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.     

Feeding behavior and nutrient intake in spiny forest-dwelling ring-tailed lemurs (Lemur catta) during early gestation and early to mid-lactation periods: compensating in a harsh environment

Strong resource seasonality in Madagascar has led to the evolution of female feeding priority and weaning synchrony in most lemur species. For these taxa, pregnancy/early lactation periods coincide with low food availability, and weaning of infants is timed with increased resources at the onset of the rainy season. Reproductive females experience high metabolic requirements, which they must accommodate, particularly when food resources are scarce. Female ring-tailed lemurs (Lemur catta) residing in spiny forest habitat must deal with resource scarcity, high temperatures (～36-40°C) and little shade in early to mid-lactation periods. Considered "income breeders," these females must use resources obtained from the environment instead of relying on fat stores; thus, we expected they would differ from same-sized males in time spent on feeding and in the intake of food and nutrients. We investigated these variables in two groups (N = 11 and 12) of Lemur catta residing in spiny forest habitat during early gestation and early to mid-lactation periods. Focal animal data and food plant samples were collected, and plants were analyzed for protein, kcal, and fiber. We found no sex differences for any feeding or nutrient intake variable for the top five food species consumed. Females in early gestation spent more time feeding compared with early/mid-lactation. Physiological compensation for spiny forest-dwelling females may be tied to greater time spent resting compared with gallery forest conspecifics, consuming foods high in protein, calories, and water, reduced home range defense in a sparsely populated habitat, and for Lemur catta females in general, production of relatively dilute milk compared with many strepsirrhines.     

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.     

The lack of female dominance in golden‐brown mouse lemurs suggests alternative routes in lemur social evolution

Female dominance is a well‐known trait of lemurs, although it has not been reported from all species and is still often unexplored, especially in the nocturnal species. We examined the intersexual dominance relationships in Microcebus ravelobensis, a congener of M. murinus who is well known for its female dominance. Given the many similarities in biology, it was predicted that M. ravelobensis should also possess female dominance. Seventeen unfamiliar male–female pairs were formed with animals captured in northwestern Madagascar and kept in a two‐cage setting (one cage for each animal) for up to 1 week. Four encounter experiments were conducted with each pair. In contrast to the expectations, females were not consistently dominant over their male partners. Only 3 of 17 dyads developed a clear agonistic asymmetry, among which were two cases of male dominance and only one case of female dominance. Because body mass differences did not explain the findings, various other possible explanations are discussed. It is suggested that food may not be the driving factor of female dominance in mouse lemurs. Instead, it is hypothesized that species‐specific differences in the quality of sleeping sites (i.e., tree holes) and in social grouping patterns may better explain why some mouse lemur species have female dominance, whereas others like the golden‐brown mouse lemur do not. It is concluded thatthese arguments and hypotheses may even hold true for other solitary foragers and may thereby lead to a better understanding of the variable social evolution in lemurs and primates in general. Am J Phys Anthropol, 2013. © 2012 Wiley Periodicals, Inc.     

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 Reproductive Status on Energy Intake,Ingestion Rates,and Dietary Composition of Female <Emphasis Type="BoldItalic">Cebus capucinus</Emphasis> at Santa Rosa,Costa Rica

The limiting factor for fitness in female primates is the acquisition of high-quality food, i.e., food that is high in energy and nutrients, such as protein, fat, and carbohydrates. Reproductive status can influence female feeding patterns, e.g., lactating females in some primate species consume greater quantities of food and specific nutrients than do nonlactating females. We examined the energy intake, ingestion rate, and composition of the diet in female white-faced capuchins in 3 reproductive states: lactating, gestating, and cycling. We observed 12 reproductively active females for 7 mo and compared their energy intake, ingestion rates, and intake rates of nutrients: protein, fat, sugar, and fiber. Lactating females took in significantly more energy per hour while feeding than pregnant and cycling females did. Lactating females also ingested significantly more food items per hour, but they did not spend more time eating than other females did, and they did not differ in the composition of their diet as measured by insect consumption and proportion of protein. Pregnant and cycling females did not differ from each other in any of the measures. We predicted that as the infants aged and began to move independently to forage and play, their mothers’ energy intake rates would decrease in concert with decreased energy demands by the infant. We found a statistically significant relationship between these 2 factors, with infant age explaining 75.4% of variation in mothers’ energy intake, supporting previous studies that found lactation to be the most energetically expensive reproductive state.     

Ecological correlates to activity and habitat use of two prosimian primates: Eulemur rubriventer and Eulemur fulvus rufus in Madagascar

In this paper, I provide data on the possible effects of group size and seasonal changes in food availability on the activity and habitat use patterns of two species of prosimian primate: the rufous lemur (Eulemur fulvus rufus) and the red-bellied lemur (Eulemur rubriventer). General and subtle seasonal differences were observed between lemur species in (1) group size and composition, (2) activity profiles, and (3) habitat use. Rufous lemur groups were larger (mean = 8 individuals) and contained more adults than red-bellied lemur groups (mean = 3 individuals). The overall degree and distribution of diurnal activity differed between lemur species and varied within species with seasonal changes in food availability. In general, rufous lemurs traveled more often than red-bellied lemurs. During food scarcity, both species increased the amount of time spent feeding, although peak feeding and traveling times differed. Both species also preferred horizontal substrates; however, during food scarcity, red-bellied lemurs used terminal branches more often while feeding than rufous lemurs. In addition, red-bellied lemurs used the lower middle story forest and rested in taller, upper canopy trees more often than rufous lemurs. Differences in activity and habitat use patterns between lemur species were more related to seasonal changes in food availability than overall differences in group size. These behavioral patterns may represent different strategies used by each lemur species to avoid directly competing for similar resources during times of food scarcity. © 1996 Wiley-Liss, Inc.     

Lactation and care for unrelated infants in forest-living ringtailed Lemurs

An adult female ringtailed lemur (Lemur catta) known not to have been pregnant showed spontaneous lactation in response to twin infants born to an unrelated female. The females had met only 7 months earlier, when they and two other unrelated adult females were released from separate locations in a forest enclosure to form a new social group. Three months after release, an adult male from an adjacent enclosure gained access to the new group for 1 day, the day of one female's estrus. No males had access to the females throughout the remainder of the breeding season. Within 2 weeks of the birth of the twins, one of the other adult females began carrying the infants frequently, typically one at a time. All three females were checked for lactation when the infants were two months old. Both their mother and the unrelated adult who had been carrying the infants were producing milk. The third adult female, who never carried either infant, had no milk. The third adult female, who never carried either infant, had no milk. This female, however, like the two maternal females, frequently attacked unfamiliar immigrating adult males when the males approached the infants. Potential implications of these observations concerning the social organization of ringtailed lemurs are discussed.     

Ontogeny of Feeding Behavior of Eulemur fulvus in the Dry Forest of Mayotte

Captive data show that juvenile mammals usually reproduce the diet of the adult. However, it is not known whether, in natural environments, the process of feeding maturation varies according to available foods and to adult choices. I examined the feeding ontogeny of Mayotte brown lemurs (Eulemur fulvus), a presumed introduced species, from birth to 12 mo old in the southern dry forest of Mayotte. The study focused on activities and diets of 4 mothers and their juveniles from 2 groups in 1999 and 2000. In 2001, we also observed 2 other dyads from each group for 2 mo (March/April). I assessed plant phenology in each group's home range. There are 4 phases of feeding ontogeny: (1) the 3 mo following birth are characterized by lactation, early bouts of exploration, and consumption; (2) the next 2 mo are characterized by feeding and social weaning; (3) during the sixth and seventh mo, the daily feeding activity rhythms of the mother and her juvenile are synchronized and diets largely overlap; (4) at 10–12 mo of age, the feeding juvenile is closer to other group members than to its mother. During their first year of life, especially before being weaned, juveniles eat several food items that are not ingested by their mothers. It appeared that the progressive overlap in dietary items and feeding synchrony between mother and juvenile depends on several types of leaming. There is a correlation between the availability of particular food resources with high nutritional quality and timing of reproduction and infant maturation.     

Plasma cortisol responses to stress in lactating and nonlactating female rhesus macaques

Lactating female rats without their pups exhibit lower HPA responsiveness to stress than nonlactating females. However, responsiveness to stress is similar when lactating females are tested with their pups and the stressor involves a potential threat to the offspring. This study constitutes the first comparison of stress responsiveness in lactating and nonlactating female nonhuman primates. Subjects were 53 multiparous female free-ranging rhesus macaques. Approximately half of the females were lactating and half were nonpregnant/nonlactating. Blood samples were obtained after capture and after overnight housing in an individual cage. Lactating females were tested with their infants. Lactating females had significantly higher plasma cortisol levels than nonlactating females on both days. Having or not having an infant was also a better predictor of plasma cortisol levels among all females than their age, dominance rank, group of origin, time of day at which the sample was obtained, and time elapsed since beginning of the sampling procedure or since anesthesia. Plasma cortisol levels of lactating females were not significantly correlated with post-partum stage or with the cortisol levels of their infants. Capture, handling, and individual housing in a cage are powerful psychological stressors for free-ranging primates. We suggest that the higher plasma cortisol levels exhibited by lactating females reflect greater responsiveness to stress associated with perception of risks to infants. Hyporesponsiveness to stress may not be a general characteristic of lactation in all mammalian species, but a short-term effect of infant suckling that is most apparent with stressors unrelated to the offspring.     

Seasonal changes in general activity, body mass and reproduction of two small nocturnal primates: a comparison of the golden brown mouse lemur ( Microcebus ravelobensis) in Northwestern Madagascar and the brown mouse lemur ( Microcebus rufus) in Eastern Madagascar

To investigate for the first time the relationship between contrasting patterns of seasonal changes of the environment and activity, body mass and reproduction for small nocturnal primates in nature, we compared a population of golden brown mouse lemur (Microcebus ravelobensis) in a dry deciduous forest of northwestern Madagascar and of the brown mouse lemur (Microcebus rufus) in an evergreen rain forest of eastern Madagascar. Both species live under similar photoperiodic conditions. Golden brown mouse lemurs (GBML) were active during the whole period (May to December) irrespective of changing environmental conditions. In contrast, a part of the population of brown mouse lemurs (BML) showed prolonged seasonal torpor, related to body mass during periods of short day length and low ambient temperatures. Differences between species might be due to differences in ambient temperature and food supply. Body weight and tail thickness (adipose tissue reserve) did not show prominent differences between short and long photoperiods in GBML, whereas both differ significantly in BML, suggesting species-specific differences in the photoperiodically driven control of metabolism. Both species showed a seasonal reproduction. The rate of growth and size of the testes were similar and preceded estrous onset in both species suggesting a photoperiodic control of reproduction in males. The estrous onset in females occurred earlier in GBML than in BML. Estrous females were observed over at least 4 months in the former, but in only 1 month in the latter species. Intraspecific variation of estrous onset in GBML may be explained by body mass. Interspecific variation of female reproduction indicates species-specific differences in the control of reproduction. Thus, environmentally related differences in annual rhythms between closely related small nocturnal lemurs emerged that allow them to cope with contrasting patterns of seasonal changes in their habitats.     

Diets of Two Lemur Species in Different Microhabitats in Beza Mahafaly Special Reserve,Madagascar

Studies of primate diets usually focus on differences that distinguish species or populations. However, variation in diet can occur at a more local level of groups within a population, especially in a non-homogeneous habitat. I compared dietary variation in food composition and toughness across groups of 2 lemur species in Beza Mahafaly special reserve, Madagascar. Beza Mahafaly contains an 80-ha reserve (Parcel 1) that, while small, hosts a dense population of Lemur catta (ring-tailed lemurs) and Propithecus verreauxi verreauxi (sifakas). Microhabitats in the eastern vs. western sides of the parcel are structurally and floristically distinct. Sifakas in this parcel have small, discrete home ranges and are morphological folivores. For these reasons, I expected that the 6 groups studied would eat a different menu of food plants but with similar toughness values. Ring-tailed lemurs have comparatively large, overlapping home ranges, and I expected that the 5 study groups would eat similar foods. Despite living in different microhabitats across the parcel, sifakas exhibit high dietary uniformity both in dietary plant species composition and the toughness of the foods. Food selection in sifakas operates on two distinct levels. Sifaka groups share many key food species that appear independent of local abundances, but the ranking of the foods within each group appears related to availability. Ring-tailed lemur groups are more heterogeneous in the composition of their diets relative to sifakas, though the time spent feeding on individual foods reveals a marked preference for the fruits of Tamarindus indica by all groups. Food toughness is consistent across the parcel with the exception of the most western group. Ring-tailed lemurs are highly specific feeders, but indiscriminate nibblers. Sifakas are targeted, balanced feeders. There does not appear to be a consistent microhabitat effect operating across species. Differences within sifaka and ring-tailed lemur populations in food composition and toughness, however, correspond to an east-west microhabitat gradient. Measures of dietary flexibility must take into account not only the plant species consumed and the different parts eaten but also their associated food properties and proportion of time spent feeding on them.     

Feeding Ecology of Sympatric Mouse Lemur Species in Northwestern Madagascar

We performed a comparative pilot study on vertical space use and feeding ecology of 2 closely related sympatric mouse lemur species in northwestern Madagascar. We recorded feeding behavior and the use of vertical forest strata for 6 gray and 4 golden-brown mouse lemur females that we followed for 120 h. We carried out feeding experiments and analyzed fecal samples for food remains. Both species showed no significant difference in use of forest strata. They had a similar overall diet, with insect secretions and gum as main plant food resources; arthropod remains occurred in about half of all fecal samples. Both lemurs used in common >50% of the plant species that each consumed, which accounts for ≥70% of all individual plants used. However, both species used >40% of their feeding plant species exclusively and seemed to differ in their degree of specialization on certain plant species.     

A comparison of salivary pH in sympatric wild lemurs (Lemur catta and Propithecus verreauxi) at Beza Mahafaly Special Reserve, Madagascar

Chemical deterioration of teeth is common among modern humans, and has been suggested for some extinct primates. Dental erosion caused by acidic foods may also obscure microwear signals of mechanical food properties. Ring-tailed lemurs at the Beza Mahafaly Special Reserve (BMSR), Madagascar, display frequent severe tooth wear and subsequent tooth loss. In contrast, sympatric Verreaux's sifaka display far less tooth wear and infrequent tooth loss, despite both species regularly consuming acidic tamarind fruit. We investigated the potential impact of dietary acidity on tooth wear, collecting data on salivary pH from both species, as well as salivary pH from ring-tailed lemurs at Tsimanampesotse National Park, Madagascar. We also collected salivary pH data from ring-tailed lemurs at the Indianapolis Zoo, none of which had eaten for at least 12 hr before data collection. Mean salivary pH for the BMSR ring-tailed lemurs (8.098, n=41, SD=0.550) was significantly more alkaline than Verreaux's sifaka (7.481, n=26, SD=0.458). The mean salivary pH of BMSR (8.098) and Tsimanampesotse (8.080, n=25, SD=0.746) ring-tailed lemurs did not differ significantly. Salivary pH for the Indianapolis Zoo sample (8.125, n=16, SD=0.289) did not differ significantly from either the BMSR or Tsimanampesotse ring-tailed lemurs, but was significantly more alkaline than the BMSR Verreaux's sifaka sample. Regardless of the time between feeding and collection of pH data (from several minutes to nearly 1 hr), salivary pH for each wild lemur was above the "critical" pH of 5.5, below which enamel demineralization occurs. Thus, the high pH of lemur saliva suggests a strong buffering capacity, indicating the impact of acidic foods on dental wear is short-lived, likely having a limited effect. However, tannins in tamarind fruit may increase friction between teeth, thereby increasing attrition and wear in lemurs. These data also suggest that salivary pH varies between lemur species, corresponding to broad dietary categories.     

Social organization of the Alaotran gentle lemur (Hapalemur griseus alaotrensis)

Knowledge of the social organization of lemurs is still limited for most species. Where there is sufficient information, it has been shown that lemur social organization differs in essential points from that of other primates. In the field study reported here, demographic structure and life-history processes were investigated in order to characterize the social organization of the Alaotran gentle lemur (Hapalemur griseus alaotrensis). Data were obtained through captures and observations. Alaotran gentle lemurs were found in small groups of up to nine individuals. Although most groups contained just one breeding female, a substantial proportion of groups (35%) had two breeding females. Therefore, Alaotran gentle lemurs cannot be classed as being organized in monogamous family groups. An extended birth season was found, and groups with two breeding females had significantly higher breeding output per adult than groups with a single adult female. Limited data suggest that females emigrate from their natal group while still subadult, whereas males can stay in the natal group until they are fully grown and disperse as adults. Variability in group composition, significantly higher reproductive output per adult in groups with two breeding females, and delayed dispersal of males suggest that Alaotran gentle lemurs pursue a resource-defense mating strategy, rather than a female-defense mating strategy. The suggestion that extant social lemurs may have evolved from a monogamous system, could explain the differences between lemur social systems and those of other primates.     

Life history traits and parental care in Lepilemur ruficaudatus

In this study we investigated the importance of biparental care for the evolution and/or maintenance of pair-living in red-tailed sportive lemurs (Lepilemur ruficaudatus), a nocturnal folivorous lemur. Between 2000 and 2005, we collected data on life history traits from a total of 14 radio-collared pairs of adults and their offspring in Kirindy forest, western Madagascar. Predation rate varied between years with a minimum of 0% and a maximum of 40% per year. Patterns of parental care were quantified during simultaneous focal observations of both pair-partners in 2003 and 2004. Mating activity was limited to the months of May and June, as indicated by conspicuous changes of vulval morphology and male mate guarding behavior. After a gestation length of about 5 months, which is much longer than expected for a lemur of this body mass, single infants were born in November. Lactation lasted for about 50 days. Apart from lactation, females provided infant care by warming, grooming and transporting infants orally. Infants were parked in dense vegetation while females foraged. Males were seen only rarely in proximity to infants and we found no evidence for direct infant care provided by social fathers. We conclude that the necessity of direct infant care cannot explain the evolution and/or maintenance of pair-living in Lepilemur ruficaudatus.     

Comparing the accuracy and precision of three techniques used for estimating missing landmarks when reconstructing fossil hominin crania

In some primate species, pelage colorations at birth contrast with adult colorations. The intensity of natal coats and their phylogenetic distribution is highly variable within primates. Natal coat coloration seems to change to adult coloration in most species when infants become independent from their mothers, but an accepted functional explanation for natal coats is not available. Here we describe pelage coloration change in sexually dichromatic redfronted lemurs (Eulemur fulvus rufus) in Kirindy Forest, and propose a new functional hypothesis for this phenomenon. In this species, infants are born with adult male coloration and female infants subsequently undergo a change in coloration. Using digital pictures and behavioral data collected on eight mother-offspring dyads from birth until the end of the coloration change, we 1) described timing and pattern of pelage developmentin redfronted lemur infants and 2) examined behavioral developmental correlates of the coloration change. The color change took place between 7 and 17 weeks of age and coincided with advanced physical independence; a pattern also found in monochromatic primate species with natal coats. No behavioral differences between male and female infants were found. Hypotheses about the ultimate function of natal coats focusing on enhanced infant care or reduced infanticide risk did not explain the pelage change in redfronted lemurs. The natal pelage pattern in this species may instead serve as sexual mimicry. Accordingly, female infants may mimic males during the most vulnerable developmental phase to avoid sex-specific aggression by adult females in a species with intense female-female aggression and competition.     

Behavior of the mayotte lemur,Lemur fulvus mayottensis,in captivity

A group of one male and two female Mayotte lemurs was observed at Chester Zoo, and their behaviors compared with published observations of wild populations of this and related forms of the brown lemur. It was found that the captive animals were active by both day and night, but that the pattern of activity was different from wild lemurs. The captive lemurs were less active than their wild counterparts and showed different timing of activity because of the feeding regimen at the zoo. The animals did not ignore the public at the zoo, but, on the contrary, directed some behaviors at the zoo visitors, particularly if the zoo visitors attempted to interact with the lemurs; under these conditions there was also an increase in the movements of the lemurs in the cage. Levels of aggression in the zoo lemurs were higher than those reported for wild populations, and aggression appeared to occur predominantly in the birth season. Levels of affiliative behaviors, particularly allogrooming, were comparable with those in the wild. Olfactory behaviors such as scent marking and anogenital sniffing also showed seasonal peaks and again appeared to occur at higher rates than in the wild.     

Microhabitat Variation and Its Effects on Dietary Composition and Intragroup Feeding Interactions Between Adult Female <Emphasis Type="Italic">Lemur catta</Emphasis> During the Dry Season at Beza Mahafaly Special Reserve,Southwestern Madagascar

We compared diets and female feeding agonism in 2 groups of ring-tailed lemurs living in markedly diverse microhabitats in and near the Beza Mahafaly Reserve, Madagascar during mid-to-late gestation periods and height of the dry season to test predictions concerning differences in resource availability, preferred foods, female rank, and degree and frequency of feeding agonism in relation to usurpability and monopolization of food resources. Quadrat sampling in disturbed forest habitat revealed a greater number of plant species than in the gallery forest home range area, but females in both groups consumed nearly equal numbers of actual food plant species. Higher-ranking females in disturbed forest consumed human food scraps from the researchers camp significantly more often than low-ranking females did, while there was no rank effect for consumption of any food type between females in the gallery forest group. Higher rates of female feeding agonism in both groups occurred around usurpable fruit and leaves, as well as over monopolizable human food scraps in the disturbed forest group. There is no association between degree of agonism and food type in either group, and rate of feeding agonism is similar for both groups. The most highly contested food items came in large packages and were high in nutrients: beneficial foods for gestating females in the height of dry season in southwestern Madagascar.     

Effects of lactation on the time-budgets and foraging patterns of female black howlers (Alouatta pigra)

Lactation is an energy demanding phase in the reproductive cycle of female mammals. For this reason, several studies have assessed the effects of lactation on female behavior. In this study we examine the influence of lactation on the time-budgets and foraging patterns of female black howlers (Alouatta pigra) in Campeche, Mexico. We observed 32 adult females and 35 infants belonging to 14 groups of black howlers for a total of 2,224 focal hours. We found that lactating females spent more time being inactive and feeding from fruits than nonlactating females. In addition, during the first two-thirds of lactation females were more active (i.e., rested less, fed more, devoted more time to social activities, and moved more) and foraged more intensively (i.e., ranged over larger distances, used more feeding trees and feeding species, and consumed more leaves) than females in the last third of lactation. Lactation seems to force black howler females to reduce activity and to maximize the intake of high-quality foods, with inactivity being the highest during late lactation, when females probably face the cumulative effects of nursing older infants and of a new pregnancy. Early lactation is probably the most energetically demanding stage of lactation for black howler females. This study demonstrates that despite being energetically constrained by a highly folivorous diet, reproductive state affects several dimensions of the behavior of black howler females. Therefore, variation in time-budgets and foraging strategies of howlers has been probably underestimated by previous research that has not considered physiological differences among individuals.