Inter‐group variability in seed dispersal by white‐handed gibbons in mosaic forest

Seed dispersers, like white‐handed gibbons (Hylobates lar), can display wide inter‐group variability in response to distribution and abundance of resources in their habitat. In different home ranges, they can modify their movement patterns along with the shape and scale of seed shadow produced. However, the effect of inter‐group variability on the destination of dispersed seeds is still poorly explained. In this study, we evaluate how seed dispersal patterns of this arboreal territorial frugivore varies between two neighboring groups, one inhabiting high quality evergreen forest and one inhabiting low quality mosaic forest. We predicted a difference in seed dispersal distance between the two groups (longer in the poor quality forest). We hypothesized that this difference would be explained by differences in home range size, daily path length, and ranging tortuosity. After 6 months of data collection, the evergreen group had a smaller home range (12.4 ha) than the mosaic group (20.9 ha), significantly longer daily path lengths (1507 m vs. 1114 m respectively) and greater tortuosity (39.1 vs. 16.1 respectively). Using gut passage times and displacement rates, we estimated the median seed dispersal distance as 163 m for the evergreen group (high quality forest) and of 116 m for the mosaic group (low quality forest). This contradiction with our initial prediction can be explained in term of social context, resource distribution, and habitat quality. Our results indicate that gibbons are dispersers of seeds between habitats and that dispersal distances provided by gibbons are influenced by a range of factors, including habitat and social context.     

Primates,Provisioning and Plants: Impacts of Human Cultural Behaviours on Primate Ecological Functions

Human provisioning of wildlife with food is a widespread global practice that occurs in multiple socio-cultural circumstances. Provisioning may indirectly alter ecosystem functioning through changes in the eco-ethology of animals, but few studies have quantified this aspect. Provisioning of primates by humans is known to impact their activity budgets, diets and ranging patterns. Primates are also keystone species in tropical forests through their role as seed dispersers; yet there is no information on how provisioning might affect primate ecological functions. The rhesus macaque is a major human-commensal species but is also an important seed disperser in the wild. In this study, we investigated the potential impacts of provisioning on the role of rhesus macaques as seed dispersers in the Buxa Tiger Reserve, India. We studied a troop of macaques which were provisioned for a part of the year and were dependent on natural resources for the rest. We observed feeding behaviour, seed handling techniques and ranging patterns of the macaques and monitored availability of wild fruits. Irrespective of fruit availability, frugivory and seed dispersal activities decreased when the macaques were provisioned. Provisioned macaques also had shortened daily ranges implying shorter dispersal distances. Finally, during provisioning periods, seeds were deposited on tarmac roads that were unconducive for germination. Provisioning promotes human-primate conflict, as commensal primates are often involved in aggressive encounters with humans over resources, leading to negative consequences for both parties involved. Preventing or curbing provisioning is not an easy task as feeding wild animals is a socio-cultural tradition across much of South and South-East Asia, including India. We recommend the initiation of literacy programmes that educate lay citizens about the ill-effects of provisioning and strongly caution them against the practice.     

Female Songs of the Nonduetting Javan Gibbons (<Emphasis Type="Italic">Hylobates moloch</Emphasis>) Function for Territorial Defense

Duets in territorial, pair-living primates may function to maintain intragroup cohesion, promote intergroup avoidance, and assist in territorial and resource defense, as well as advertising and reinforcing pair bonds. Despite the absence of duetting in Javan gibbons (Hylobates moloch), recent playback experiments suggested that Javan gibbon songs also play a role in pair-bond advertisement as well as territorial and mate defense. However, playback experiments only assess motivations of the listener, which may not reflect the motivations of the caller. We conducted an observational study of naturally occurring female songs in two groups of Javan gibbons from July 2009 to March 2010 and from March to November 2011 in Gunung Halimun–Salak National Park, Indonesia. We investigated female singing rates in relation to singing location, daily path length, occurrence of intergroup encounters, feeding rate, allogrooming rate, and distance between pair mates. The two females produced 47 songs during 164 observation days. Females in the area of their home range that overlapped with neighboring groups sang more frequently than expected based on time spent in the area of overlap vs. the home range interior. Groups also had longer daily path lengths on days when females sang than on nonsinging days, and on days when they visited the area of overlap than on nonvisiting days. Our findings indicate that, like the duets of other pair-living territorial primates, female Javan gibbon songs function for territorial defense, but we found no support for other functions such as intergroup avoidance, resource defense, and pair-bond reinforcement.     

Gut microbiota composition of Japanese macaques associates with extent of human encroachment

In recent decades, human–wildlife interaction and associated anthropogenic food provisioning has been increasing and becoming more severe due to fast population growth and urban development. Noting the role of the gut microbiome in host physiology like nutrition and health, it is thus essential to understand how human–wildlife interactions and availability of anthropogenic food in habitats can affect an animal's gut microbiome. This study, therefore, set out to examine the gut microbiota of Japanese macaques (Macaca fuscata) with varying accessibility to anthropogenic food and the possibility of using gut microbiota as indicator for macaques’ reliance on anthropogenic food. Using 16S ribosomal RNA gene sequencing, we described the microbial composition of Japanese macaques experiencing different types of human disturbance and anthropogenic food availability—captive, provisioned, crop‐raiding, and wild. In terms of alpha diversity, our results showed that observed richness of gut microbiota did not differ significantly between disturbance types but among collection sites, whereas Shannon diversity index differed by both disturbance types and sites. In terms of beta diversity, captive populations harbored the most distinctive gut microbial composition, and had the greatest difference compared with wild populations. Whereas for provisioned and crop‐raiding groups, the macaques exhibited intermediate microbiota between wild and captive. We identified several potential bacterial taxa at different taxonomic ranks whose abundance potentially could help in assessing macaques’ accessibility to anthropogenic food. This study revealed the flexibility of the gut microbiome of Japanese macaques and provided possible indices based on the gut microbiome profile in assessing macaques’ accessibility to/reliance on anthropogenic foods.     

The Hand That Feeds the Monkey: Mutual Influence of Humans and Rhesus Macaques (<Emphasis Type="Italic">Macaca mulatta</Emphasis>) in the Context of Provisioning

Historically, humans and other primates (primates henceforth) have coexisted across cultures and contexts, and many primate populations use anthropogenic food sources as their main or supplementary food. While primates may actively forage for such food, they are also directly provisioned by humans in many regions. Ethnoprimatology views humans and primates as cohabitants of integrated socioecological spaces who mutually influence each other’s ecologies and social lives. We contextualized provisioning of primates by humans within an ethnoprimatological framework and examined if the availability of anthropogenic food affected primate diets or the amount of time primates spent in anthropogenic habitats and whether primates influenced the human act of provisioning. To this end, we used scan sampling on a group of rhesus macaques across a year, and conducted interviews with 86 people who paused at a nearby tea shop for refreshments. We found that the macaques’ consumption of natural resources and dietary diversity decreased, and they spent more time in human-modified habitats when provisioned food was available. We also found that particular behaviors of the provisioned macaques stimulated provisioning by humans. Our findings show that provisioning influences macaque feeding ecology and habitat use, and that the behavior of the macaques themselves drives people to provide them with food subsidies, illustrating a complex web of interactions between the sympatric species.     

Diet and ranging behavior of the endangered Javan gibbon (Hylobates moloch) in a submontane tropical rainforest

Altitude influences forest structure and food abundance and distribution, which in turn affect primate feeding and ranging patterns. Javan gibbons (Hylobates moloch) are endemic to forests spanning a broad range of altitudes on Java, Indonesia. Most information about Javan gibbon behavior comes from studies in lowland forests, while the vast majority of wild gibbons remaining inhabit hill and lower montane forests. We studied the diets, activity patterns, and ranging behavior of three gibbon groups in hill/lower montane (950-1,100?m asl) forest in the Gunung Halimun-Salak National Park (GHSNP) from April 2008 to March 2009. The mean home range size was 37?ha and the mean daily path length was 1,180?m. The study groups spent 36% of time feeding, 41% resting, 15% traveling, 6% engaging in social behavior, and 2% in aggressive interactions. Fruit was the most important food (63% of feeding time) followed by leaves (24%), and flowers (12%). Our results suggest that Javan gibbons in higher elevation habitats have substantially larger home ranges than lowland populations, despite broad similarity in their activity budgets and diets. Conservation managers should consider the effects of altitude and habitat quality on gibbon ranging behavior when developing habitat corridors, selecting sites for translocation or reintroduction projects, and designating and managing protected areas.     

Ranging behavior of eastern hoolock gibbon (Hoolock leuconedys) in a northern montane forest in Gaoligongshan,Yunnan, China

Generally, food abundance and distribution exert important influence on primate ranging behavior. Hoolock gibbons (genus Hoolock) live in lowland and montane forests in India, Bangladesh, Myanmar and China. All information about hoolock gibbons comes from studies on western hoolock gibbons (Hoolock hoolock) living in lowland forest. Between August 2010 and September 2011, we studied the ranging behavior of one habituated group of eastern hoolock gibbon (H. leuconedys) living in a seasonal montane forest in Gaoligongshan, Yunnan, China. Results show that the study group did not increase foraging effort, calculated in this study as the daily path length, when fruit was less available. Instead, the gibbons fed more on leaves and decreased traveling to conserve energy. They relied heavily on a single food species in most study months which was patchily distributed within their total (14-month) home range, and during most months they used only a small portion of their total home range. In order to find enough food, the group shifted its monthly home range according to the seasonal availability of food species. To satisfy their annual food requirements, they occupied a total home range of 93 ha. The absence of neighboring groups of gibbons and the presence of tsaoko cardamom (Amomum tsaoko) plantations may also have influenced the ranging behavior of the group. Further long-term studies of neighboring groups living in intact forests are required to assess these effects.     

Effects of food and topography on ranging behavior of black crested gibbon (Nomascus concolor jingdongensis) in Wuliang Mountain, Yunnan, China

We studied the ranging behavior of a habituated group of black crested gibbons (Nomascus concolor jingdongensis) in a high, seasonal habitat on Mt. Wuliang, central Yunnan, China, between March 2005 and April 2006. Our results indicated that the total home range size for the study group was 129 ha, or 151 ha if the lacunae within the borders in which gibbons were not observed were included. This is a much bigger range size than that of other gibbon species. However, 69.7% of their activities occurred within 29 ha. The intensity of quadrant use was significantly correlated with the distribution of important food patches. The mean yearly daily path length was 1,391 m. Gibbons traveled farther when they spent more time feeding on fruit. To avoid often passing through ridges with little food, gibbons usually stayed in the same valley for successive days, and then moved on to another valley for another several days, which resulted in a concentrated ranging pattern.     

Darwin's Finch Begging Intensity Does Not Honestly Signal Need in Parasitised Nests

Parental care should be selected to respond to honest cues that increase offspring survival. When offspring are parasitised, the parental food compensation hypothesis predicts that parents can provision extra food to compensate for energy loss due to parasitism. Chick begging behaviour is a possible mechanism to solicit increased feeding from attending parents. We experimentally manipulated parasite intensity from Philornis downsi in nests of Darwin's small ground finch (Geospiza fuliginosa) to test its effects on chick begging intensity and parental food provisioning. We used in‐nest video recordings of individually marked chicks to quantify nocturnal parasite feeding on chicks, subsequent diurnal chick begging intensity and parental feeding care. Our video analysis showed that one chick per brood had the highest parasite intensity during the night (supporting the tasty chick hypothesis) and weakest begging intensity during the day, which correlated with low parental care and rapid death. We observed sequential chick death on different days rather than total brood loss on a given day. Our within‐nest video images showed that (1) high nocturnal larval feeding correlated with low diurnal begging intensity and (2) parent birds ignored weakly begging chicks and provisioned strongly begging chicks. Excluding predation, all parasite‐free chicks survived (100% survival) and all parasitised chicks died in the nest (100% mortality). Weak begging intensity in parasitised chicks, which honestly signalled recent parasite attack, was not used as a cue for parental provisioning. Parents consistently responded to the strongest chick in both parasitised and parasite‐free nests.     

猕猴觅食策略的地域性差异

本文通过1年连续野外观察（2015年12月至2016月11月）,对比研究了广东省内伶仃—福田国家级自然保护区野生群和台山上川岛猕猴省级自然保护区内半野生投食群的猕猴觅食策略异同,分析了食物数量和食物营养对猕猴觅食行为的影响,探讨了猕猴对生境变化的应对策略及其适应机制。研究结果表明,野生群和半野生群猕猴均偏好取食低灰分果实。另外,比起取食较少的果实,半野生群猕猴主要取食的果实具有低含水量和低蛋白等特点。全年来看,野生群猕猴偏好取食分布数量多的果实;半野生群猕猴食果频率月间波动受环境果实量影响,符合能量最大化理论。野生群和半野生群猕猴采食乔木树叶的行为均不受乔木数量的影响;而且两者采食所有生活型植被的叶的行为不受叶营养的影响。本研究表明猕猴会根据环境的食物资源数量和质量调整其觅食策略,具有对不同生活环境灵活适应的能力。     

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.     

人工投食行为对非人灵长类社会生态的影响

近年来,国内开展了对几种非人灵长类投食群的研究。借鉴日本猴(Macaca fuscata)、恒河猴(M.mulatta)和黑猩猩(Pan troglodytes)长期积累的研究经验,有利于我们减少人工投食对非人灵长类带来的不利影响,也有利于推动国内非人灵长类社会生态学的深入研究。本文通过比较这三种灵长类野生群和投食群之间的差异,量化地分析了人工投食对灵长类动物社会生态学方面的影响。结果表明在社会行为方面,人工投食会导致灵长类动物的聚集、增加争斗频率、改变个体间等级关系、延缓分群现象、影响活动域和增加新兴行为。在种群数量方面,人工投食会缩短雌性性成熟时间、增加出生率和婴猴存活率、减少成年猴死亡率。而人工投食对雄性性成熟、出生间隔等方面的影响仍需要进一步考证。最后我们讨论了人工投食行为的利弊,并建议应谨慎和妥善地实施人工投食,和对非人灵长类投食群进行合理的监管。     

Food provisioning and stone handling tradition in Japanese macaques: a comparative study of ten troops

By addressing the influence of food provisioning on stone handling (SH), a behavioral tradition in Japanese macaques, this study contributes to the ongoing debate in cultural primatology by asking whether human intervention influences the emergence or propagation of behavioral traditions. SH is a form of object play consisting of the manipulation of stones by performing various behavioral patterns. We tested the hypothesis that the frequency of food provisioning affects the daily performance, form, and context of occurrence of SH by influencing a troop's feeding-related activity budget. We used a standardized observation procedure to investigate SH in ten troops of Japanese macaques. In troops provisioned several times a day, SH was more frequent, longer, and more prevalent during provisioning than nonprovisioning periods. These effects of provisioning were not significant in troops provisioned less frequently. SH was more frequently integrated with food-related activities in troops supplied with food several times a day than in the other troops. Food provisioning may be a key factor in the innovation and transformation phases of the SH tradition in Japanese macaques.     

Changes in Demographic Parameters of <Emphasis Type="Italic">Macaca fuscata</Emphasis> at Takasakiyama in Relation to Decrease of Provisioned Foods

Though researchers have studied population dynamics extensively in many free-ranging primate populations, information on the relationship between food abundance and demographic parameters based on long-term data is sparse. We examined changes in demographic parameters in relation to decreased provisioning of foods based on data gathered for >50 yr in a provisioned, free-ranging Japanese macaque (Macaca fuscata) population at Takasakiyama, Oita Prefecture, in the southern region of Japan. At Takasakiyama, population size increased linearly because of heavy provisioning in the 1950s and 1960s. The provisioning of food to macaques decreased from 1965. We examined changes in the physique index of full adult females, primiparous age, birth percentage, infant mortality rate, population density, and annual population growth rate relative to the decrease of provisioned foods between the 1970s and 1990s. As a result of the drastic decrease in the amount of foods provisioned, the physique index of full adult females decreased, primiparous age increased, birth percentage decreased, and infant mortality rate increased, such that annual population growth rate diminished substantially. Ours is the first study to demonstrate quantitatively the relationship between provisioned food abundance and demographic parameters based on long-term data.     

Interspecific spacing between gibbons (Hylobates klossii) and langurs (Presbytis potenziani) on Siberut Island,Indonesia

Spatial arrangement and social interactions of two sympatric and ecologically similar primate species, Hylobates klossii and Presbytis potenzianai, are described from field observations made between July 1972 and October 1974 on Siberut Island, Indonesia. Gibbon territories and langur home ranges overlap extensively. Because gibbons have the ability to supplant langurs at shared food sources, langurs are at a competitive disadvantage. To avoid or decrease the frequency of hostile interactions with gibbons, langurs locate their core areas on boundaries between adjacent gibbon territories, which permits langurs to retreat across these barriers in response to gibbon movements. Langurs further enhance segregration by leaving their sleeping trees earlier than gibbons, gaining additional feeding time on contested food sources. This form of interspecific spatial organization between gibbons and langurs resembles certain predator-prey spacing systems, where territorial boundaries between adjacent predators serve as sanctuaries for prey populations.     

Dominance Style Among Macaca thibetana on Mt. Huangshan, China

The dominance style concept has proven useful for understanding covariation patterns in relationship qualities, particularly among macaques. However, the dominance styles of many macaques, including Tibetan macaques (Macaca thibetana), have not been examined in detail. We describe patterns of bidirectionality of aggression, postconflict affiliation and kin bias in a group of wild, but provisioned Tibetan macaques over a 2-yr period in order make an initial assessment of their dominance style. Bidirectional aggression, including percentage of counteraggression (1.9%), and conciliatory tendencies (6.4%) were consistently low across partner combinations, seasons and locations (forest vs. provisioning area). In addition, females consistently displayed high levels of kin bias in affiliation and tolerance. Compared with macaque species with better known dominance styles, the Tibetan data generally fell within the range for despotic species and outside the range for relaxed species. Although other researchers have tentatively classified them as tolerant or relaxed, we conclude that Tibetan macaques display a despotic dominance style. This conclusion poses complications to explanations based both on phylogenetic inertia and socio-ecological models.     

Nutrient contents predict the bamboo‐leaf‐based diet of Assamese macaques living in limestone forests of southwest Guangxi,China

Determining the nutrient factors influencing food choice provides important insight into the feeding strategy of animals, which is crucial for understanding their behavioral response to environmental changes. A bamboo‐leaf‐based diet is rare among mammals. Animals’ food choice and nutritional goals have been explained by several frameworks; however, the influence of nutrients on food choice in bamboo‐leaf‐based macaques is not yet available. Assamese macaques (Macaca assamensis) inhabiting limestone forests are characterized by such a bamboo‐leaf‐based diet, predominantly consuming young leaves of Bonia saxatilis, a shrubby, karst‐endemic bamboo. We studied the feeding behavior of one group of Assamese macaques using instantaneous scan sampling in limestone forests of the Guangxi Nonggang National Nature Reserve in southwest Guangxi, China. We compared the nutrient content of staple food and nonfood items and examine the role of key nutrients in the food selection of macaques. Our results showed that young leaves of bamboo B. saxatilis contained more water, crude protein, phosphorus, and less tannin than nonfood items. Furthermore, staple foods contained a higher content of water and less content of calcium than nonfood items. More specifically, quantities of water, crude protein, calcium, and phosphorus in food items were critical factors affecting feeding time on a specific plant item. Our results suggest that young bamboo leaves could meet macaques’ required protein and water intake, while enabling them to maintain their mineral balance, consequently facilitating to maintain the primates’ bamboo‐leaf‐diet in the limestone forest. Our findings confirm the effects of nutrient contents in food choice of Assamese macaques, highlighting the importance of the nutrient contents in maintaining their bamboo‐based diet and the need to increase the knowledge on their nutritional strategy adapted to the bamboo‐dominated diet inhabiting the unique limestone habitat.     

Responses of Javan Gibbon (Hylobates moloch) Groups in Submontane Forest to Monthly Variation in Food Availability: Evidence for Variation on a Fine Spatial Scale

Primates tend to prefer specific plant foods, and primate home ranges may contain only a subset of food species present in an area. Thus, primate feeding strategies should be sensitive to the phenology of specific species encountered within the home range in addition to responding to larger scale phenomena such as seasonal changes in rainfall or temperature. We studied three groups of Javan gibbons (Hylobates moloch) in the Gunung Halimun‐Salak National Park, Indonesia from April 2008 to March 2009 and used general linear mixed models (GLMM) and a model selection procedure to investigate the effects of variation in fruit and flower availability on gibbon behavior. Preferred foods were defined as foods that are overselected relative to their abundance, while important food species were those that comprised >5% of feeding time. All important species were also preferred. Season and measurements of flower and fruit availability affected fruit‐feeding time, daily path lengths (DPL), and dietary breadth. Models that included the availability of preferred foods as independent variables generally showed better explanatory power than models that used overall fruit or flower availability. For one group, fruit and preferred fruit abundance had the strongest effects on diets and DPL in the models selected, while another group was more responsive to changes in flower availability. Temporal variation in plant part consumption was not correlated in neighboring groups. Our results suggest that fine‐scale local factors are important determinants of gibbon foraging strategies. Am. J. Primatol. 74:1154‐1167, 2012. © 2012 Wiley Periodicals, Inc.     

Winter feeding of birds increases productivity in the subsequent breeding season

Supplementary food given to birds can have contemporary effects by reducing the risk of starvation, increasing survival and altering movements and reproductive performance. There is, however, a widely held perception that birds benefit from extra food over winter, but that it is better that they 'look after themselves' during breeding. Here we describe a landscape-scale experiment showing for the first time that the effects of increasing food availability only during the winter can be carried over to the subsequent breeding season. Even though food supplementation stopped six weeks prior to breeding, birds living on sites provisioned over winter had advanced laying dates and increased fledging success compared with birds living on unprovisioned sites. Thus, supplemental feeding of wild birds during winter, in a manner mimicking householders provisioning in gardens and backyards, has the potential to alter bird population dynamics by altering future reproductive performance. With levels of bird feeding by the public continuing to increase, the impacts of this additional food supply on wild bird populations may be considerable.     

Influence of Human Food Consumption on the Time Budget of Vervets

In nonhuman primate groups that utilize crop and food-raiding strategies, the inclusion of human food in the diet can have a pervasive impact on the activity budget. Human food is usually of higher quality and energy per unit than wild food. Thus, metabolic demands may be reached sooner on a human food diet, which in turn leads to improved foraging efficiency. In particular, a reduction in time spent feeding is associated with an increase in time spent resting. We investigated changes in time budget in a group of vervets (Chlorocebus aethiops pygerthrus) living in a tourist and cultivated area of Entebbe, Uganda (Zoo group). Saj collected 123 hr of focal observations on 17 individuals. Human food constituted half of the diet of Zoo group. Access to human food resulted in a high proportion of time resting and a low proportion of time feeding. The group had a small average daily range and a small home range in relation to other reports on unprovisioned groups. The time budget differences suggest that the juveniles spent more time moving and feeding and less time resting than adults did, and that adult females were more social than adult males were. These patterns have been observed in both provisioned and unprovisioned vervet groups. In contrast to a report from an unprovisioned group (Harrison, 1983), lactating females in this provisioned situation were able to maintain feeding and activity levels similar to those of other adults. However, adult females spent significantly less time consuming human food than juveniles did. We suggest that the risk associated with human food consumption may explain the low consumption of human food by females with dependent offspring. The differences may also reflect the different nutritional requirements of lactating females or the tendency for juveniles to be more exploratory than other group members.