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Many plants use colour to attract pollinators, which often possess colour vision systems well-suited for detecting flowers. Yet, to isolate the role of colour is difficult, as flowers also produce other cues. The study of florivory by Neotropical primates possessing polymorphic colour vision provides an opportunity to investigate the importance of colour directly. Here we determine whether differences in colour vision within a mixed population of wild dichromatic and trichromatic white-faced capuchins (Cebus capucinus imitator) affect flower foraging behaviours. We collected reflectance data for flower foods and modelled their chromatic properties to capuchin colour vision phenotypes. We collected behavioural data over 22 months spanning four years, determined the colour vision phenotype of each monkey based on amino acid variation of the L/M opsin gene from fecal DNA, and compared foraging behaviours of dichromats and trichromats. Most flowers were more conspicuous to trichromats, and trichromats foraged in small flower patches significantly more often. These data demonstrate a difference in wild primate foraging patterns based on colour vision differences, supporting the hypothesis that trichromacy enhances detection of small, ephemeral resources. This advantage, which may also extend to other foods, likely contributes to the maintenance of colour vision polymorphism in Neotropical monkeys.

Documenting primate life history characteristics is important because it provides information about traits that affect the timing and rate of reproduction in these long-lived species. This study describes the hormonal correlates of female reproductive events and quantifies for the first time key life history variables for Colobus vellerosus, using hormonal and observational data. This study also biologically validates that the reproductive events determined in the hormone profiles correspond to observed reproductive events for each female. We collected behavioural data on 18 females in our four study groups during 12 months (May 2012-2013) at the Boabeng-Fiema Monkey Sanctuary, using 10-min continuous focal and ad libitum sampling. We concurrently collected faecal samples (n = 1866) every 2-3 days from these 18 females (prepubescent n = 2, cycling n = 2, lactating n = 12, pregnant, n = 7, and post-reproductive n = 1) and extracted oestrogen (E2) and progesterone (P) metabolites in the field using solid-phase extraction cartridges. We created a hormone profile for each female by analyzing 1586 of our samples for E2 using radio-immuno assays, and P using enzyme-immunoassays at the Wisconsin National Primate Research Center. Mean ovarian cycle length was 24 days ± 1 (n = 2 cycles). Mean gestation length was 23 weeks (range = 21-25 weeks, n = 2 complete pregnancies). For females whose infants survived to nutritional independence, the mean inter-birth interval (IBI) was significantly longer than for females whose infants died prior to reaching nutritional independence (Mann-Whitney U Test; U = 14.5, p = 0.006; IBI surviving infants: 17.75 months, range = 8-20.75 months, n = 11 vs. IBI infant death: 11.89 months, range = 8-18.5 months, n = 9). The values for most life history traits reported in this study are similar to those documented in other similarly sized colobine species. Some values are on the lower end of the range for similarly sized colobines; C. vellerosus shows a cycle of 24 days and gestation length of 5.75 months vs. a range of 24-29 days for cycle length and 5.25-7.5 months for gestation length in other colobines. This may be due to C. vellerosus' smaller body size, or their limited access to higher quality food resources.

Much attention has been paid to hormonal variation in relation to male dominance status and reproductive seasonality, but we know relatively little about how hormones vary across life history stages. Here we examine fecal testosterone (fT), dihydrotestosterone (fDHT), and glucocorticoid (fGC) profiles across male life history stages in wild white-faced capuchins (Cebus capucinus). Study subjects included 37 males residing in three habituated social groups in the Área de Conservacíon Guanacaste, Costa Rica. Male life history stages included infant (0 to <12months; N=3), early juvenile (1 to <3years; N=10), late juvenile (3 to <6years; N=9), subadult (6 to <10years; N=8), subordinate adult (⩾10years; N=3), and alpha adult (⩾10years; N=4, including one recently deposed alpha). Life history stage was a significant predictor of fT; levels were low throughout the infant and juvenile phases, doubled in subadult and subordinate adults, and were highest for alpha males. Life history stage was not a significant predictor of fDHT, fDHT:fT, or fGC levels. Puberty in white-faced capuchins appears to begin in earnest during the subadult male phase, indicated by the first significant rise in fT. Given their high fT levels and exaggerated secondary sexual characteristics, we argue that alpha adult males represent a distinctive life history stage not experienced by all male capuchins. This study is the first to physiologically validate observable male life history stages using patterns of hormone excretion in wild Neotropical primates, with evidence for a strong association between fT levels and life history stage., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Efficient detection and selection of reddish fruits against green foliage has long been thought to be a major selective pressure favoring the evolution of primate trichromatic color vision. This has recently been questioned by studies of free-ranging primates that fail to show predicted differences in foraging efficiency between dichromats and trichromats. In the present study, we use a unique approach to evaluate the adaptive significance of trichromacy for fruit detection by undertaking a functional substitution model. The color vision phenotypes of neotropical monkeys are simulated for human observers, who use a touch-sensitive computer interface to search for monkey food items in digital images taken under natural conditions. We find an advantage to trichromatic phenotypes - especially the variant with the most spectrally separated visual pigments - for red, yellow and greenish fruits, but not for dark (purple or black) fruits. These results indicate that trichromat advantage is task-specific, and that shape, size and achromatic contrast variation between ripe and unripe fruits cannot completely mitigate the advantage of color vision. Similarities in fruit foraging performance between primates with different phenotypes in the wild likely reflect the behavioral flexibility of dichromats in overcoming a chromatic disadvantage., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The rapid disruption of tropical forests probably imperils global biodiversity more than any other contemporary phenomenon. With deforestation advancing quickly, protected areas are increasingly becoming final refuges for threatened species and natural ecosystem processes. However, many protected areas in the tropics are themselves vulnerable to human encroachment and other environmental stresses. As pressures mount, it is vital to know whether existing reserves can sustain their biodiversity. A critical constraint in addressing this question has been that data describing a broad array of biodiversity groups have been unavailable for a sufficiently large and representative sample of reserves. Here we present a uniquely comprehensive data set on changes over the past 20 to 30 years in 31 functional groups of species and 21 potential drivers of environmental change, for 60 protected areas stratified across the world’s major tropical regions. Our analysis reveals great variation in reserve ‘health’: about half of all reserves have been effective or performed passably, but the rest are experiencing an erosion of biodiversity that is often alarmingly widespread taxonomically and functionally. Habitat disruption, hunting and forest-product exploitation were the strongest predictors of declining reserve health. Crucially, environmental changes immediately outside reserves seemed nearly as important as those inside in determining their ecological fate, with changes inside reserves strongly mirroring those occurring around them. These findings suggest that tropical protected areas are often intimately linked ecologically to their surrounding habitats, and that a failure to stem broad-scale loss and degradation of such habitats could sharply increase the likelihood of serious biodiversity declines.

The frequency of anointing bouts and the materials used for self- and social anointing vary across capuchin species in captivity, but there is little published data on capuchin anointing in the wild. Here we present previously unpublished data on anointing behaviors from capuchin monkey populations at ten different field sites and incorporate these data into a review of the anointing literature for captive and wild capuchins. Using a comparative phylogenetic framework, we test four hypotheses derived primarily from captive literature for variation in anointing between wild untufted capuchins (Cebus) and tufted capuchins (Sapajus), including that (1) the frequency of anointing is higher in Cebus, (2) Cebus uses a higher proportion of plant species to insect species for anointing compared with Sapajus, (3) anointing material diversity is higher in Cebus, and (4) social indices of anointing are higher in Cebus. We found that wild Cebus anoints more with plant parts, including fruits, whereas wild Sapajus anoints more with ants and other arthropods. Cebus capucinus in particular uses more plant species per site for anointing compared with other capuchins and may specialize in anointing as an activity independent from foraging, whereas most other capuchin species tend to eat the substances they use for anointing. In agreement with captive studies, we found evidence that wild Cebus anoints at a significantly higher frequency than Sapajus. However, contrary to the captive literature, we found no difference in the range of sociality for anointing between Cebus and Sapajus in the wild. We review anointing in the context of other Neotropical primate rubbing behaviors and consider the evidence for anointing as self-medication; as a mechanism for enhanced sociality; and as a behavioral response to chemical stimuli., (© 2011 Wiley Periodicals, Inc.)

We report on the responses of Cebus capucinus in the Santa Rosa Sector of the Area de Conservación Guanacaste, Costa Rica, to the presence of observers over a 4-week period. Study groups were habituated to different degrees: (1) Cerco de Piedra (CP): continuous observations began in 1984; (2) Exclosure (EX): focus of an 18-month study on males from 1998 to 1999; and (3) NBH: never studied/followed but the group frequently encounters researchers. We collected three types of data: group scans (group state was coded as calm or agitated at observer presence), focal animal data (observer-directed behaviors were recorded), and fecal cortisol levels. The two less-habituated groups (NBH and EX) differed significantly from the habituated group (CP) in their behavioral and cortisol responses, and they showed an increase in habituation over the study period (agitation and cortisol levels both dropped). Individuals in NBH also decreased their responses to observers during focal follows; however, at the end of the study the responses of the two less-habituated groups (NBH and EX) remained elevated in comparison to the habituated group (CP), suggesting the need for further habituation. Unlike capuchin groups that rarely encounter humans, NBH and EX never fled from observers and they rarely emitted observer-directed alarm calls. We suggest that the permanence of habituation and the ability to habituate animals passively through a neutral human presence are both important considerations for researchers conducting studies in areas where animal safety from poachers, etc. cannot be guaranteed., (© 2007 Wiley-Liss, Inc.)

Observed patterns of variability in the food-processing behavior of white-faced capuchins (Cebus capucinus) across populations may reflect foraging traditions. However, there has been relatively little attention given to intrapopulation variability in food processing among groups and age/sex classes, making recent cross-population comparisons difficult to interpret. In this paper, we provide data on patterns of object use in foraging that we observed at Santa Rosa National Park, Costa Rica, for comparison with published data from a neighboring research site, Palo Verde National Park. We also describe the techniques used to process two food items consumed by Cebus capucinus at Santa Rosa, and discuss the factors that may underlie observed variability. We conducted a 6-month study on two groups of capuchins in 2001, and collected data on general activity and feeding patterns, rates and forms of object use, and distinctive processing techniques employed for two specific foods (Sloanea terniflora and Luehea candida). Rates of object-use behavior at Santa Rosa were considerably higher than those reported for Palo Verde and showed significant variation between groups and age/sex classes, as did patterns of Sloanea and Luehea processing. Observed differences in feeding rates between groups may reflect food availability or relative profitability, whereas variation between age/sex classes seems to reflect differences in the physical capabilities, foraging strategies, and the relative experience of mature and immature animals. Further research is needed to identify how a social context may influence the acquisition of food-processing techniques in juveniles and the development of foraging traditions in social groups., ((c) 2005 Wiley-Liss, Inc.)

This study explores the question of reproductive termination (loss of reproductive ability) in female Japanese macaques (Macaca fuscata) from the Arashiyama West (Texas) troop. We used a large sample of completed lives to identify reproductively terminated female Japanese macaques and to consider reproductive termination in Japanese macaques from a comparative life history perspective, which permits meaningful comparisons to be made with human female menopause. We classified a female as reproductively terminated if the time lag between last parturition and death exceeded two standard deviations of the female's own mean lifetime interbirth interval (Caro et al. [1995] Int. J. Primatol. 16:205-220). Seventy of the 95 females in the sample had at least 3 infants over their lifetime (the minimum required for the calculation of a mean and standard deviation), and thus were included in the analysis. Of these 70 females, 20 showed reproductive termination. Reproductively terminated females ranged in age from 14.5-32.7 years, although in females under age 25, reproductive termination was unlikely. The majority of females up to age 25 showed continued parturition. However, after age 25, reproductive termination was population-wide. Length of postreproductive life for reproductively terminated females varied from 0.07-4.4 years, with a mean of 2.08 years. Variation in length of postreproductive life was not related to the age at death of the female. While the occurrence of population-wide reproductive termination after 25 years does suggest similarities with human female menopause, the age at which this termination occurs is very late in the life span, and it was experienced by only 2.9% of the population. Female Japanese monkeys over age 25 are visibly aged and show outward signs of weakness and deterioration, quite unlike the healthy middle age of menopausal human females. Accordingly, as a life history characteristic, reproductive termination in Japanese macaques does not appear to coincide with menopause as experienced by human females., (Copyright 1999 Wiley-Liss, Inc.)

Theorists argue that mortality in male mammals should be higher than that of females, and many studies of primates followed across the life course have found this to be the case. This study examines mortality patterns in the rapidly expanding Arashiyama West (Texas) population of Japanese macaques (Macaca fuscata) and finds that males have a significantly lower median survival age (12.2 years) in comparison to females (20.5 years). Males and females are born in equal proportions, but by adulthood there are 2-5 females to every male. Males are at higher risk of falling victim to infectious diseases and human-related causes of death, and they are more likely to "disappear" from the population, which is inferred to result largely from emigration. There are no significant sex differences in the risks of dying from predation, non-infectious illnesses, neonatal defect, or social stress. Males become more susceptible to mortality than females once they reach sexual maturity, and they remain at greater risk than females until their old age. There is no evidence that one sex or the other is at greater risk of dying as infants, or as juveniles. Comparing males of different age classes, adolescent and adult males are more likely to die and to disappear than are juvenile males. These findings support the "high-risk, high-gain" hypothesis that males are mainly lost to the population because of their risk-taking behaviors after sexual maturity, rather than the "fragile male" hypothesis that males are more vulnerable to mortality during the period of growth and development.

A total of 54 free-ranging monkeys were captured and marked in Santa Rosa National Park, Costa Rica, during May 1985, and an additional 17 were captured during March 1986. The animals were darted using a blowpipe or a CO2 gun. The drugs used were Ketaset, Sernylan and Telazol. Ketaset was effective for Cebus capucinus but unsuccessful for Alouatta palliata and Ateles geoffroyi. Sernylan was successful for A. geoffroyi and A. palliata but is no longer commercially available. Telazol proved to be an excellent alternative capture drug for both A. palliata and A. geoffroyi.

The feeding patterns of three neighboring groups of Cebus capucinus were documented over a 3-year period in Santa Rosa National Park, Costa Rica. We describe the diets of the three groups and examine whether dietary differences between groups could be attributed to environmental differences in food abundances, to differences in the profitability of what was available or to learned local traditions. Diets were variable among groups; group A primarily ate fruit (81.2% of feeding time) and spent little time eating insects (16.9%), while group C was more heavily reliant on insects (44.3%) and ate less fruit (53.0%). Group B had a diet that was somewhat intermediate (69.8% fruit, 29.0% insects). By measuring the densities of all major plant foods, we were able to determine that many of the dietary differences between groups could not be attributed to simple measures of food abundance, but we could not distinguish between the food profitability hypothesis and local tradition hypothesis.

A long-term study of two groups of white-faced capuchins (Cebus capucinus) in Santa Rosa National Park in Costa Rica provides evidence of unusually high levels of vertebrate predation compared to those reported in other field studies of Cebus. The hunting techniques for different prey types are described, and several questions concerning vertebrate predation in primates are addressed. Why is there variation between individuals and between groups in the rate of predation? Why do males hunt more than females? Previous hypotheses to explain hunting in Old World primates are applied to this Neotropical example. Finally, I argue that successful vertebrate predation can readily arise in species like Cebus, which are characterized by opportunistic foraging patterns, manipulative and cognitive skills and well-developed techniques for locating and subduing invertebrate prey.

Lifetime reproductive success, measured by the number of offspring surviving to age five, varied from 0 to 10 in a group of 33 provisioned female Japanese macaques. Of the three contributors to reproductive success, the number of reproductive years, fecundity per year and survivorship of offspring to reproductive age, the first accounted for two-thirds of the variation. Fecundity per year and survivorship were negatively correlated, indicating reproductive costs of reducing interbirth interval. No other demographic measure used, nor the behavioral measure 'dominance rank', significantly correlated with lifetime reproductive success or its components. Age-specific changes in fecundity and infant survival were not found for this sample, neither could cessation of reproduction, even in very old females, be demonstrated.

The ranging patterns of two male and five female spider monkeys (Ateles geoffroyi) were studied with the use of radio telemetry in Santa Rosa National Park, Costa Rica. The average size of a spider monkey home range was 62.4 hectares; however, range size varied with sex, and, for females, with the presence of a dependent infant. The probability of encountering a radio-collared spider monkey in a three-hour search using radio telemetry (0.91) was much greater than using a visual search (0.20), and telemetric data resulted in a larger estimate of mean home range size than did observational data, when all subjects were compared. However, the difference appeared to be owing to the presence of male ranges in the telemetric, but not the observational, data. When the size of home ranges derived from radio-tracking data for adult females was compared to size of ranges for adult females derived from observations, the results were not significantly different. Adult males had larger home ranges than adult females, thus lending support to the hypothesis that males have adapted to the dispersion of females by occupying a large home range that overlaps the ranges of several adult females. The smallest home ranges were occupied by low-weight females with dependent infants, perhaps reflecting social and energetic constraints., (Copyright © 1988 Wiley‐Liss, Inc., A Wiley Company.)

Thirteen cases of a spontaneous neuropathy were observed among 150 free-ranging Japanese snow monkeys (Macaca fuscata). Necropsy of 4 individuals revealed lesions in the peripheral and central nervous systems. Pathologic changes were associated with ingestion of coyotillo berries (Karwinskia humboldtiana).

The Arashiyama West troop of Macaca fuscata consists of 150 monkeys transported from Kyoto, Japan, in February 1972 to their present location near Laredo, Texas. At this site the animals range over 108 acres and during the study period were only minimally provisioned and disturbed. In the report of a 3-year study of consort partner selection it was suggested that yearlong social bonds within the troop appeared to be distinct from consort bonds formed during mating season. Like Sade, we consider grooming to be a good measure of yearround affinitive bonding in macaques. This study compares the pairs formed for consorting with those formed for yearlong grooming activities, by the 94 individuals who formed consort relationships during the mating season of 1973--1974. It was found that yearlong grooming involved pairs of monkeys which were significantly different from those for consorting. Grooming was relatively frequent between related monkeys, while consort partners were seldom related. In addition, yearlong grooming patterns were not disrupted during mating season. These results suggest that consort and grooming activities are behavioral expressions of two important social networks, involving mutually exclusive social bonds.

This study describes and defines play in a laboratory colony of vervet monkeys (Cercopithecus aethiops). Play is divided into 26 units of behavior and the frequencies of these behaviors are recorded. Analysis shows that sex, age, and dominance have effects on some play behaviors. The play behavior of each age/sex group within the colony is described both quantitatively and qualitatively. Adult females are characterized by reinforcement of the play of immature monkeys, adult males by rough play with older male juveniles, four-year-old females and three-year-old males by stimulation of play in young infants, and males in general by a preference to play within their peer group. It is suggested that modified repetition of behaviors, diversified interactions, and innovative behaviors, are important qualities of play, and are essential to the adaptive plasticity of behavior in primates.