Ecological correlates to social structure in two lemur species in Madagascar

In this study, I tested two hypotheses regarding the relationship of ecological variables (size, density, and distribution of patches) and infant developmental patterns to lemur social structure using two prosimian primates in Ranomafana, Madagascar: the rufous lemur (Eulemur fulvus rufus) and the red-bellied lemur (Eulemur rubriventer). Three predictions regarding the general effects of patch size and subgroup size on lemur feeding rates were supported: (1) Rufous lemurs used large patches; red-bellied lemurs used smaller patches; (2) larger subgroups of rufous lemurs used larger patches; and (3) rufous lemur feeding rates decreased significantly with increases in subgroup size and patch size, whereas size and patch size had no significant effect on red-bellied lemur feeding rates. However, food item size (fruit) had a more significant effect on rufous and red-bellied lemur feeding rates than either patch size or subgroup size. When similar-sized fruits were compared, rufous lemur feeding rates on small fruit were most affected by patch size, yet feeding rates on medium-sized fruit were most affected by subgroup size. Neither lemur species used patches in consistent ways seasonally. During periods of food abundance, rufous lemurs used many small, common, and clumped patches. In food scarcity periods, they used fewer, larger, rarer, and less clumped patches; groups migrated when food became most scarce. Red-bellied lemurs also used patches in variable ways, but these patterns were not linked with food availability. Finally, infant development patterns differed between lemur species; red-bellied lemur males cared for offspring and infants reached developmental landmarks faster than rufous lemur infants. Therefore, red-bellied lemur group size may be constrained by the need for additional infant care by other group members. In contrast, rufous lemur group size may be constrained by patch availability during the most critical period of food scarcity. © 1996 Wiley-Liss, Inc.     

Effect of habitat structure on positional behavior and support use in three species of lemur

This study compares locomotor and postural behavior and substrate use of three species of lemur, the diademed sifaka (Propithecus diadema edwardsi), the brown lemur (Eulemur fulvus rufus), and the red-bellied lemur (Eulemur rubriventer) at two different localities within Ranomafana National Park. The object of the study is to see if there are quantitative differences in the behavior of the lemurs or their choice of substrates in forests that have different structural attributes. Analysis of the physical characteristics of the habitat demonstrates that compared to the Talatakely area, the forest at Vatoharanana has a higher proportion of larger, taller trees. The behavior of the lemurs also differs in the two areas: all species leap less and climb and move quadrupedally more at Vatoharanana. All species use small size supports less frequently at Vatoharanana, choosing insted medium size supports (all three species) or tiny supports (Propithecus andE. fulvus) found in tree crowns and terminal branches. The lemurs prefer (i.e. use more often than would be expected based on abundance) large trees at both sites. At Vatoharanana however, they are more frequently observed higher in trees and in taller trees with greater trunk breadth. The differences in locomotor behavior are in part due to the fact that at Vatoharanana, more bouts are collected during feeding and foraging than during travel. The relationship between this difference in activity pattern and the structural differerences in the two forests, however, is not clear. This study points out the need for longer term field studies of positional behavior and substrate use that incorporate the variety of forest types the subject species inhabit.     

Better few than hungry: flexible feeding ecology of collared lemurs Eulemur collaris in littoral forest fragments

Background

Frugivorous primates are known to encounter many problems to cope with habitat degradation, due to the fluctuating spatial and temporal distribution of their food resources. Since lemur communities evolved strategies to deal with periods of food scarcity, these primates are expected to be naturally adapted to fluctuating ecological conditions and to tolerate a certain degree of habitat changes. However, behavioral and ecological strategies adopted by frugivorous lemurs to survive in secondary habitats have been little investigated. Here, we compared the behavioral ecology of collared lemurs (Eulemur collaris) in a degraded fragment of littoral forest of south-east Madagascar, Mandena, with that of their conspecifics in a more intact habitat, Sainte Luce.

Methodology/Principal Findings

Lemur groups in Mandena and in Sainte Luce were censused in 2004/2007 and in 2000, respectively. Data were collected via instantaneous sampling on five lemur groups totaling 1,698 observation hours. The Shannon index was used to determine dietary diversity and nutritional analyses were conducted to assess food quality. All feeding trees were identified and measured, and ranging areas determined via the minimum convex polygon. In the degraded area lemurs were able to modify several aspects of their feeding strategies by decreasing group size and by increasing feeding time, ranging areas, and number of feeding trees. The above strategies were apparently able to counteract a clear reduction in both food quality and size of feeding trees.

Conclusions/Significance

Our findings indicate that collared lemurs in littoral forest fragments modified their behavior to cope with the pressures of fluctuating resource availability. The observed flexibility is likely to be an adaptation to Malagasy rainforests, which are known to undergo periods of fruit scarcity and low productivity. These results should be carefully considered when relocating lemurs or when selecting suitable areas for their conservation.     

Group Size, Home Range Use, and Seasonal Variation in the Ecology of Eulemur mongoz

We collected data during a 10-month study carried out on the mongoose lemur, Eulemur mongoz, at Anjamena in northwestern Madagascar, which provide baseline information on seasonal variation in the ecology, home range use and some aspects of the behavior of two neighboring groups. We monitored group size of nine groups in the study area and assessed them for seasonal variation. We present additional information collected during short-term surveys in other areas before and during the study for comparison. The study groups were small family units, and changes in group size were limited to births and emigrations of sexually mature progeny. In spite of clear seasonal changes in climate and vegetation, there is no variation in grouping patterns, so it is not possible to correlate variation in group size with seasonal variation of ecological variables. Comparison with ecological data from other field studies on lemurids reveals differences in food resource distribution in western forests versus other types of Malagasy forest. This distribution of food resources may predict home range size in mixed frugivorous–folivorous lemurs. Small home ranges, mainly in the West, could be correlated with a uniform distribution of food resources. Finally, we suggest that the dry season in the West may not present frugivorous–folivorous lemurs with major problems in finding an adequate food supply. This is supported by the lack of seasonal differences in ranging behavior of mongoose lemurs.     

Feeding enrichment by self‐operated food boxes for white‐fronted lemurs (Eulemur fulvus albifrons) in the Masoala exhibit of the Zurich Zoo

In the new Masoala exhibit of the Zurich Zoo four self‐operated food boxes were installed to encourage arboreal behavior and higher activity levels, and to increase the attractiveness to visitors of a group of three white‐fronted lemurs (Eulemur fulvus albifrons) and one Alaotran gentle lemur (Hapalemur griseus alaotrensis). Data obtained by direct observations with and without food boxes present were compared. In addition, visitors were surveyed to investigate attractiveness of the lemurs. Overall activity and locomotor behavior increased due to food box presentation. Furthermore, the visitor survey documented that the lemurs were spotted more often in trees when the food boxes were present. Because behavior patterns of the subjects approached natural levels with food boxes, the presentation of self‐operated food boxes seems a valuable tool to improve the captive environment of lemurs. Zoo Biol 0:1–10, 2005. © 2005 Wiley‐Liss, Inc.     

Maintenance of female dominance in blue‐eyed black lemurs (Eulemur macaco flavifrons) and gray bamboo lemurs (Hapalemur griseus griseus) under semi‐free‐ranging and captive conditions

Several species of Malagasy prosimians are characterized by female dominance, an unusual trait among mammals. We compare the extent to which female dominance is displayed and the mechanisms that are used to maintain dominance in the frugivorous blue‐eyed black lemurs (Eulemur macaco flavifrons) and the folivorous gray bamboo lemurs (Hapalemur griseus griseus) housed at the Duke Lemur Center. All dominant–submissive interactions were recorded during 448 hr of focal animal observations. Both species of lemurs exhibited clear patterns of female dominance. However, the two species used aggressive dominance (defined as aggression+submission) and social dominance (defined as submission in the absence of aggression) to different extents in maintaining hierarchies within each group. The adult female blue‐eyed black lemurs used aggressive dominance (e.g., chase, cuff, bite) in more of their dominance interactions (66%) than did the adult female gray bamboo lemurs (40%). In both species, rates of aggressive dominance interactions were higher during feeding versus nonfeeding periods and while in smaller outdoor runs versus larger natural habitat enclosures, but the differences were not significant. Overall blue‐eyed black lemurs exhibit a more aggressive form of female dominance compared to the gray bamboo lemur. Zoo Biol 26:345–361, 2007. © 2007 Wiley‐Liss, Inc.     

Habitat shifting by the common brown lemur (Eulemur fulvus fulvus): a response to food scarcity

During periods of food scarcity, most primates display behavioral responses, such as dietary switching or adjustment of traveling and foraging efforts, within home ranges. In rare cases, several primate species leave their home ranges for other remote habitats to seek alternative resources; this migration-like behavior is termed “habitat shifting.” Reports of habitat shifting have concentrated on platyrrhines, but this behavior has rarely been observed among prosimians. During 1 year of observation of a troop of common brown lemurs (Eulemur fulvus fulvus) in Ankarafantsika National Park, northwestern Madagascar, habitat shifting occurred twice. To understand the causes of this behavior, I examined the seasonal availability of fruit resources in the range continuously used by the troop during the year (defined as the annual range) and compared feeding activities and vegetation types between the annual range and new areas. The troop usually stayed within a 38.7-ha annual range, defined by a 95 % fixed kernel analysis based on GPS location data collected at 5-min intervals. In April 2007, the lemurs suddenly moved to a habitat 1.0–1.5 km south of their annual range and concentrated on the consumption of Grewia triflora fruits for 2 weeks. In November 2007, they visited a habitat 0.8–1.7 km southeast of the annual range and exploited fruits of Landolphia myrtifolia. These new areas were open habitats with high densities of the respective fruit species. The density of fruiting trees was low in the annual range during these periods; thus, habitat shifting to areas with different phenological productivity appeared to be an effective response to fruit scarcity. Brown lemurs are generally categorized as a nonterritorial species, and the lemurs observed here showed no agonistic behavior in intergroup encounters during range shifting. Such nonterritoriality may allow brown lemurs to shift habitats, a behavior resulting in long-term absence from their annual range.     

Comparison of Reproductive Characteristics and Changes in Body Weight Between Captive Populations of Rufous and Gray Mouse Lemurs

The nocturnal Malagasy mouse lemurs are among the smallest primates worldwide. Several sibling species are known. Of these, the rufous and the gray mouse lemur differ with respect to morphology, genetics, and communication. They might also differ in seasonal reproduction and body weight changes. We investigated and compared reproductive activities and changes in monthly body weight in males and females of successfully breeding colonies of both species under the same photoperiodic conditions. Females of both species showed estrous cycles only during the long-day period. Rufous mouse lemur females seemed to have a shorter gestation than their sibling species (57 vs. 62 days). The number of estrous cycles (2.25 vs. 2.5/season) and their lengths (59 vs. 52 days) were similar. Litter size (2) seemed to be similar. Latency of estrous occurrence after photoperiodic stimulation was longer in Microcebus rufus than in its sibling species (71.6 and 42.3 days). The same was true for the onset of the growth of the testes. The rate of growth and size of the testes were similar, and precede the estrous onset in both species. The reproductive activity was shorter in both sexes of the rufous than of the gray mouse lemurs. In both species, body weight showed similar seasonal changes. Males lost more weight during the breeding season than females did. In rufous mouse lemurs, body weight was similar in both sexes during the nonbreeding season. In gray mouse lemurs, sexes differed throughout the year.     

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.     

Food Intake and Dietary Overlap in Native Lemur catta and Propithecus verreauxi and Introduced Eulemur fulvus at Berenty,Southern Madagascar

The introduction of Eulemur fulvus in 1975 into the Berenty Reserve and their recent attainment of population densities comparable to those of Lemur catta led us to analyze food partitioning among the 3 large prosimian species in the gallery forest. We assessed the diets of ring-tailed lemurs (Lemur catta), brown lemurs (Eulemur fulvus) and sifakas (Propithecus verreauxi) from food intake measurements during 3 successive short-term studies. All species exhibited marked seasonal changes in their major food categories. Dietary overlap was very high between ring-tailed lemurs and brown lemurs during 2 of 3 seasons, including the middle of the dry season. During the latter period, Eulemur appeared to compensate for a low quality diet by increasing the amount of food eaten. In contrast, Lemur fed on lower amounts of food and seemed more efficient at coping with fibrous plant materials. There is low dietary overlap of Lemur catta and Eulemur fulvus versus Propithecus, which exhibit by far the highest dietary diversity of the 3 species. We discuss sustainable coexistence among them, based on respective dietary adaptations and potential for dietary flexibility.     

Coordination of Group Movements in Wild Red-fronted Lemurs (<Emphasis Type="Italic">Eulemur rufifrons</Emphasis>): Processes and Influence of Ecological and Reproductive Seasonality

Group-living species have to coordinate collective actions to maintain cohesion. In primates, spatial movements represent a meaningful model to study group coordination processes across different socio-ecological contexts. We studied 4 groups of red-fronted lemurs (Eulemur rufifrons) in Kirindy Forest, Madagascar, between 2008 and 2010 across different ecological and reproductive seasons. We collected data on ranging patterns using GPS collars and observational data on different predefined parameters of group movements, including initiation, leadership, followership, overtaking events, termination, and travel distances. Cohesion of these relatively small, egalitarian lemur groups was high year-round, but daily path length and home range size varied considerably between ecological seasons, presumably due to long-distance migrations of some groups at the beginning of the rainy season. Individuals of different age and sex classes successfully initiated group movements. However, stable female leadership prevailed year-round, irrespective of ecological and reproductive season, which might be due to higher or more specific energetic requirements of reproduction. In contrast to lemur species with a more despotic social structure, female red-fronted lemurs did not recruit more followers than males. Adult leaders recruited more followers than subadult ones. Further, recruitment success was higher during the peak of the dry season, when predation risk appeared to be higher. Distances of single group movements did not depend on the initiator’s sex and age or on ecological seasons. Our results provide new insights into seasonal variability of coordination processes and the role of social dominance in lemur group movements, thereby contributing to a comparative perspective from a primate radiation that evolved group living independently of anthropoids.     

Use of Vocal Fingerprinting for Specific Discrimination of Gray (Microcebus murinus) and Rufous Mouse Lemurs (Microcebus rufus)

Advertisement calls are often important noninvasive tools for discriminating cryptic species and for assessing specific diversity and speciation patterns in nature. We investigated the contribution of these calls to uncover specific diversity in nocturnal Malagasy lemurs. We compared sexual advertisement and predator advertisement calls of two mouse lemur species, western gray and eastern rufous mouse lemurs (Microcebus murinus and M. rufus, respectively) living in two contrasting habitats (dry deciduous vs. rain forest), and analyzed them statistically. Both species emitted several highly variable whistle calls in the context of predator-avoidance. Intrapopulation variation was high and overlapped interspecific variation. Sexual advertisement calls, given in the mating context, displayed a totally distinct, species-specific acoustic structure. Whereas gray mouse lemurs produced rapidly multifrequency modulated, long trill calls, rufous mouse lemurs gave slowly frequency-modulated short chirp calls. Our results suggest specific status for gray and rufous mouse lemurs and indicate the importance of predation and social needs in shaping vocal communication.     

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.     

Effects of Habitat Quality and Seasonality on Ranging Patterns of Collared Brown Lemur (Eulemur collaris) in Littoral Forest Fragments

Degraded forest habitats typically show low fruit availability and scattered fruit tree distribution. This has been shown to force frugivorous primates either to move further in search of food, resulting in large home ranges, or to use energy saving strategies. Malagasy lemurs are known to face pronounced seasonality and resource unpredictability, which is amplified by the overall reduction in food availability due to the human-driven habitat disturbance on the island. To explore lemur flexibility to habitat disturbance, we examined the ranging behavior of collared brown lemurs (Eulemur collaris) in two differently degraded fragments of littoral forest of southeastern Madagascar. We collected data from February 2011 to January 2012 on two groups living in a degraded area and two groups living in a less disturbed forest. We calculated annual ranges, monthly ranges, and daily distance traveled. We then ran repeated measures ANOVAs using seasonality as dichotomous, intrasubject factor and site/group as intersubject nested factors. In the degraded forest, the lemurs had larger monthly ranges, and their annual ranges were either fragmented or characterized by multiple core areas. They were able to use a habitat mosaic that also included nonforested areas and swamps. In addition, they shortened their daily path length, possibly to preserve energy, and used different areas of their annual home ranges seasonally. Although a number of possible confounding factors may have been responsible for the observed differences between sites, our findings highlight the ranging flexibility of collared brown lemurs in littoral forest fragments.     

Social structure and seasonal variation in the behaviour of Eulemur mongoz

Data presented here were collected during a 10-month study carried out on the mongoose lemur, Eulemur mongoz, at Anjamena in northwestern Madagascar. Baseline information is provided on behaviour and assessed for seasonal variation. Given the highly seasonal nature of the environment, remarkably little behavioural variation was observed. This may be directly linked to the cathemeral activity pattern, where differential movement into either the diurnal or the nocturnal 'niche' could reduce the effect of selective seasonal pressures, thus reducing behavioural and ecological adaptation to seasonal variation. The behavioural mechanisms of reinforcement of the pair bond through close proximity, mutual anogenital marking, paternal participation in the care of offspring and territoriality are all in accord with the general characteristics of monogamy in primates. The feature of mate monopolization by males, proposed as typical of monogamous social systems, seems to be more attributable to females in E. mongoz, as only they exhibit high levels of intrasexual aggression. Female dominance, characteristic of most species of lemur, may thus include more than just female feeding priority in E. mongoz.     

Habitat Degradation and Seasonality Affect Physiological Stress Levels of Eulemur collaris in Littoral Forest Fragments

The littoral forest on sandy soil is among the most threatened habitats in Madagascar and, as such, it represents a hot-spot within a conservation hot-spot. Assessing the health of the resident lemur fauna is not only critical for the long-term viability of these populations, but also necessary for the future re-habilitation of this unique habitat. Since the Endangered collared brown lemur, Eulemur collaris, is the largest seed disperser of the Malagasy south-eastern littoral forest its survival in this habitat is crucial. In this study we compared fecal glucocorticoid metabolite (fGCM) levels, a measure of physiological stress and potential early indicator of population health, between groups of collared brown lemurs living in a degraded forest fragment and groups occurring in a more preserved area. For this, we analysed 279 fecal samples collected year-round from 4 groups of collared brown lemurs using a validated 11-oxoetiocholanolone enzyme immunoassay and tested if fGCM levels were influenced by reproductive stages, phenological seasons, sex, and habitat degradation. The lemurs living in the degraded forest had significantly higher fGCM levels than those living in the more preserved area. In particular, the highest fGCM levels were found during the mating season in all animals and in females during gestation in the degraded forest. Since mating and gestation are both occurring during the lean season in the littoral forest, these results likely reflect a combination of ecological and reproductive pressures. Our findings provide a clear indication that habitat degradation has additive effects to the challenges found in the natural habitat. Since increased stress hormone output may have long-term negative effects on population health and reproduction, our data emphasize the need for and may add to the development of effective conservation plans for the species.     

Mother-Young Feeding Synchrony and Early Food Selection Differences in Eulemur fulvus

I tested the hypothesis that mother-young cofeeding correlates with a systematic similar food selection in wild Mayotte brown lemurs (Eulemur fulvus). I simultaneously recorded the feeding behavior of 4 maternal dyad members, from infant birth to weaning, and 10–12 mo of juvenile age, during 10-min focal periods, each separated by 5-min intervals over 10 mo. I recorded the solid food items selected for each feeding behavior at the level of specific plant part, along with the distance between the 2 individuals. I considered feeding behaviors simultaneous (between the young lemurs and their mothers) when the young began to eat a food item while the mother was already eating, regardless of the separation distance. During their first year, most of the feeding events of the young brown lemurs were initiated by their mothers. However, infants also selected different food items from those chosen by their mothers in almost one-third of cases, and the food selection dissimilarity was not attributable to an increase in the distance of separation. For juveniles, dissimilar food selection was significantly lower and linked to an increase in mother-juvenile separation distance. Thus, the substantial proportion of dissimilarity in solid foods selected by the infant during the synchronized feeding behaviors strongly suggests that social influences were not food-specific and might instead drive the acquisition of group feeding activity rhythm rather than food selection habits.     

Ecological Flexibility as Measured by the Use of Pioneer and Exotic Plants by Two Lemurids: <Emphasis Type="BoldItalic">Eulemur collaris</Emphasis> and <Emphasis Type="BoldItalic">Hapalemur meridionalis</Emphasis>

Primate responses to habitat alteration vary depending on the species’ dietary guild and forest type. Leaves from secondary vegetation can provide nutritious resources to folivorous primates, whereas frugivores, burdened with a scattered spatial and temporal distribution of fruiting resources, require larger home ranges, potentially limiting their ability to cope with altered landscapes. Within coastal southeastern Madagascar, we sought to determine whether two lemur species occupying contrasting ecological niches respond differently to the changing features of their degraded and fragmented habitat. We conducted behavioral observations between 2011 and 2013 on frugivorous collared brown lemurs (Eulemur collaris) and folivorous southern bamboo lemurs (Hapalemur meridionalis). To estimate the ability of lemurs to use pioneer species, we categorized all plants used for feeding and resting as fast growing, mid-growing, or slow growing. We fitted general linear mixed-effects models, one for each plant growth category with monthly proportional use rates as the dependent variable, and included species (E. collaris and H. meridionalis), activity (feeding and resting), and season (dry and wet) as fixed effects. Our results show that E. collaris used both slow- and mid-growing plant species most often, while H. meridionalis were more likely to use fast-growing plants, which indicated an ability to use secondary/disturbed vegetation. Frugivorous E. collaris appear more limited by climax plants, while folivorous H. meridionalis appear to be slightly more adaptable, a finding that is consistent with that for other primate folivores.     

Measuring social tolerance: An experimental approach in two lemurid primates

Social tolerance crucially affects the life of group‐living animals as it can influence, among other things, their competitive regimes, access to food, learning behavior, and recruitment. However, social tolerance tests were mainly conducted in semi‐free or captive populations, and we know little about the behavioral mechanisms and consequences of social tolerance under natural conditions. We therefore developed a co‐feeding experiment to measure social tolerance in groups of wild and captive animals across two primate species. Specifically, we recorded the social tolerance level of redfronted lemurs (Eulemur rufifrons, four wild, one captive group) and ringtailed lemurs (Lemur catta, three wild, three captive groups) by presenting a clumped food resource in an experimental arena, and recorded patterns of resource use during the experiment. Because redfronted lemurs exhibit lower levels of decided conflicts than ringtailed lemurs, we predicted that they would be socially more tolerant. The probability for an individual to feed in the arena was higher in redfronted lemurs than in ringtailed lemurs. In addition, in both species, the probability for an individual to feed in the arena was higher in the captive populations than in their wild counterparts, suggesting that proximate factors, such as a relaxation of feeding competition in captivity, may adapt species‐specific levels of social tolerance to local levels of food availability. Hence, the number of individuals co‐feeding on a valuable food resource appears to be a useful proxy of social tolerance that could be measured with this experimental setup in other wild and captive species as well.     

Niche separation in Varecia variegata rubra and Eulemur fulvus albifrons: II. Intraspecific patterns

Based on a year-long field study in northeastern Madagascar, I summarize annual patterns of niche use (food patch size, diet, forest height, and forest site) in two sympatric lemurs, Varecia variegata rubra and Eulemur fulvus albifrons. Furthermore, I examine intraspecific patterns of niche use according to sex, season, and reproductive stage in these two lemurs that differ in terms of energetic investment in reproduction. Lemurs as a group provide a special opportunity to test hypotheses concerning sex differences in niche use. Due to their body size monomorphism and seasonal, synchronous pattern of breeding, it is possible to directly evaluate whether sex differences in diet reflect high energetic investment in reproduction by females. Results confirm the hypothesis that intraspecific variation in niche use (e.g., sex differences, seasonal differences) would be more pronounced in V. v. rubra than in E. f. albifrons, due in large measure to the former's relatively high energetic investment in reproduction: 1a) Dietary sex differences in V. v. rubra are most pronounced during costly reproductive stages and involve acquisition of low-fiber, high-protein plant foods. Females of both species consume more seasonally available low-fiber protein (young leaves, flowers) relative to conspecific males during the hot dry season, but only in V. v. rubra females is this pattern also evident during gestation and lactation. 1b) The diets of female V. v. rubra and female E. f. albifrons are more similar to each other than are the diets of conspecific males and females in the case of V. v. rubra. This is not uniformly the case for female E. f. albifrons. This finding confirms a hypothesis put forward in Vasey ([2000] Am J Phys Anthropol 112:411-431) that energetic requirements of reproductive females drive niche separation more than do the energetic requirements of males. 1c) Both species synchronize most or all of lactation with seasonal food abundance and diversity. E. f. albifrons shows a more protracted period of synchrony, and this may contribute to its wide biogeographic distribution in Madagascar. 2) Sex differences and seasonal differences in microhabitat use reflect intraspecific patterns of thermoregulation, predator avoidance, and in the case of V. v. rubra, reproduction. One important factor selecting for body size monomorphism in lemurs appears to be the tight synchrony between lactation and periods of food abundance afforded by annual, seasonal breeding.