Species-Specific Responses to Tourist Interactions by White-Faced Capuchins (Cebus imitator) and Mantled Howlers (Alouatta palliata) in a Costa Rican Wildlife Refuge

Ecotourism shows great potential for primate conservation, but further investigation through an ethnoprimatological lens is vital to understanding species-specific variation in human–nonhuman primate interactions at ecotourism sites. This study measured the rates and types of human–monkey interactions, the participants in these encounters, and the association between tourist numbers and interaction rates with white-faced capuchin monkeys (Cebus imitator) and mantled howlers (Alouatta palliata) at the Curú Wildlife Refuge in western Costa Rica. I collected data through 15-min all-occurrence samples for human–monkey interactions between January 2006 and December 2007. I recorded and analyzed a total of 1949 discrete interactions, representing one tourist group and one non-tourist group for each species. For both species, tourist groups showed more varied and more intense—but not more frequent—human–monkey interactions than the non-tourist groups. White-faced capuchins differed from mantled howlers in their greater frequency and more variable forms of human–monkey interactions. White-faced capuchins also showed a more gregarious pattern of interactions than mantled howlers, with most capuchin interactions being initiated by the monkeys and involving multiple actors. Although mantled howler human interaction rates correlated positively with levels of human traffic, white-faced capuchins did not show this relationship. These findings demonstrate that the differences in human–monkey interactions across species are an important consideration for the management of primate tourism sites. This study suggests that species-specific guidelines for ecotourism would reduce visitor impact on nonhuman primates.     

The Ontogeny of Prehensile‐Tail Use in Cebus capucinus and Alouatta palliata

A study of the platyrrhine prehensile tail provides an opportunity to better understand how ecological and biomechanical factors affect the ability of primates to distribute mass across many different kinds of arboreal supports. Young individuals experience ontogenetic changes in body mass, limb proportions, and motor skills that are likely to exert a strong influence on foraging strategies, social behaviors, support use, and associated prehensile‐tail use. In this research, I examine ontogenetic patterns of prehensile‐tail use in Cebus capucinus and Alouatta palliata. I collected behavioral data on activity, positional context, support size, and prehensile‐tail use in five age categories of white‐faced capuchins and mantled howlers during a 12‐month period at Estación Biológica La Suerte in northeastern Costa Rica. Infant and juvenile howlers and capuchins were found to use their prehensile tails significantly more often than adults during feeding, foraging, and social behavior. Prehensile‐tail use did not show predictable increases during growth. In both species, adults used their prehensile tails in mass‐bearing modes significantly less often than juveniles. Despite differences in tail anatomy in Cebus and Alouatta, prehensile‐tail use was observed to follow an increasing trajectory from infancy, peaking during juvenescence, and then decreasing in older juveniles and adults. In both species, it appeared that adult patterns of prehensile‐tail use reflected the demands placed on young juveniles. Am. J. Primatol. 74:770‐782, 2012. © 2012 Wiley Periodicals, Inc.     

Diets of some French guianan primates: Food composition and food choices

We studied food choices of black spider monkeys (Ateles paniscus) and red howlers (Alouatta seniculus) in an undisturbed tropical forest of French Guiana for 6 months in the rainy season. We made additional observations on tufted capuchins (Cebus apella) and examined the differences and similarities in feeding behavior with respect to the plant specific composition of the habitat and the biochemical characteristics of their food. Capuchins and spider monkeys mainly fed on ripe fruit pulp, to which they added invertebrates (capuchins) or young leaves (spider monkeys); their plant diet was more varied than that of howlers, which contained approximately equal proportions of ripe fruits and young leaves of many species, with large monthly variations in these food categories. BothAteles andAlouatta tended to feed preferentially on abundant plant species, with a large overlap in their fruit choices. The former species included a high proportion of soluble sugars and a low proportion of protein in its diet compared, to howlers. Leaves of several species selected byAlouatta reacted positively when screened for alkaloids and phenolic compounds. Besides specific metabolic requirements and adaptations to deriving nutrients from distinct food types, we hypothesize that the ability to taste sugars, which varies among primates, affects the range of foods appearing to be palatable and, consequently, contributes to the differentiation of feeding niches.     

Group size in folivorous primates: ecological constraints and the possible influence of social factors

The ecological-constraints model assumes that food items occur in depletable patches and proposes that an increase in group size leads to increased day range due to more rapid patch depletion. Smaller groups become advantageous when an increase in travel costs is not repaid by an increase in energy gained or some other fitness advantage. On the other hand, we also know that group size can be influenced by social factors. Here we contrast the diet and group size of red colobus (Procolobus badius) and black-and-white colobus (Colobus guereza) in Kibale National Park, Uganda to consider how ecological and social factors are affecting their group sizes. Subsequently, we examine whether the insights gained from this detailed comparison can provide an understanding of why the social organization and group size of mantled howlers (Alouatta palliata) and black howlers (A. pigra) differ. Two groups of red colobus and two groups of black-and-white colobus were studied over 10 months. Red colobus groups were larger (48 and 24) than black-and-white colobus groups (9 and 6). The two groups of red colobus overlap home ranges with the two groups of black-and-white colobus; 75% and 95% of their home ranges were within red colobuss home range. There was a great deal of similarity in the plant parts eaten by the two species and both species fed primarily on young leaves (red colobus 70%, black-and-white colobus 76%). In terms of the actual species consumed, again there was a great deal of similarity between species. The average dietary overlap among months for the two neighboring groups of red colobus was 37.3%, while the dietary overlap between the red colobus and the black-and-white colobus group that had its home range almost entirely within the home range of the red colobus groups averaged 43.2% among months. If ecological conditions were responsible for the difference in group size between the two colobine species, one would expect the density of food trees to be lower in the home ranges of the black-and-white colobus monkeys, since they have the smaller group size. We found the opposite to be true. Both black-and-white colobus groups had more food trees and the cumulative size of those trees was greater than those in the red colobuss home ranges. We quantify how these differences parallel differences in mantled and black howlers. The average group size for mantled howlers was 12.9 individuals, and for black howlers it was 5.3 individuals. We explore possible social constraints, such as infanticide, that prevent black-and-white colobus and black howlers from living in large groups.This revised version was published online in April 2005 with corrections to the cover date of the issue.     

Seasonal variation in the positional behavior of red howling monkeys (Alouatta seniculus)

Recent studies on the positional behavior of primates reveal that significant seasonal variation occurs in both locomotion and postures that is related to changes in diet and foraging techniques. Howling monkeys (genusAlouatta), which also have a seasonally varied diet, are predicted to have correspondingly varied positional behaviors. Two groups of red howling monkeys were studied in a primary rain forest in French Guinana during the dry and wet seasons. During the dry season, when howler diet is based mainly on leaves, howlers traveled more frequently by quadrupedal walking on large supports, a mode of progression that is probably inexpensive energetically and relatively stable. During feeding, quadrupedal and tripedal stand contributed considerably, a posture probably associated with the equal distribution of leaves within a tree crown. In contrast, during the wet season, when fruit was abundant, howlers fed very frequently by sitting on large supports, probably because fruit consumption required more time for special manipulation. However, most seasonal changes in feeding postures, and in travel and feeding locomotion, were difficult to associate directly with dietary shifts. These behavioral changes may be more highly correlated with slight modifications in microhabitat use (horizontal and vertical daily ranges, similar and alternative arboreal pathways) that are not considered in this paper.     

Diet and activity in black howler monkeys (<Emphasis Type="Italic">Alouatta pigra</Emphasis>) in southern Belize: does degree of frugivory influence activity level?

This study reports on the diet and activity budgets of Central American black howling monkeys (Alouatta pigra) at Monkey River, Belize. This is a previously unstudied population, close to the southern boundary of the species range, and it provides comparative data on A. pigra from a new study site. Both diet and activity are within the ranges reported for other A.pigra sites and for mantled howlers (A. palliata). No age-sex differences could be discerned in either diet or activity, though monthly variation was apparent. The monkeys switch from consuming leaves 86% of the time in January to March to consuming 67% fruit in April to July. This difference was statistically significant, and provided the opportunity to compare activity levels of the monkeys over two dietary periods, one characterized primarily by folivory, the other by frugivory. Howlers are often seen as a relatively inactive species, something that is associated with a low quality, folivorous diet. However, A. pigra have been described as being as frugivorous as possible and as folivorous as necessary. Yet, despite the opportunistic consumption of large quantities of high-energy foods, A. pigra has been observed as conforming to the howler lifestyle, resting as much as 80% of the day. The data in this paper support both of these reports. Black howlers at Monkey River Belize are typically inactive, maintaining high levels of inactivity even during months characterized by frugivory, suggesting that diet is more flexible and varied than is behavior and calling into question the assumption that howler inactivity is due to the digestion of large quantities of leaves.     

The Repertoire and Social Function of Facial Displays in Cebus capucinus

Systematic studies on facial displays in capuchins are limited and based mainly on studies of tufted capuchins (Cebus apella). Despite the great social-morphological variability within Cebus suggesting possible morphological and functional variations in the facial displays of different species, no study has considered thoroughly visual communication in the genus. Our aim was to describe the facial displays of white-faced capuchins and to assess their distribution and communicative function. We observed 15 captive white-faced capuchins in the Primate Centre of the Louis Pasteur University of Strasbourg, for a total of 198 h. We described the following facial displays: relaxed open-mouth, lip-smacking, open-mouth threat-face, silent bared-teeth, open-mouth silent bared-teeth, protruded-lip face, and tongue-out. We never observed the scalp-lifting display, one of the most common displays characterizing tufted capuchins. White-faced capuchins use the majority of facial displays in an affiliative or playful context; only the open-mouth threat-face display is associated with aggressive behaviors. White-faced capuchins lack ritualized signals of submission. The fact that in white-faced capuchins the silent bared-teeth display conveys only a positive message, while in tufted capuchins it signals submission as well as affiliation, supports the covariation hypothesis (Thierry 2004 Social epigenesis. In B. Thierry, M. Singh, & W. Kaummans (Eds.), Macaque societies: A Model for the study of social organization, pp. 267–294. Oxford University Press).     

Forest Fragmentation Modifies Habitat Quality for Alouatta palliata

Fragmentation reduces habitat area, increases the number of habitat patches, decreases their size, and increases patch isolation. For arboreal mammals such as howlers (Alouatta palliata), canopy modifications from fragmentation processes could also negatively affect habitat quality. We analyzed changes in the composition and plant structure of 15 fragments (1–76 ha) and compared them with vegetation from a continuous tropical rain forest reserve (700 ha) in Los Tuxtlas, Mexico. At each site, we sampled 1000 m² of all trees, shrubs, and lianas with a diameter at breast height (DBH) ≥10 cm. We obtained estimates of species richness, density, and basal area for different ecological groups, DBH ranges, and top food resources for howlers. We used a stepwise multiple regression analysis to determine relationships between fragment characteristics (size, shape index, and isolation) and plant variables. Compared to continuous forest, fragments have altered composition and plant structure, with large trees absent from the canopy. The basal area of top food resources is higher in continuous forest. Fragment size is the best explanation for the differences in composition and plant structure. The largest fragments had greater basal area of top food resources and more large primary trees in the canopy. Overall, our results suggest that fragmentation altered the habitat quality for howlers.     

Cougar Predation on Black-and-Gold Howlers on Mutum Island,Southern Brazil<Superscript>1</Superscript>

Researchers consider predation rates by terrestrial animals to be lower in the case of arboreal primates, particularly among large-bodied species. We recorded the consumption of black-and-gold howlers (Alouatta caraya) by cougars (Puma concolor) as evidence of predation on an island of the upper Paraná River. We collected and processed fecal samples of the felid in 2004 and 2005. We identified items in the laboratory by comparison with museum specimens. We considered each species in a fecal sample as a single occurrence. Based on analysis of the cuticle scale pattern, we identified the felid as cougar. Howlers occurred in 4 out of the 8 fecal samples (40% of the occurrences). In addition to howlers, we also recorded 5 occurrences of agouti (Dasyprocta azarae; 50%) and a small unidentified sigmodontine rodent (10%). The abundance of howlers and the low forest canopy in a successional vegetation might have facilitated the predation of the large primates by a primarily terrestrial predator. The versatility of cougars is corroborated by the consumption of prey species that were abundant in the region and that were available in different forest strata, such as howlers and agoutis.     

Look before leaping: foraging selectivity of capuchin monkeys on acacia trees in Costa Rica

Acacia trees in Costa Rica have an obligate mutualism with three species of Pseudomyrmex ants, which vigorously defend their host tree from insect and mammalian herbivores. Depending on the size and species of ant colony, individual acacia trees may be differentially protected. For animals able to discern between weakly and highly aggressive ant colonies, costs of ant stings from less active colonies might be offset by nutritional value acquired from feeding on acacia fruit or ant larvae in swollen thorns. We examined foraging selectivity of capuchin monkeys on acacia trees in Santa Rosa National Park, Costa Rica. We measured four characteristics of the acacia trees from which capuchins fed and of acacias immediately adjacent to those in which the monkeys fed: diameter at breast height (DBH), accessibility, species of closest tree and ant species present. We found that capuchins prefer to forage in acacias that are large and accessible. We also made two measurements of ant colony activity on each tree, one before and one after disturbing the ant colony. We found that the three species of mutualistic ants differ in baseline activity levels and that mutualistic ants are more active than non-mutualistic ant species found in acacia trees. We also found that capuchins foraged more frequently in trees colonized by non-mutualistic ants, but the explanatory value (r ²) of this model was low. Furthermore, monkeys did not discriminate between acacias on the basis of baseline ant activity or the ant colony’s response to disturbance. We conclude that these monkeys select acacia trees in which to forage based on characteristics of the trees rather than the ants. In addition, our study suggests that white-faced capuchins act as predators on the acacia ants but they probably benefit the dispersal and reproductive success of acacia trees. Capuchins may in fact function as an additional mutualistic partner for acacia trees via seed dispersal, but they must overcome the ants’ defense of the trees to do so.     

Serum Nutritional Profiles of Free-Ranging <Emphasis Type="Italic">Alouatta Caraya</Emphasis> in Northern Argentina: Lipoproteins; Amino Acids; Vitamins A,D, and E; Carotenoids; and Minerals

Quantifying circulating nutrient concentrations in sera of free-ranging subjects will help to establish a basis from which we can evaluate the nutritional status and needs of the captive population. We collected serum samples from 26 free-ranging black-and-gold howlers (Alouatta caraya) in San Cayetano forest in northern Argentina. We analyzed them for concentrations of lipoproteins; amino acids; vitamins A, D, and E; carotenoids; and minerals. There are a few significant differences between sexes in concentrations of high-density lipoprotein cholesterol, certain amino acids, vitamin E, lutein + zeaxanthin, and copper. Most nutritional parameters are similar to the ones measured in free-ranging Mexican mantled howlers (Alouatta palliata mexicana) and in captive New World primates (NWPs). Carotenoid, vitamin D, and phosphorus concentrations are the exceptions. Carotenoid concentrations are higher in free-ranging Alouatta caraya than reported for other free-ranging and captive species. Vitamin D concentrations are 14 times greater in the free-ranging black-and-gold howlers than in captive NWPs. Phosphorus concentrations are also higher than expected and higher than typically occur in captive primates, leading to a 1:1.6 calcium:phosphorus ratio. Because we based our study on a small number of free-ranging howlers, additional samples from different regions and throughout the year would better define desirable nutritional parameters for captive howlers.     

Species differences in substrate use and diet between sympatric galagos in two Kenyan coastal forests

Two sympatric species of galago,Galago zanzibaricus andG. garnettii, differed significantly in diet (judged mainly by fecal analysis) and substrate use at each of two coastal forest sites in Kenya. The larger-bodiedG. garnettii tended to use branches of a bigger diameter and to be found higher in the canopy than the smallerG. zanzibaricus. The latter species spent most of its time in the thick undergrowth. The larger species ate more fruit than the smaller. There were differences between the species in the type of invertebrate prey remains found in the feces. It is suggested that the differences in habitat use were influenced by the different body sizes of the two galagos, which so enabled these closely related species to coexist. Order of authorship is alphabetical.     

Tropical rain forest fragmentation and wild populations of primates at Los Tuxtlas,Mexico

In view of the extensive destruction, fragmentation, and conversion of primate habitats to anthropogenic vegetation, information on Neotropical primate ability to use a landscape consisting of forest fragments and agricultural habitats is necessary to understand the ecological flexibility of the species involved and it is of relevance to the design of conservation scenarios at the landscape level. We censused howlers and spider monkeys in 126 forest fragments and in 44 agricultural sites at Los Tuxtlas, southern Veracruz, Mexico, and used the IDRISI Geographic Information System to assess the extent of primate habitat remaining. We conducted economic surveys to assess the productivity of several systems of land management, including cattle ranching. Seventy- five percent of native habitat has been lost at Los Tuxtlas, 20% remains in the form of isolated fragments of vegetation, and only 5% consists of contiguous rain forest at high elevations (> 800 m). Howlers and spider monkeys were present in only 60 and 8% of the forest sites investigated, respectively, attesting to extensive local extinction. Populations of both species are small and found in a fragmented and isolated condition throughout the landscape. A large number of howlers were detected in artifactual habitats such as cacao, coffee and mixed (cacao and coffee) plantations shaded by rain forest trees. Residual rain forest vegetation along streams, rivers, and lakes facilitated the interfragment and interhabitat movement of howlers. Economic surveys showed that some of the agricultural habitats were more productive than cattle ranching. Conservation of spider monkeys requires large areas of contiguous forest, but only small areas are present at high elevations. Howlers are restricted to elevations < 800 m, where most of the forest has disappeared, but can apparently exist in a matrix of forest fragments, arboreal agricultural habitats, and pasture lands. Structural aspects of the vegetation and connectivity among habitat islands may be indispensable components of both landscape scenarios.     

Effective Solutions for Howler Conservation

All species of howlers are at risk due to habitat destruction. I review the three most serious threats to howler survival—agricultural disturbance, logging disturbance, and hunting—both in a general context and in species accounts. Withal, the adaptability of howlers has allowed them to be a widespread genus throughout Central and South America. Alouatta is especially adaptable due to its generalized folivorous diet. I discuss how this adaptability relates to survival in situations of habitat destruction. In addition, I examine howler preference for riverine habitats and their ability to capitalize on secondary forests and secondary growth. I take a practical approach, suggesting some solutions for species survival, which include habitat management and reintroductions. Successful conservation models highlighted are the Community Baboon Sanctuary in Belize, in which villagers manage their lands for the benefit of the black howler and for their own economic benefit from ecotourism, and a village-sustainable logging system in Quintana Roo, Mexico, which has reduced levels of deforestation while benefiting both villagers and howlers, as well as other wildlife. A successful translocation of black howlers in Belize in order to reestablish a viable population is also described. The paper concludes with further suggestions to encourage howler conservation.     

Resource use by howler monkeys (Alouatta palliata) in the rain forest of Los Tuxtlas,Veracruz, Mexico

The feeding preferences of howler monkeys at their northernmost distribution in the Neotropics are reported for an annual cycle. A remarkable selectivity for 27 species representing 15 families was observed. The Moraceae and Lauraceae plant families were the most important in the diet. The howlers spent an almost equal proportion of their feeding time eating leaves and fruit, and displayed a marked preference for young leaves and mature fruit. The consumption of different plant parts was markedly seasonal and the howlers’ ranging behavior was closely associated with the availability of young leaves and mature fruit. Their home range was unusually large (ca. 60 ha) for howlers and the food species exploited occur at very low densities (93%, ≤ 4 ind/ha). They chose food items richer in protein and energy. Alkaloid compounds, present in some of the leaves, play a secondary role in their dietary selectivity.     

Seismic activity response as observed in mantled howlers (Alouatta palliata), Cuero y Salado Wildlife Refuge, Honduras

This report documents the response of wild mantled howlers (Alouatta palliata) to coseismic activity (seismic activity at the time of an earthquake). During field work on the north coast of Honduras, data were collected on a habituated troop of mantled howlers as they responded to coseismic activity. The seismic event occurred on 13 February 2001 at 0822 hours local time with a magnitude of Richter scale 6.6, focus depth of approximately 15 km at a distance of 341 km from the epicentre to the field site, Cuero y Salado. At the field site, based upon Jeffreys and Bullen (1988), body waves, noted as P and S waves, arrived at 60 and 87 s, respectively, with surface waves arriving approximately 103 s post-origin time of the seismic event. While there are three reports on non-human primate response to coseismic activity in the literature, they report on captive non-human primates. This is the first documented response on a non-captive troop. In addition, this report compares the intensity measure encountered by a wild troop of howlers and one captive group of orangutans as set out by the Modified Mercalli Intensity scale. The Modified Mercalli measure of intensity is one of two standard measures of seismic activity and rates what a person sees and feels at their location (Wood and Neumann 1931; Richter 1958). Thus, arboreal nonhuman primates are found to respond to coseismic activity ranging from Level IV to Level VI as based upon the modified Mercalli intensity scale.     

Fruit diet of Alouatta guariba and Brachyteles arachnoides in Southeastern Brazil: comparison of fruit type, color, and seed size

Fruit is an important food resource for neotropical primates. In this study I compare the fruit diet of sympatric brown howlers (Alouatta guariba) and southern muriquis (Brachyteles arachnoides). Feeding behavior was studied over 12 months and fruit species consumed were identified and assigned to the categories fruit type, fruit color, and seed size. Observed-fruit feeding records were compared with expected records determined from local availability of the fruit of the tree species. I also determined dietary overlap. Fruit consumption occupied 8 and 12% of the feeding time of A. guariba and B. arachnoides, respectively. Fruit from eight tree species were consumed by the former and fruit from twenty-two species by the latter. Patterns of fruit selection of A. guariba and B. arachnoides varied widely. Although howlers and muriquis converge behaviorally by selecting fruit with common attributes (fleshy/unprotected, violet and brown/black-colored), unlike A. guariba, B. arachnoides fed on immature seeds of fleshy/protected and dry fruit. Large seeds were ingested, and defecated intact, by B. arachnoides only. There was little overlap of fruit diet even within categories that had been selected by both, suggesting that dietary divergence is occurring at the interspecific level. Different resource exploitation probably mediates the coexistence of A. guariba and B. arachnoides in low diversity, semideciduous forests, where the environment imposes narrow limits on primate food choices.     

Telemetry System for Assessing Jaw-Muscle Function in Free-ranging Primates

In vivo laboratory-based studies describing jaw-muscle activity and mandibular bone strain during mastication provide the empirical basis for most evolutionary hypotheses linking primate masticatory apparatus form to diet. However, the laboratory data pose a potential problem for testing predictions of these hypotheses because estimates of masticatory function and performance recorded in the laboratory may lack the appropriate ecological context for understanding adaptation and evolution. For example, in laboratory studies researchers elicit rhythmic chewing using foods that may differ significantly from the diets of wild primates. Because the textural and mechanical properties of foods influence jaw-muscle activity and the resulting strains, chewing behaviors studied in the laboratory may not adequately reflect chewing behaviors of primates feeding in their natural habitats. To circumvent this limitation of laboratory-based studies of primate mastication, we developed a system for recording jaw-muscle electromyograms (EMGs) from free-ranging primates so that researchers can conduct studies of primate jaw-muscle function in vivo in the field. We used the system to record jaw-muscle EMGs from mantled howlers (Alouatta palliata) at Hacienda La Pacifica, Costa Rica. These are the first EMGs recorded from a noncaptive primate feeding in its natural habitat. Further refinements of the system will allow long-term EMG data collection so that researchers can correlate jaw-muscle function with food mechanical properties and behavioral observations. In addition to furthering understanding of primate feeding biology, our work will foster improved adaptive hypotheses explaining the evolution of primate jaw form.     

Behavior, Diet, and Movements of the Sulawesi Crested Black Macaque (Macaca nigra)

We present the first field study of activity budgets, diet and ranging patterns of the Sulawesi crested black macaque, Macaca nigra, one of seven macaque species endemic to Sulawesi, Indonesia. We studied three crested macaque groups, ranging in size from 50 to 97 individuals, for 18 months in the Tangkoko-DuaSudara Nature Reserve, North Sulawesi. They spent 59% of the day moving and procuring food, especially fruits, and 41% of the day resting and socializing. Their diet is composed of more than 145 species of fruit (66% of observed feeding bouts), vegetative material (2.5%), invertebrates (31.5%), and occasional vertebrate prey. Group differences were more pronounced than seasonal or diurnal differences. Specifically, the largest group moved farther during the day, moved at a faster and more uniform rate, ate less fruit, rested more, and socialized less than the smaller groups did. The largest group had the largest home range, but it included less primary forest and more disturbed habitat than the ranges of smaller groups. There are individual differences in activity budgets of adult males and females in time spent moving, resting, feeding, and socializing that may reflect differences in reproductive strategies of males versus females. The behavior of large juveniles is more similar to that of adults than to that of small juveniles. Daily movements and use of home range are correlated with diet. Macaques moved shorter distances as the proportion of time spent feeding on fruit increased, and the top four dietary items accounted for most of the variance in entry into hectare blocks of home range.     

Habitat selection and seasonal patterns of activity and foraging of mantled howling monkeys(Alouatta Palliata) in Northeastern Costa Rica

I conducted a 15- month ecological study of habitat preferences and activity and foraging patterns of two troops of mantled howling monkeys, Alouatta palliata,in a lowland rain forest at La Selva Biological Reserve in northeastern Costa Rica. The two troops specialized on different habitats in spite of the fact that both of them had all habitats available and were not constrained by neighboring troops since the population density of howlers is low (7- 15 howlers/km ² ).Troop 1 spent the majority of time in primary forest (80%) followed by secondary forest (10%), while troop 2 spent the majority of time in undisturbed riparian habitat (60%) followed by primary forest (30%). Habitat sampling indicates that neither the total number of stems, species, or families nor the diversity (Shannon index) or evenness is a good indicator for howler habitat selection. Instead the density of trees from the 12 species most commonly consumed by each troop is the most important factor. Activity and foraging patterns were not dependent upon the season as has been described for howling monkeys in forests with a more pronounced dry season at Barro Colorado Island, Panama, and La Pacifica in northwestern Costa Rica. This is likely a result of the more constant food supply at La Selva, combined with less intraspecific competition due to the low howler density. The intraspecific variability of foraging patterns and troop- specific habitat specialization observed in Alouatta palliatashould be considered in the conservation biology of primates. Primate relocation programs should include not only an ecological assessment of the release site but also a comparison of the release site with the habitat that the groups currently occupy.