gorilla) living in the Nouabalé-Ndoki National Park, northern Congo, was surveyed for one full year. This is the first record to make clear the seasonal changes in the feeding habits of gorillas in a whole year, living in the primary lowland forest almost completely undisturbed. Fecal contents, feeding traces, and direct observation were analyzed with reference to a fruit availability survey. Although the gorillas fed largely on fruits in the forest, their basic diet was fibrous parts of plants, including shoots, young leaves, and bark. Terrestrial herbaceous vegetation, such as monocotyledons of the Marantaceae and aquatic herbs having much protein content and minerals, were frequently eaten even in the fruiting season. As these highly nutritious fibrous foods were superabundant all year, the major foods of the Ndoki gorillas seemed to be those plants. However, they selected fruits as their alternative food resources in the fruiting season. Gorillas foraged on many fruit species, while showing strong preferences for some particular species. The swamp forest, including marshy grasslands, was an important and regular habitat for the Ndoki gorillas.     

Minerals in the Foods Eaten by Mountain Gorillas (Gorilla beringei)

Minerals are critical to an individual’s health and fitness, and yet little is known about mineral nutrition and requirements in free-ranging primates. We estimated the mineral content of foods consumed by mountain gorillas (Gorilla beringei beringei) in the Bwindi Impenetrable National Park, Uganda. Mountain gorillas acquire the majority of their minerals from herbaceous leaves, which constitute the bulk of their diet. However, less commonly eaten foods were sometimes found to be higher in specific minerals, suggesting their potential importance. A principal component analysis demonstrated little correlation among minerals in food items, which further suggests that mountain gorillas might increase dietary diversity to obtain a full complement of minerals in their diet. Future work is needed to examine the bioavailability of minerals to mountain gorillas in order to better understand their intake in relation to estimated needs and the consequences of suboptimal mineral balance in gorilla foods.     

Diet of Grauer's Gorillas in the Montane Forest of Kahuzi,Democratic Republic of Congo

We describe the diet of a semihabituated group of Grauer's gorillas (Gorilla beringei graueri) inhabiting the montane forest of Kahuzi-Biega National Park, Democratic Republic of Congo, based on direct observations, feeding remains in their fresh trails, and fecal samples collected over 9 yr. We examined fruit availability in their habitat; consumption of fruit, vegetative, and animal food; and daily intake of vegetative plant food using a transect, fruit monitoring trails, fecal analysis, and tracing of the animal's daily trails between consecutive nest sites. The fruit food repertoire of Kahuzi gorillas resembles that of western and eastern lowland gorillas inhabiting lowland tropical forests, while their vegetative food repertoire resembles that of mountain gorillas inhabiting montane forests. Among 236 plant foods (116 species), leaves, pith, and barks constitute the major parts (70.2%), with fruit making up the minor part (19.7%). About half (53.2%) of the total fecal samples included fruit remains. The gorillas used leaves, stems and other vegetative plant parts as staples. Their fruit intake was similar to that reported for mountain gorillas in Bwindi. They ate animal foods, including earthworms, on rare occasions. Variation in fruit consumption was positively associated with variation in fruit production. The gorillas ate fig fruits frequently; fig intake is positively correlated with that of other fruits, and figs were not fallback foods. They relied heavily on bamboo shoots on a seasonal basis; however, no bamboo shoots were available for several years after a major flowering event. Our results support the argument that variation in gorilla diets mostly reflects variation in vegetational composition of their habitats.     

Condensed tannins in the diets of primates: a matter of methods?

To understand the ways in which condensed tannins (CT) affect primate diet selection and nutritional status, correct measurements are essential. In the majority of studies of the CT contents of primate foods, a tannin source such as "quebracho" is used to standardize CT assays, but the CT in quebracho tannin may not be similar to those in the plants of interest. We investigated how the choice of standard to calibrate CT assays affects the estimation of CT in the diets of mountain gorillas (Gorilla beringei). We purified the CT from gorilla foods and compared the actual amounts of CT in the foods with estimates produced by using the quebracho tannin. When quebracho was used, the estimates of CT contents of gorilla foods were, on average, 3.6 times the actual content of CT so that the amounts in frequently eaten gorilla foods were substantially overestimated. The overestimation for a given plant could not be predicted reliably and the ranking of plants by tannin content differed according to the standard used. Our results demonstrate that accurate measurements of CT necessitate the use of tannins purified from the plant species of interest. A reevaluation of primatology studies using interspecific comparisons of tannin content will provide new insights into primate food selection and nutritional ecology.     

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. Am. J. Primatol. 70:927–938, 2008. © 2008 Wiley‐Liss, Inc.     

Ecological Constraints on the Foraging Effort of Western Gorillas (Gorilla gorilla gorilla) at Bai Hoköu, Central African Republic

Recent studies demonstrate that western lowland gorillas incorporate much more fruit into their diet than Virunga mountain gorillas do. Very little is known, however, about how the frugivorous behavior of western gorillas influences their daily ranging behavior, which may ultimately affect social factors such as group size and structure. I examined the influence of diet and the spatiotemporal availability of plant foods on the foraging effort of nonhabituated western lowland gorilla groups during 17 months at Bai Hoköu in the Dzanga-Ndoki National Park, Central African Republic. I determined diet from indirect methods and gorilla plant food availability and spatial distribution from phenology and line transects. Daily path length gives an estimate of foraging effort and was the distance paced, following fresh gorilla trails, from morning to evening nest sites. The availability and distribution of fruit and its consumption by gorillas varied seasonally. When concentrating on fruits, gorillas traveled significantly farther (mean = 3.1 km/day) than when their diet consisted mostly of nonfruit vegetation, such as leaves and woody pith, stems, and bark (mean = 2.1 km/day). The amount of herbaceous vegetation in the diet did not vary seasonally and did not influence daily path length. The best environmental predictor of foraging effort was fruit density, or a measure combining both density and spatial pattern: coefficient of dispersion. In addition, when fruit patches were small, path length tended to increase but not significantly. Compared with results of other studies, gorillas at Bai Hoköu travel farther (mean = 2.6 km/day) than gorillas in Gabon (mean = 1.7 km/day) and five times farther than mountain gorillas in the Virungas (mean = 0.5 km/day). Increased foraging effort of gorillas in this region, especially during the fruiting season, may have profound effects on group size and structure.     

Foliage acceptability to browsing ruminants in relation to seasonal changes in the leaf chemistry of woody plants in a South African savanna

Summary We investigated seasonal changes in food selection by hand-reared kudus and impalas in savanna vegetation in northern Transvaal, South Africa. The acceptability of the leaves of woody plants to these animals was compared with leaf concentrations of nutrients, fibre components and old leaf phenophases. No consistently significant correlation was found between acceptability and any single chemical factor. Based on an a priori palatability classification, discriminant function analysis separated relatively palatable species from unpalatable species in terms of a linear combination of protein and condensed tannin concentrations. The high acceptability of certain otherwise unpalatable species during the new leaf phenophase was related to elevation of protein levels relative to condensed tannin contents. Species were added to the diet during the dry season approximately in the order of their relative protein-condensed tannin difference.     

Dietary adaptations of temperate primates: comparisons of Japanese and Barbary macaques

Habitat, diet and leaf chemistry are compared between Japanese and Barbary macaques to reveal the similarities and differences in dietary adaptations of temperate primates living at the eastern and western extremes of the genus Macaca. Tree species diversity and proportion of fleshy-fruited species are much higher in Japan than in North Africa. Both species spend considerable annual feeding time on leaves. Japanese macaques prefer fruits and seeds over leaves, and Barbary macaques prefer seeds. These characteristics are adaptive in temperate regions where fruit availability varies considerably with season, since animals can survive during the lean period by relying on leaf and other vegetative foods. The two species are different with respect to the higher consumption of herbs by Barbary macaques, and the leaves consumed contain high condensed and hydrolysable tannin for Barbary but not for Japanese macaques. Barbary macaques supplement less diverse tree foods with herbs. Because of the low species diversity and high tannin content of the dominant tree species, Barbary macaques may have developed the capacity to cope with tannin. This supports the idea that digestion of leaves is indispensable to survive in temperate regions where fruit and seed foods are not available for a prolonged period during each year.     

Mountain gorilla geophagy: A possible seasonal behavior for dealing with the effects of dietary changes

Geophageous Rwandan mountain gorillas excavate and eat weathered leucite-rich regolith (subsoil C horizons) from the slopes of Mount Visoke in the Virunga Mountains. In the months of the dry season, the gorillas reportedly ingest a halloysitic natural earth having a chemical composition similar to that of Kaopectate, a pharmaceutical used by humans to treat gastrointestinal upsets. Several plants known to contain potential toxins are consumed more heavily by gorillas in these months. New information from geochemical and mineral analyses suggests that geophagy may alleviate intestinal problems associated with changes in their diet because the ingested weathered regolith,containing halloysitic clay minerals, may act as a pharmaceutical agent that helps to adsorb toxins and to control dehydration in the dry season.     

Mature leaf selection of Japanese macaques: effects of availability and chemical content

Mature leaf food selection of Japanese macaques was studied in two different altitudinal zones of Yakushima: in the coniferous (1000–1200 m) and coastal forests (0–200 m). Logistic regression analysis was conducted to reveal the effect of chemical [neutral detergent fibre (NDF), crude protein, crude ash, crude lipid, condensed tannin, hydrolysable tannin and crude protein/NDF ratio] and availability properties (density of trees and total basal area) on the macaques' choice between the major food leaves and the rarely eaten or non-food leaves. In both forests, macaques selected leaves with a high crude ash and crude protein content, or a high crude protein/NDF ratio, as major foods. In the coniferous forest, macaques selected leaves with less condensed tannin, but this tendency was absent in the coastal forest. This was because macaques in the coniferous forest suffer from a greater risk of ingesting large amounts of condensed tannins, as their feeding time on mature leaves is seven times as long as that in the coastal forest. Among the chemical and availability properties, stepwise multiple regression analysis revealed that the number of trees (ha⁻¹) was the only significant factor explaining the variations in feeding time among major food leaves, both in the coniferous and coastal forests. The present results suggest that the effect of travelling cost, which can be reduced by selecting common trees, exceeds the benefits gained by selecting high-quality foods.     

Day-journey length and daily diet of solitary male gorillas in lowland and highland habitats

We compared day-journey length and daily diets of solitary male gorillas in lowland versus highland habitats. Solitary males in tropical forests of Zaire tend to travel longer distances, to visit more types of vegetation, and to consume more kinds of food than a solitary male mountain gorilla in the Virunga Volcanoes did. The number of feeding sites per day is larger and the mean distance between feeding sites is far longer for the former than the latter. These observations may reflect differences in food breadth and availability between highland and lowland habitats. The herbaceous plants that are eaten by mountain gorillas are densely and evenly distributed in the higher montane forest of the Virungas, where gorillas need not cover long distances to search for food. In contrast, herbaceous plants are scarce in primary and ancient secondary forests of lowland habitats, where gorillas travel long distances and eat various fruits and insects. The patchy and unpredictable distribution of foods may extend the distances over which gorillas search for food in the lowland habitat. However, solitary males showed a prominent reduction in day-journey length and changed their choices of food during the nonfruiting season (the long rainy and dry seasons) in the lowland habitats. This strategy may have developed during the Pleistocene and may have enabled them to enlarge their ranges to the higher montane forests, where fruits are sparse throughout the year.     

Diet and seasonal changes in sympatric gorillas and chimpanzees at Kahuzi–Biega National Park

Based on 8 years of observations of a group of western lowland gorillas (Gorilla beringei graueri) and a unit-group of chimpanzees (Pan troglodytes schweinfurthii) living sympatrically in the montane forest at Kahuzi–Biega National Park, we compared their diet and analyzed dietary overlap between them in relation to fruit phenology. Data on fruit consumption were collected mainly from fecal samples, and phenology of preferred ape fruits was estimated by monitoring. Totals of 231 plant foods (116 species) and 137 plant foods (104 species) were recorded for gorillas and chimpanzees, respectively. Among these, 38% of gorilla foods and 64% of chimpanzee foods were eaten by both apes. Fruits accounted for the largest overlap between them (77% for gorillas and 59% for chimpanzees). Gorillas consumed more species of vegetative foods (especially bark) exclusively whereas chimpanzees consumed more species of fruits and animal foods exclusively. Although the number of fruit species available in the montane forest of Kahuzi is much lower than that in lowland forest, the number of fruit species per chimpanzee fecal sample (average 2.7 species) was similar to that for chimpanzees in the lowland habitats. By contrast, the number of fruit species per gorilla fecal sample (average 0.8 species) was much lower than that for gorillas in the lowland habitats. Fruit consumption by both apes tended to increase during the dry season when ripe fruits were more abundant in their habitat. However, the number of fruit species consumed by chimpanzees did not change according to ripe fruit abundance. The species differences in fruit consumption may be attributed to the wide ranging of gorillas and repeated usage of a small range by chimpanzees and/or to avoidance of inter-specific contact by chimpanzees. The different staple foods (leaves and bark for gorillas and fig fruits for chimpanzees) characterize the dietary divergence between them in the montane forest of Kahuzi, where fruit is usually scarce. Gorillas rarely fed on insects, but chimpanzees occasionally fed on bees with honey, which possibly compensate for fruit scarcity. A comparison of dietary overlap between gorillas and chimpanzees across habitats suggests that sympatry may not influence dietary overlap in fruit consumed but may stimulate behavioral divergence to reduce feeding competition between them.     

Fracture toughness of mountain gorilla (Gorilla gorilla beringei) food plants

Mountain gorillas, the largest extant primates, subsist almost entirely on plant matter. Moreover, their diet includes a substantial amount of structural material, such as bark and stems, which other primates tend to avoid. Accordingly, the robust masticatory apparatus of gorillas may be adaptive to this presumably tough diet; however, quantitative information on this subject is lacking. In this study the fracture toughness of mountain gorilla foods was examined for the first time. Samples of 44 food plants from Bwindi-Impenetrable National Park (BINP) and Mgahinga Gorilla National Park (MGNP) were tested. These parks are inhabited by two gorilla populations that regarded by some as being distinct at the subspecific taxonomic level. Although food toughness did not differ between the two populations, both diets contained tough items. Tree barks were the toughest food items (varying from 0.23 to 8.2 kJ/m2), followed by shrub barks, pith, and stems. The toughness of leaves and fruit was negligible compared to that of bark. The toughness of bamboo was low in comparison to the toughest food items. Accordingly, the prominent toughness of bark, pith, and stems may be key factors in the evolution of orofacial robusticity in mountain gorillas.     

Nutritional geometry: gorillas prioritize non-protein energy while consuming surplus protein

It is widely assumed that terrestrial food webs are built on a nitrogen-limited base and consequently herbivores must compensate through selection of high-protein foods and efficient nitrogen retention. Like many folivorous primates, gorillas' diet selection supports this assumption, as they apparently prefer protein-rich foods. Our study of mountain gorillas (Gorilla beringei) in Uganda revealed that, in some periods, carbohydrate-rich fruits displace a large portion of protein-rich leaves in their diet. We show that non-protein energy (NPE) intake was invariant throughout the year, whereas protein intake was substantially higher when leaves were the major portion of the diet. This pattern of macronutrient intake suggests that gorillas prioritize NPE and, to achieve this when leaves are the major dietary item, they over-eat protein. The concentrations of protein consumed in relation to energy when leaves were the major portion of the diet were close to the maximum recommended for humans and similar to high-protein human weight-loss diets. By contrast, the concentrations of protein in relation to energy when gorillas ate fruit-dominated diets were similar to those recommended for humans. Our results question the generality of nitrogen limitation in terrestrial herbivores and provide a fascinating contrast with human macronutrient intake.