Leaf chemistry as a predictor of primate biomass and the mediating role of food selection: a case study in a folivorous lemur (Propithecus verreauxi).

Folivorous primate biomass has been shown to positively correlate with the average protein-to-fiber ratio in mature leaves of tropical forests. However, studies have failed to explain the mismatch between dietary selection and the role of the protein-to-fiber ratio on primate biomass; why do not folivores always favor mature leaves or leaves with the highest protein-to-fiber ratio? We examined the effect of leaf chemical characteristics and plant abundance (using transect censuses; 0.37 ha, 233 trees) on food choices and nutrient/toxin consumption in a folivorous lemur (Propithecus verreauxi) in a gallery forest in southern Madagascar. To assess the nutritional quality of the habitat, we calculated an abundance-weighted chemical index for each chemical variable. Food intake was quantified using a continuous count of mouthfuls during individual full-day follows across three seasons. We found a significant positive correlation between food ranking in the diet and plant abundance. The protein-to-fiber ratio and most other chemical variables tested had no statistical effect on dietary selection. Numerous chemical characteristics of the sifaka's diet were essentially by-products of generalist feeding and "low energy input/low energy crop" strategy. The examination of feeding behavior and plant chemistry in Old World colobines and folivorous prosimians in Madagascar suggests that relative lack of feeding selectivity and high primate biomass occur when the average protein-to-fiber ratio of mature leaves in the habitat exceeds a threshold at 0.4.

Age-dependent patterns of intensive observation on elders by free-ranging juvenile Japanese macaques (Macaca fuscata yakui) within foraging context on Yakushima.

Early learning about edible food in the environment is a critical survival task for young nonhuman primates. Social learning and social facilitation are often cited to explain how youngsters learn to select and find their food. In this framework, we observed eight mother-youngster pairs of free-ranging Japanese macaques divided into two sets according to the age of the young (infants aged between 7 and 12 months and juveniles aged between 1.5 and 2 years) during three winter months. We systematically investigated the intensive observation directed by the youngsters toward elders by recording the target's identity (e.g. mother, subadult), the items manipulated by the elder and those items closely observed by the youngster, along with the behavior of the youngster preceding and immediately following an intensive observation period. The diet of the mothers and juveniles was estimated from time records of each feeding occurrence for each food item (identified to species level) and from the quantity of fresh matter ingested. The results show that intensive observation by both infants and juveniles were directed toward those elders engaged in plant and invertebrate foraging. Such behavior was age-dependent, being more frequent in infants than in juveniles. The majority of the intensive observations were directed toward the mother. Intensive observations also shaped a change in the behavior of infants by significantly stimulating the investigation of food items and locations otherwise not investigated by juveniles. Moreover, infants showed a particular interest in rare food items and especially invertebrates. Age differences between the two sets of young and their interest in rare foods are discussed with reference to the occurrence of intensive observation within the framework of kin relationships, social organization, and social transmission of information about food type and food location and its survival values.

Feeding behavior of lactating brown lemur females (Eulemur fulvus) in Mayotte: influence of infant age and plant phenology.

Altmann [Baboon Mothers and Infants, University of Chicago Press, 1980] and Dunbar and Dunbar [Animal Behavior 36:970-980, 1988] provided a model that predicts the amount of time spent feeding by lactating baboon females, as related to infant age. Dunbar's model further suggests that food quality affects the amount of time that females devote to feeding activity, and is predictable from rainfall and temperature data. In this study the model was tested with data recorded from births of the Mayotte brown lemur from four maternal dyads (Eulemur fulvus). This study also examines the correlation between female activity budget, quantities of fresh plant matter ingested, and suckling duration using data collected from focal animal samples on the mother-infant dyads. The relationships among rainfall, food supply, and food quality were also tested. It appears that female brown lemurs do not devote more time to feeding during the infant growth period. The data show that female brown lemurs increased their food intake during the early-lactating period when the frequency of suckling is the highest, and before infants begin to eat substantial amounts of solid foods. Thus, the frequency of feeding reflects the cost of lactation better than suckling duration. Furthermore, females did not appear to select foods according to their availability or rainfall levels. I hypothesize that the lack of convergence between lemur data and baboon observations is due to differences between their respective environments and their feeding ecology. These data also indicate that the most significant lactating cost for the brown lemur occurs during the early lactation period.

Cathemerality in the Mayotte brown lemur (Eulemur fulvus): seasonality and food quality.

In past decades, cathemerality--as defined by Tattersall [1987]--has been documented in two primate families: Cebidae and Lemuridae. In the Lemuridae, in particular the genus Eulemur, cathemeral activity seems to be a regular behavioural trait. Nevertheless, ultimate and proximate determinants responsible for this behaviour remain unclear. In this study, in a dry and deciduous forest on Mayotte (Comoro Archipelago), activities of 4 female brown lemurs (Eulemur fulvus fulvus) were recorded by focal animal sampling during the daylight period and by scan sampling on their respective groups during the night. Horizontal distances travelled by females and groups were measured using GPS. During the daylight period, food intakes were estimated in grams by extrapolation of counting of mouthfuls after weighing a large sample of plant parts eaten. Crude protein, crude lipid, soluble sugar and crude fibre were analyzed for each seasonal reconstituted diet. Records of temperature and rainfall were supplied by a local meteorological station. Observations confirmed cathemerality in the Mayotte brown lemur as reported by Tattersall in 1977. During the dry season, the animals increased their nocturnal activity--substantially increasing the time devoted to feeding and moving overall, but especially at night--and were less active during the daylight period. The quality of their diet in the dry season was poorer than that in the wet season, with soluble sugar content and protein content decreasing and fibre content increasing slightly. As a result, Mayotte brown lemurs may need to extend their foraging activity over the 24-hour cycle to balance nutritional requirements.