Differences in Activity Budgets and Diet Between Semiprovisioned and Wild‐Feeding Groups of the Endangered Barbary Macaque (Macaca sylvanus) in the Central High Atlas Mountains,Morocco

The Barbary macaque, Macaca sylvanus is a very adaptable primate species occupying a wide range of habitats in Morocco and Algeria. Several groups of this endangered macaque can be found in tourist sites, where they are affected by the presence of visitors providing food to them. We compare the activity budgets and the diet of semiprovisioned and wild‐feeding groups of Barbary macaques in the central High Atlas Mountains of Morocco from February to August 2008. We used instantaneous scan sampling at 15‐min intervals. The behaviors included in the activity budget were feeding, moving, foraging, resting, and aggressive display. Food items were grouped into seven categories. We found no differences between the two groups in the daily percentages of records attributed to feeding. The semiprovisioned group spent significantly more time engaged in resting and aggressive behavior, and foraged and moved significantly less than the wild‐feeding group. There was no significant difference between the two groups in time spent eating leaves, fruits, or roots and bark. The semiprovisioned group, however, spent significantly less time per day feeding on herbs, seeds, and acorns than the wild‐feeding group. Human food accounted for 26% of the daily feeding records for the semiprovisioned group and 1% for the wild‐feeding group. Our findings agree with previous studies and indicate that in the tourist site, where food is highly clumped, macaques decreased foraging time yet showed higher levels of contest competition. Our results support the common claim that the diet of the Barbary macaque is highly flexible, differing among its varied habitats. Conservation efforts for the Barbary macaques should take into account the changes in behavior that human‐modified environments may cause.     

Comparative density,demography, and ranging behavior of Barbary macaques (Macaca sylvanus) in marginal and prime conifer habitats

A semiisolated study population of 162 Barbary macaques (six groups) inhabiting the Ghomaran fir forests of the Moroccan Rif mountains has a density of 6.73 individuals/km ². The adult sex ratio is 0.725, and immatures comprise 46.9% of the population. Births are seasonal, occurring from April to June, and the adult female birth rate is 0.58 per annum. Mortality appears relatively low in all age classes until old age. Group size ranges from 12 to 59 individuals, with a median value of 24. Home-range sizes vary between 3 and 9 km², with a mean of 7.2 km². Home-range overlap is approximately 80%. On the basis of macaque density, conifer density, and herding competition from domestic animals, the Ghomaran environment can be considered “marginal” compared to the Moyen Atlas. Despite the marginal habitat of the Ghomaran population, it is surprisingly similar in demographic characteristics to a Barbary macaque population in the Moyen Atlas. Two characteristics of the population dynamics in the Ghomara differentiate it from the former. (1) The mean home range is five times larger in the Ghomara, roughly inversely proportional to the sixfold decrease in macaque density, confirming Caldecott’s (1986) principle that, in macaque species, range size adjustments are a primary proximate response to poor-quality habitat. (2) Smaller groups in the marginal habitat of the Ghomara appear to have better rates of growth than small groups in prime habitat. This may result from an overall decreased home-range defensibility in marginal habitat (larger home ranges), resulting in an ecological and demographic release of small groups from the levels of intergroup competition they would normally experience in prime habitat.     

Geographic distribution and habitat diversity of the Barbary Macaque sylvanus L.

During a 15-month behavioral study in Morocco and a 3-month survey in Morocco and Algeria, the present distribution of the Barbary macaque was determined. In Algeria, monkeys are found in seven constricted and disjunct localities in the Grande and Petite Kabylie mountain ranges. These localities are severely restricted in space and are located in remote or inaccessible areas which support only small populations. Their habitats include mixed cedar and holm oak forests, humid Portuguese and cork oak mixes and gorges dominated by scrub vegetation. In only two regions (Guerrouch and Agfadou) can population of reasonable size be found; even there they do not approach the abundance found in the central Middle Atlas zone of Morocco. Distribution was more extensive earlier in this century and some areas have become unoccupied within the past 15 years. Today, their absolute numbers and population densities are low in all but two locations. Algeria contains approximately 23% (5,500 maximum) of the total number of surviving Barbary macaques in North Africa. About 77% of the total number of Barbary macaques occur in Morocco. Moroccan habitats include high cedar forests, cedar/holm-oak mixtures and pure holm oak forests. Macaque distribution in the High Atlas is restricted to the Ourika valley where only a small relict population survives. There are between five and eight small, disjunct forest pockets in the Rif which support small groups of monkeys. In the Middle Atlas, monkeys are found in high numbers and in relatively wide stretches of distribution, although there are constricted areas of low densities in this region also. 65% (14,000 maximum) of the animals and their highest densities occur in the high mixed cedar forests of the Central zone, and mixed cedar forest appears to be the preferred habitat for the species. With the exception of the Central zone, their remaining distribution is typically disjunct and constricted, and population densities aer low. As in Algeria, distribution in Morocco was wider earlier this century, and several areas have recently become unoccupied.     

Habitat distribution and habitat preference in Barbary macaques (Macaca sylvanus)

Barbary macaques (Macaca sylvanus,L.) occur at varying densities in a number of different habitat types in Morocco and Algeria. Taub (1977) has argued that the abundance of the species in cedar forests of the Moyen Atlas, Morocco, is an indication of a habitat preference. A reexamination of available data on the distribution and abundance of Barbary macaques suggests that monkey numbers reflect the distribution of habitat size rather than habitat types. Differences between populations relative to habitat types can be seen only between forest and scrub localities. Human factors are considered more important in determining the present status of this eclectic feeder. The modern distribution of the Barbary macaque is inadequate evidence for a habitat preference.     

Effect of habitat quality on the ecological behaviour of a temperate-living primate: time-budget adjustments

Barbary macaques, like other non-human primates living in highly seasonal temperate environments, display high monthly variations in their diet. In addition, their diet changes according to the habitat type they colonize and to the degree of habitat degradation due to resource exploitation by local people, in particular through pastoralism. We studied the time-budget adjustments of wild Barbary macaques in three cedar–oak forests impacted by different intensities of grazing pressure from goats and sheep. We examined how diet variations influenced the time monkeys spent in their activities and their day range lengths (i.e. their energy costs). At three studied sites, diet composition and time budgets showed marked seasonal variations. Diet composition had a strong influence on monkeys’ time budget. In the forest where pastoralism was the highest, diet included a greater proportion of underground resources, shrub fruit and acorns, which led to an increase in the time spent foraging and moving, as well as an important increase in day range lengths. Energy costs were therefore higher in a degraded environment than in a suitable habitat. The monkeys living in forests subjected to pastoralism took advantage of increased day lengths to spend more time searching for food. However, in the forest with the highest pastoralism pressure, although monkeys spent more time foraging, they spent less time feeding than monkeys at the other sites. In addition, they appeared to have reached the limits of the available time they could devote to these activities, as their diurnal resting time was at its lowest level over several months. Temperature variations did not appear to modify monkeys’ time budgets. In the least favourable habitat, saving time from resting activity allowed monkeys to maintain a relatively high level of social activity, partly linked to rearing constraints.     

Possible use of heterospecific food-associated calls of macaques by sika deer for foraging efficiency

Heterospecific communication signals sometimes convey relevant information for animal survival. For example, animals use or eavesdrop on heterospecific alarm calls concerning common predators. Indeed, most observations have been reported regarding anti-predator strategies. Use of heterospecific signals has rarely been observed as part of a foraging strategy. Here, I report empirical evidence, collected using playback experiments, showing that Japanese sika deer, Cevus nippon, use heterospecific food calls of Japanese macaques, Macaca fuscata yakui, for foraging efficiency. The deer and macaques both inhabit the wild forest of Yakushima Island with high population densities and share many food items. Anecdotal observations suggest that deer often wait to browse fruit falls under the tree where a macaque group is foraging. Furthermore, macaques frequently produce food calls during their foraging. If deer effectively obtain fruit from the leftovers of macaques, browsing fruit fall would provide a potential benefit to the deer, and, further, deer are likely to associate macaque food calls with feeding activity. The results showed that playback of macaque food calls under trees gathered significantly more deer than silence control periods. These results suggest that deer can associate macaque food calls with foraging activities and use heterospecific calls for foraging efficiency.     

Is Diet Flexibility an Adaptive Life Trait for Relictual and Peri-Urban Populations of the Endangered Primate Macaca sylvanus?

Habitat loss, fragmentation and urban expansion may drive some species to marginal habitats while others succeed in exploiting urban areas. Species that show dietary flexibility are more able to take advantage of human activities to supplement their diet with anthropogenically abundant and accessible resources. The Barbary macaque (Macaca sylvanus) is an endangered species due to the loss of its habitat, and human pressure. The population of Gouraya National Park (Algeria) lives in a relictual habitat that constitutes about 0.6% of the species range. In addition, this population is a unique case where urban expansion favours contact zones between Barbary macaque habitats and a big city (Bejaia). We quantified the dietary composition of Gouraya macaques over an annual cycle with the objective to understand how diet flexibility of this species may help it adapt to a relictual habitat or cope with urban expansion. We recorded the phenology of plant species every month. This study shows that Gouraya macaques, compared to those living in other forest types of the distribution area, are under lower seasonal constraints. They consume a greater amount of fruit and seeds that are available throughout much of the year, and a lesser amount of costly to find and extract subterranean foods. Therefore the Gouraya relictual habitat appears as a favourable environment compared to other major habitats of that species. This study also shows that colonizing peri-urban zones increases the availability and species richness of diet resources for Barbary macaques as they consume more human foods and exotic plants than in farther sites. Adult males eat more human foods than adult females and immatures do. The exploitation of high-energy anthropogenic food could favour macaque population growth and expansion towards the city center associated with human/macaque conflicts. We recommend applying management actions to restore macaques back to their natural habitat.     

The Effect of Climatic Factors on the Activity Budgets of Barbary Macaques (Macaca sylvanus)

Climatic conditions can significantly affect the behavior of animals and constrain their activity or geographic distribution. Barbary macaques (Macaca sylvanus) are one of the few primates that live outside the tropics. Here we analyze if and how the activity budgets of Barbary macaques are affected by climatic variables, i.e., air temperature, relative humidity, rainfall, and snow coverage. We collected scan sampling data on the activity budgets of four groups of macaques living in the Middle Atlas Mountains of Morocco from June 2008 to January 2011. This habitat is characterized by extreme seasonal changes, from cold and snowy winters to hot and dry summers. The activity budgets of the macaques differed across months but not across the time of day (with the exception of time spent feeding). The monkeys spent significantly more time feeding or foraging when there was no snow than when snow coverage was moderate or major. Daily rainfall was positively related to resting time and negatively to time spent moving or in social behavior. Air temperature was negatively related to time spent feeding or foraging. Finally, time spent on social behavior was significantly lower when relative humidity was high. These data indicate that environmental factors significantly affect the time budgets of endangered Barbary macaques, a species that has been little studied in the wild. Our findings support previous studies on temperate primates in showing that snow coverage can have negative consequences on the feeding ecology and survival of these species.     

Influence of Fruit Availability on Fruit Consumption in a Generalist Primate,the Rhesus Macaque <Emphasis Type="Italic">Macaca mulatta</Emphasis>

Fluctuations in resource availability occur in all ecosystems. To survive, species must alter their foraging strategies according to the quantity, quality, and distribution of available food. The rhesus macaque (Macaca mulatta), a commensal primate, is considered a generalist omnivore and very few studies have addressed how its feeding strategies change with respect to resource availability. We examined dietary diversity and frugivory levels in a group of rhesus macaques at the Buxa Tiger Reserve in northern India across one year. Using behavioural observations of diet and phenological monitoring, we found that although rhesus macaques fed on 107 food items including leaves, flowers, fruits, seeds, and insects, fruits made up ca. 74% of their diet. Fruit consumption correlated positively with fruit availability, but fruit preference appeared to play an important role; 16% of all the fruit species they fed on accounted for >50% of all fruit feeding observations. We suggest that afforestation programs involving preferred fruit species at the agricultural land–forest interface would prevent forest groups of rhesus macaques from gravitating toward human habitations and reduce conflict over anthropogenic resources. We further propose that the movement of certain primates in the direction of human habitations may be contingent on resource availability and food preference rather than an inherent propensity to gravitate to anthropogenic areas.     

Current Status of Macaca mulatta in Taihangshan Mountains Area, Jiyuan, Henan, China

Primatologists have classified rhesus macaques of the Taihangshan Mountains area as Macaca mulatta tcheliensis, a subspecies confined to China. The northernmost area of its distribution is in the southern part of the Taihangshan Mountains, which includes a National Nature Reserve established in 1998. To investigate the current status of the subspecies, we surveyed the population and distribution of macaque groups in Jiyuan, Henan, China. In addition, we investigated habitat preferences and the food items foraged by macaques during winter and early spring. The results show that: 1) ca. 2100 macaques currently inhabit the Taihangshan Mountains National Nature Reserve, mainly in Jiyuan; 2) fragmentation of habitats of rhesus macaques have become severe in recent years; and 3) food resources are very poor during the long winter and the early spring. Macaques in the Taihangshan Mountains feed mainly on seeds, twigs, and bark and the eggs of some insects.     

Behavioral responses of Barbary macaques (Macaca sylvanus) to variations in environmental conditions in Algeria

In this study, the behavioral responses of Barbary macaques to seasonal and interhabitat variations in resource availability were analyzed over an entire annual cycle. Two groups, one in an evergreen cedar–oak forest (Djurdjura) and the other in a deciduous oak forest (Akfadou), were observed. In this paper, references to data on resource availability published elsewhere are made. Time budget has been studied. Variations in foraging and moving time, in day–range lengths, and in time moving in trees have been considered to estimate the variations in foraging effort and thus energy expenditure. Great monthly variations in foraging effort and other activities were observed in both habitats. In early spring, when resource availabilities were maximal, foraging effort was low while monkeys maximized their feeding time (about 5 h/day). In June, during the peak of the birth season and the rearing period, monkeys minimized their feeding time to the benefit of social interactions (to 1.6–2.7 h/day), whatever the food availability, which was low in Akfadou and high in Djurdjura. In addition, foraging effort remained low in Djurdjura, while it increased in Akfadou. Thus, at the beginning of the dry summer period, monkeys in Akfadou were in a less favorable position than those in Djurdjura. At both sites, in periods of food shortage in summer or in winter, monkeys displayed two different strategies. In the former case, their foraging effort increased, while in the second one it remained relatively low. Whatever the foraging effort, monkeys did not reach the same amount of feeding time as in early spring. In the poorest site of Akfadou, foraging effort was globally greater than in the richest site of Djurdjura, especially for adults. At both sites, adult males spent more time feeding than juveniles and less time in social interactions. Results are discussed according to rearing period, temperatures, and day length constraints. The limits of adaptability to different habitats are considered in light of the demographic parameters. Am. J. Primatol. 43:285–304, 1997. © 1997 Wiley-Liss, Inc.     

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.     

Effects of Food Type and Number of Feeding Sites in a Tree on Aggression During Feeding in Wild <Emphasis Type="Italic">Macaca fuscata</Emphasis>

It is important to understand the effects of ecological factors on aggression during feeding in order to link habitat characteristics to competitive regime and social relationships. Multiple habitat characteristics are likely to affect aggression, but few researchers have examined the effect of multiple factors on intragroup competition simultaneously. I examined the effect of 8 factors on aggression during feeding in wild Japanese macaques living in a coniferous forest in Yakushima: density of the tree species, feeding time, number of feeding sites within a feeding tree, number of cofeeding animals, intratree macaque density, food type, and rank and age of the focal individual. When macaques cofed with other individuals, food type, the number of feeding sites, and their interactions significantly influenced aggression. Aggression increased when macaques ate fruits/seeds versus other foods and as the number of feeding sites decreased. Primate socioecological models highlight the importance of clumped distribution of food patches as a correlate of intragroup contest. However, my study indicates that primatologists need to pay attention to the factors related to the current feeding tree—food type and feeding tree size with respect to monopolizability—in addition to the distribution of food in the entire home range.     

Niche separation of sympatric macaques, Macaca assamensis and M. mulatta, in limestone habitats of Nonggang, China

Comparative studies of sympatric species are essential in understanding those species’ behavioral and ecological adaptations as well as the mechanisms that can reduce resource competition enough to allow coexistence. We collected data on diet, activity budget and habitat use from two sympatric macaque species, the Assamese macaque (Macaca assamensis) and the rhesus macaque (M. mulatta), in a limestone seasonal rainforest of Nonggang Nature Reserve, southwestern Guangxi, China. Our results show that the two sympatric macaques differ in diet, activity budget, and habitat use: (1) out of the 131 plant species that were used by both macaque species as food over the year, only 15 plant species (11 %) were shared. Rhesus macaques used more plant species as major foods, and had higher dietary diversity and evenness indexes than Assamese macaques. (2) Assamese macaques fed predominantly on leaves, whereas rhesus macaques fed more selectively on fruits. The rhesus macaques’ diet varied according to season, and was significantly correlated to season fluctuation in fruit availability. (3) Assamese macaques devoted more time to resting, and less time to feeding than rhesus macaques (4) Assamese macaques were present mostly on the cliff, and tended to stay on the ground, whereas rhesus macaques were present mostly on the hillside, and showed preference to lower and middle canopy. The observed differences in diet and habitat use between the two macaque species represent behavioral patterns enabling their coexistence.     

Spatial patterns in the diet of the Japanese macaque Macaca fuscata and their environmental determinants

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野生藏酋猴冬季食物组成及营养成分

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Diet and feeding behavior of Rhinopithecus bieti at Xiaochangdu, Tibet: adaptations to a marginal environment

The diet and feeding ecology of a wild subpopulation of black-and-white snub-nosed monkeys (Rhinopithecus bieti) were studied at Xiaochangdu in Honglaxueshan Nature Reserve, Tibet. This region is climatologically harsher than any other inhabited by non-human primates. Black-and-white snub-nosed monkeys fed on 48 parts of 25 plant species, at least three species of lichens and seven species of invertebrates. The number of food items exploited varied markedly among seasons, with dietary diversity being greatest in spring and summer. In winter, black-and-white snub-nosed monkeys had to subsist on fallback foods such as dried grass and bark. Ubiquitous lichens formed a major dietary constituent throughout the year, contributing about 75% of feeding records. Even though lichens act as a staple, our findings signify that the monkeys at Xiaochangdu prefer feeding on foliage, which is higher in protein content than the former. We provide evidence that black-and-white snub-nosed monkeys are able to cope with an array of food items other than lichens and hence can be regarded as feeding generalists. We discuss the results with reference to previous studies on other subpopulations living in habitats that are floristically more diverse and offer more plant food items than the marginal habitat at Xiaochangdu.     

Linking summer foraging to winter survival in yellow pine chipmunks (Tamias amoenus)

This study links summer foraging and scatter-hoarding to winter larder-hoarding and winter survival in yellow pine chipmunks (Tamias amoenus) by comparing patterns of time allocation and winter larder contents in 2 years with very different levels of resource availability. In 2003, seed production and the number of trees and shrubs producing seeds were high. In 2004 seed crops were small. Chipmunks allocated more time to foraging when food resources were scarce (66% in 2004) compared to when they were abundant (39% in 2003). Increased time allocated to foraging in 2004 corresponded to significant decreases in time allocated to vigilance, resting, and social interactions. When seeds were scarce (i.e., in autumn 2004), chipmunks spent more time searching for cached food items than gathering seeds from plants or the ground surface. Despite the increase in foraging effort, the edible mass and caloric contents of larders were significantly smaller in 2004. In the year with low seed production, the diversity of seed species found in larders increased, and many of these seeds were of species that ripened in summer. When autumnal seed production by Jeffrey pine seeds was high, Jeffrey pine seeds were nearly the exclusive food item found in larders. Larder contents would have provisioned chipmunks for an estimated 116-257 days in 2003 and but only 6-111 days in 2004. It is likely that all chipmunks would have survived the winter of 2003 (duration ~110-120 days). However, none of the larders recovered in 2004 contained enough food to have provisioned the inhabitant for the ~148-158 days of winter.     

Northern pigtailed macaques rely on old growth plantations to offset low fruit availability in a degraded forest fragment

Space-use and foraging strategies are important facets to consider in regard to the ecology and conservation of primates. For this study, we documented movement, ranging, and foraging patterns of northern pigtailed macaques (Macaca leonina) for 14 months in a degraded habitat with old growth Acacia and Eucalyptus plantations at the Sakaerat Biosphere Reserve in northeastern Thailand. We used hidden Markov models and characteristic hull polygons to analyze these patterns in regard to fruit availability. Macaques' home range (HR) was 599 ha and spanned through a natural dry-evergreen forest (DEF), and plantation forest. Our results showed that active foraging increased with higher fruit availability in DEF. Macaques changed to a less continuous behavioral state during periods of lower fruit availability in DEF, repeatedly moving from foraging to transiting behavior, while extending their HR further into plantation forest and surrounding edge areas. Concomitantly, macaques shifted their diet from fleshy to dry fruit such as the introduced Acacia species. Our results showed that the diet and movement ecology adaptations of northern pigtailed macaques were largely dependent on availability of native fruits, and reflected a “high-cost, high-yield” foraging strategy when fresh food was scarce and dry fruit was available in plantation forest. Conversely, wild-feeding northern pigtailed macaque populations inhabiting pristine habitat approached a “low-cost, low-yield” foraging strategy. Our results outline the effects of habitat degradation on foraging strategies and show how a flexible species can cope with its nutritional requirements.     

Molecular phylogeny of macaques: implications of nucleotide sequences from an 896-base pair region of mitochondrial DNA

We determined the nucleotide sequences of an 896-base pair region of mitochondrial DNA (mtDNA) from 20 primates representing 13 species of macaques, a baboon, and a patas. We compared these sequences and the homologous sequences from four macaques and a human against each other and deduced the phylogenetic relationships of macaques. The results from the phylogenetic analyses revealed five groups among the macaques: (1) Barbary macaque, (2) two species of Sulawesi macaques, (3) Japanese, rhesus, Taiwanese, crab-eating, and stump-tailed macaques, (4) toque, pig-tailed, and lion-tailed macaques, and (5) Assamese and bonnet macaques. The phylogenetic position of Tibetan macaque remains ambiguous as to whether it belongs to the fourth or fifth group. Phylogenetic trees revealed that Barbary macaque diverged first from the other Asian macaques. Subsequently, the four groups of Asian macaques diverged from one another in a relatively short period of time. Within each group, most of the species diverged in a relatively short period of time following the divergence of the groups. Assuming that the Asian macaques diverged from the outgroup Barbary macaque three million years ago (MYA), the divergence times among groups of Asian macaques were estimated at 2.1-2.5 MYA and within groups at 1.4- 2.2 MYA. The intraspecific nucleotide diversity observed among three rhesus macaques was so large that they did not form a monophyletic cluster in the phylogenetic trees. Instead, one of them formed a cluster with Japanese and Taiwanese macaques, whereas the other two formed a separate cluster. This implies that either polymorphisms of mtDNA sequences that existed before the divergence of these three species (ca. 700,000 years ago) have been retained in rhesus macaques or introgression has occurred among the three species.