Survival rates and mortality risks of Plecturocebus cupreus at the California National Primate Research Center.

This article describes survivorship and explores factors affecting mortality risks in a captive colony of coppery titi monkeys (Plecturocebus cupreus) housed at the California National Primate Research Center (CNPRC), at UC Davis, in Davis, CA. We analyzed data collected on individuals since the colony's creation in the 1960s, with a sample of 600 animals with partially complete information (date of birth, age at death, body mass, parental lineage). We used three methods: (1) Kaplan-Meier regressions followed by a log-rank test to compare survival in male and female titi monkeys, (2) a breakpoint analysis to identify shifts in the survival curves, and (3) Cox regressions to test the effect of body mass change, parental pair tenure, and parental age on mortality risk. We found that males tend to have a longer median lifespan than females (14.9 and 11.4 years; p = 0.094) and that survival decreases earlier in males than in females during adulthood (9.8 and 16.2 years). A body mass loss of 10% from adulthood to the time of death led to a 26% higher risk of dying (p < 0.001) as compared to an individual with stable body mass. We found no evidence of sociobiological factors on mortality risks (parental age, parental pair tenure), but an exploratory analysis suggested that a higher rate of offspring conceptions increases mortality risks. This description of factors influencing survival and mortality in titi monkeys is a first step toward understanding aging in this species to consider titi monkeys as a primate model for socioemotional aging.

Do zoo-housed primates retreat from crowds? A simple study of five primate species.

An animal's welfare state is directly influenced by the mental state, which is shaped by experiences within the environment throughout the animal's life. For zoo-housed animals, visitors to the zoo are a large part of that environment and a fluctuating influence within it. This study examines the impact of zoo visitors on the space use of five species of zoo-housed primates (Eastern black-and-white colobus monkeys, Colobus guereza, n = 5, Allen's swamp monkeys, Allenopithecus nigroviridis, n = 2, DeBrazza's monkeys, Cercopithecus neglectus, n = 3, Bolivian gray titi monkeys, Callicebus donacophilus, n = 3, and crowned lemurs, Eulemur coronatus, n = 3). Specifically, we considered whether primates' distance from visitor areas changed as crowd sizes increased. Data were collected using the ZooMonitor app. Observers recorded spatial coordinates for each animal over periods ranging from 12 to 32 months. Data were analyzed using two types of regression models (linear and logistic) to examine the influence of visitors on the location of the primates. Both analyses revealed a statistically significant but small decrease in primate distance from visitor viewing glass as the number of visitors increased. Behavioral indicators of welfare were also unaffected by the presence of visitors. These results suggest that, with additional validation, distance from visitors may be one promising, simple way to evaluate the influence of visitors on primate welfare.

Individuality in the vocalizations of infant and adult coppery titi monkeys (Plecturocebus cupreus).

As social animals, many primates use acoustic communication to maintain relationships. Vocal individuality has been documented in a diverse range of primate species and call types, many of which have presumably different functions. Auditory recognition of one's neighbors may confer a selective advantage if identifying conspecifics decreases the need to participate in costly territorial behaviors. Alternatively, vocal individuality may be nonadaptive and the result of a unique combination of genetics and environment. Pair-bonded primates, in particular, often participate in coordinated vocal duets that can be heard over long distances by neighboring conspecifics. In contrast to adult calls, infant vocalizations are short-range and used for intragroup communication. Here, we provide two separate but complementary analyses of vocal individuality in distinct call types of coppery titi monkeys (Plecturocebus cupreus) to test whether individuality occurs in call types from animals of different age classes with presumably different functions. We analyzed 600 trill vocalizations from 30 infants and 169 pulse-chirp duet vocalizations from 30 adult titi monkeys. We predicted that duet contributions would exhibit a higher degree of individuality than infant trills, given their assumed function for long-distance, intergroup communication. We estimated 7 features from infant trills and 16 features from spectrograms of adult pulse-chirps, then used discriminant function analysis with leave-one-out cross-validation to classify individuals. We correctly classified infants with 48% accuracy and adults with 83% accuracy. To further investigate variance in call features, we used a multivariate variance components model to estimate variance partitioning in features across two levels: within- and between-individuals. Between-individual variance was the most important source of variance for all features in adults, and three of four features in infants. We show that pulse-chirps of adult titi monkey duets are individually distinct, and infant trills are less individually distinct, which may be due to the different functions of the vocalizations.

Observations of infant conflict and avoidance in San Martin titi monkeys (Plecturocebus oenanthe).

Conflict between caregivers and infants typically centers on disagreements over the amount and frequency of care provided. Prior research has identified many variables that impact patterns of mother-infant conflict. These include wide-ranging factors such as individual temperament, reproductive status, and availability of resources. By contrast, no studies have investigated the variables that influence father-infant conflict. To better understand the nature of caregiver-infant conflict in a species with obligate biparental care, I observed two groups of San Martin titi monkeys inhabiting disturbed secondary forest fragments in the San Martín region of Peru. Using instantaneous focal sampling of infants, I recorded physical conflict between infants and caregivers and instances of infant avoidance (leaving the infant) by adult males. I summarized data as the percentage of records in which these activities occurred for each focal day and report the estimates for caregivers. I further calculated mean percentages by month of infant age to assess the relative timing of infant conflict, for each group and age/sex class, and infant avoidance by males. Percentages of conflict and avoidance were markedly higher in the larger group living in a smaller habitat than in the other group. This pattern occurred across all age/sex classes. In both groups, the greatest amount of infant conflict occurred with siblings. I discuss the substantial variation in conflict and avoidance in relation to the various socioecological conditions that may have played a role. This study provides an in-depth description and exploration of parent-offspring and sibling conflict, which has not been examined previously in this species.

Effects of aggressive temperament on endogenous oxytocin levels in adult titi monkeys.

Coordination of oxytocin (OT) activity and partner interactions is important for the facilitation and maintenance of monogamous pair bonds. We used coppery titi monkeys (Callicebus cupreus) to identify the effects of male aggressive temperament on OT activity, affiliative partner-directed behaviors, aggressive partner-directed behaviors, anxiety-related behaviors, and hormone-behavior interactions. We used a mirror technique, simulating an intruder in the home territory of pairs to elicit behavioral responses, and quantified behaviors using an established ethogram. Plasma concentrations of OT (pg/ml) were quantified using enzyme immunoassay. We used general linear mixed models to predict 1) percent change in OT as a function of aggression score, and 2) percent change in behaviors as a function of aggression, OT, and OT by aggression interactions. High-aggressive males exhibited a significant drop in OT concentration relative to control when exposed to the front of the mirror (ß = -0.22, SE = 0.10, t = -2.20, p = 0.04). High-aggressive males spent significantly less time in contact with their mates (ß = -1.35, SE = 0.60, t = -2.26, p = 0.04) and lip-smacked less (ß = -1.02, SE = 0.44, t = -2.32, p = 0.03) relative to control. We also saw a trend toward an interaction effect between OT and proximity such that High-aggressive males displaying a drop in OT exhibited a smaller percent increase in social proximity (ß = 6.80, SE = 3.48, t = 1.96, p = 0.07). Males exhibiting a decrease in OT also trended toward back-arching and tail-lashing less in response to the mirror (ß = 4.53, SE = 2.5, t = 1.82, p = 0.09). To our knowledge, this is the first empirical study to examine interactions between OT and temperament in adult monogamous primates. Future studies should incorporate measures of pair-mate interactions and early-life experience to further understand variation in responses to social stressors and their effects on pair bonding.

Effect of reward type on object discrimination learning in socially monogamous coppery titi monkeys (Callicebus cupreus).

Highly valued food items are often used as rewards to reinforce an animal's behavior. For social species, social interaction is rewarding and can drive an individual's behavior as well. In the currently study, we wanted to compare the efficacy of a food reward and a social reward on object discrimination learning in socially monogamous titi monkeys. We hypothesized that titi monkeys would perform more accurately for a social reward (their pair mate) than for a food reward (a highly desired food item). Eleven adult titi monkeys were tested with a two-object visual discrimination task for both types of reward. The colors and shapes of the objects in the two-object discrimination task were counterbalanced across subjects. During each trial, subjects were shown two objects, and the trial ended when the subject touched the reinforced shape (S+) or after 5 min. A correct trial was defined as one when the subject touched S+ first. We found that 45.5% of subjects were able to learn the task with a social reward, and 83.3% were able to learn the task with a food reward. We found that subjects balked more often and had fewer correct trials for the social reward. Finally, subjects took longer to approach the shapes for a social reward, possibly indicating lower motivation to engage in the task when a social reward is used compared to a food reward. Although significantly fewer subjects met criteria of success with the social reward than with the food reward, our results show that titi monkeys can learn a visual discrimination task with either type of reward.

Delayed Dispersal and Immigration in Equatorial Sakis (Pithecia aequatorialis): Factors in the Transition from Pair- to Group-Living.

Saki monkeys (Pithecia spp.) live in pairs and small groups, sometimes with more than 1 same-sex adult. Previous studies have not been able to distinguish additional, unrelated adults from adult-sized offspring, but both can influence social relationships and mating strategies, albeit in different ways. In this study, we documented the immigration of an adult male equatorial saki (P. aequatorialis) into a group following the departure of the previous resident male. At immigration, the group contained an adult female, her 5-year-old (adult age) and 1.5-year-old daughters, and her 1-month-old infant. We used nearest neighbor, approach, grooming, playing, aggression, and copulation data to describe the social dynamics between the immigrant male and the 2 adult females. In the 12 months following his arrival, the immigrant male tended to be closer to and groom the adult daughter more than the mother, but he mated with both females. Both females interacted more with the immigrant male than with each other, and both females eventually reproduced. These observations provide evidence that in equatorial sakis, adult offspring may delay dispersal and reproduce within their natal group, thus transitioning from groups of reproductive pairs to groups with more than 1 reproductive adult of the same sex.

Ecological and Behavioural Factors Influencing Territorial Call Rates for the Bolivian Titi Monkeys, Plecturocebus modestus and Plecturocebus olallae.

Primate territorial calls have been used to locate groups in censuses. Daily variations in call rates are a potential source of error in these studies. To obtain more accurate estimations of population density it is necessary to determine how much variation there is in group call rates and to identify the factors that influence them. We present data on the emission of territorial calls by two threatened and endemic titi monkeys in Bolivia: Plecturocebus olallae and P. modestus. We found interspecific differences in daily call rates (52% P. modestus and 33% P. olallae). Groups inhabiting more continuous forests vocalized more frequently than groups in fragmented forests, which might be linked to the higher abundance of groups in less fragmented forests. We found seasonal differences in call rates between species, with more frequent calling in P. modestus during the dry season, while P. olallae called more frequently in the rainy season. The study groups emitted territorial calls mostly from the central zones of their territories, suggesting they do not face intense spatial competition with neighbouring groups. Our results improve the general ecological knowledge on P. olallae and P. modestus, and can also be used to improve population abundance studies and ongoing conservation efforts.

Serotonin 1A agonism decreases affiliative behavior in pair-bonded titi monkeys.

Relatively little is known about serotonergic involvement in pair-bonding despite its putative role in regulating social behavior. Here we sought to determine if pharmacological elevation of serotonin 1A (5-HT1A) receptor activity would lead to changes in social behavior in pair-bonded male titi monkeys (Callicebus cupreus). Adult males in established heterosexual pairs were injected daily with the selective 5-HT1A agonist 8-OH-DPAT or saline for 15days using a within-subjects design. Social behavior with the female pair-mate was quantified, and plasma concentrations of oxytocin, vasopressin, and cortisol were measured. When treated with saline, subjects showed reduced plasma oxytocin concentrations, while 8-OH-DPAT treatment buffered this decrease. Treatment with 8-OH-DPAT also led to decreased plasma cortisol 15minutes post-injection and decreased social behavior directed toward the pair-mate including approaching, initiating contact, lipsmacking, and grooming. The reduction in affiliative behavior seen with increased activity at 5-HT1A receptors indicates a substantial role of serotonin activity in the expression of social behavior. In addition, results indicate that the effects of 5-HT1A agonism on social behavior in adulthood differ between rodents and primates.

Determinants of spatial behavior of a tropical forest seed predator: The roles of optimal foraging, dietary diversification, and home range defense.

Specialized seed predators in tropical forests may avoid seasonal food scarcity and interspecific feeding competition but may need to diversify their daily diet to limit ingestion of any given toxin. Seed predators may, therefore, adopt foraging strategies that favor dietary diversity and resource monitoring, rather than efficient energy intake, as suggested by optimal foraging theory. We tested whether fine-scale space use by a small-group-living seed predator-the bald-faced saki monkey (Pithecia irrorata)-reflected optimization of short-term foraging efficiency, maximization of daily dietary diversity, and/or responses to the threat of territorial encroachment by neighboring groups. Food patches across home ranges of five adjacent saki groups were widely spread, but areas with higher densities of stems or food species were not allocated greater feeding time. Foraging patterns-specifically, relatively long daily travel paths that bypassed available fruiting trees and relatively short feeding bouts in undepleted food patches-suggest a strategy that maximizes dietary diversification, rather than "optimal" foraging. Travel distance was unrelated to the proportion of seeds in the diet. Moreover, while taxonomically diverse, the daily diets of our study groups were no more species-rich than randomly derived diets based on co-occurring available food species. Sakis preferentially used overlapping areas of their HRs, within which adjacent groups shared many food trees, yet the density of food plants or food species in these areas was no greater than in other HR areas. The high likelihood of depletion by neighboring groups of otherwise enduring food sources may encourage monitoring of peripheral food patches in overlap areas, even if at the expense of immediate energy intake, suggesting that between-group competition is a key driver of fine-scale home range use in sakis.

A Saki Saga: Dynamic and Disruptive Relationships among Pithecia aequatorialis in Ecuador.

Saki monkeys live in socially monogamous groups and in groups containing more than one same-sex adult. As part of a 10-year study of equatorial sakis (Pithecia aequatorialis) in Ecuador, we documented the immigration of a second adult male into a group containing a resident male-female pair that had associated with one another for seven years and the resident female's two daughters. In the first month after immigration, the resident male spent more time closer to and grooming his putative adult daughter than the resident female, and the two males were seen performing a cooperative territorial display. After two months, the resident male interacted more with the resident female than with his putative adult daughter, while that daughter interacted more with the immigrant male and copulated with him. After three months, the males left the group together and associated with an unfamiliar female, leaving the resident females and a neonate behind. The resident male then paired with a new female, while the immigrant male joined another group, again as a second male. Compared to other socially monogamous primates, sakis appear to have a more variable social system whereby additional males can join established groups and form relationships with putatively unrelated males.

Spatial foraging in free ranging bearded sakis: traveling salesmen or Lévy Walkers?

According to optimal foraging theory and most current models of primate socioecology, primate foraging involves a series of decisions concerning when is the most optimal time to leave a food patch, how to travel to the next patch in an efficient manner, and how to minimize the time and distance traveled to all patches throughout the course of the day. In this study, I assess how bearded sakis solve these challenges by presenting data on their patch use, distance minimization, and by comparing their movements with non-deterministic foraging patterns. The study group, composed of 38 ± 15 individuals, fed significantly longer in higher quality patches (quality defined by patch size and productivity) and in those that contained ripe fruit pulp. However, group size was not a significant predictor of patch occupancy. Bearded sakis traveled relatively directly between food patches, sometimes over distances > 300 m. In addition, they chose the optimal daily path among all patches visited on 9 of 17 occasions, and on average traveled only 21% more than the least distance route. Bearded saki step lengths were consistent with a Brownian rather than a Lévy Walk pattern while waiting times were consistent with a Lévy pattern. However, the distribution of their turning angles indicated a high degree of directional persistence between patches. These results suggest that bearded sakis exploit food patches that are randomly distributed spatially but heterogenous in patch quality. They appear to encode the locations of high quality food patches and minimize travel between them, despite opportunistically feeding from more abundant and randomly distributed, lower quality patches en route.

Differences in titi monkey (Callicebus cupreus) social bonds affect arousal, affiliation, and response to reward.

Titi monkeys (Callicebus cupreus) are a monogamous, New World primate. Adult pair-mates form a bidirectional social bond and offspring form a selective unidirectional bond to their father. Some of the neurobiology involved in social bonds and maternal behavior is similar to the neural circuitry involved in nonsocial reward. Due to these overlapping mechanisms, social states may affect responses to external rewarding stimuli. We sought to determine whether having a social attachment, and/or being in the presence of that attachment figure, can affect an individual's response to a rewarding stimulus. In addition, we compared affiliative bonds between pair-mates to those between offspring and fathers. Eighteen adult male titi monkeys were either living alone (Lone), with a female pair-mate (Paired), or with the natal group (Natal; N = 6/condition). Each individual went through eight 30-min preference tests for a sweet substance, Tang. For Paired and Natal males, half of the test sessions were with their attachment figure and half were alone. Lone males were always tested alone. Preference scores for Tang, time spent drinking, affiliative, and arousal behaviors were measured. Paired and Natal males emitted significantly more isolation peeps and locomoted more when tested alone compared to when tested with their attachment figure, and paired males engaged in more affiliative behavior than Natal males. Lone males engaged in significantly more behaviors indicative of behavioral arousal such as locomotion and piloerection compared to Paired and Natal males. Finally, Paired males drank significantly more Tang and had a significantly greater preference for Tang compared to Lone and Natal males. These results indicate that offspring undergo a behavioral separation response upon separation from their father that persists into adulthood, Lone males are more behaviorally reactive, and that living with an attachment figure and the type of attachment relationship result in different responses to a rewarding sweet stimulus.

Sleeping site selection by golden-backed uacaris, Cacajao melanocephalus ouakary (Pitheciidae), in Amazonian flooded forests.

In Amazonian seasonally flooded forest (igapó), golden-backed uacaris, Cacajao melanocephalus ouakary, show high selectivity for sleeping trees. Of 89 tree species in igapó, only 16 were used for sleeping (18%). Hydrochorea marginata (Fabaceae) and Ormosia paraensis (Fabaceae) were used most frequently (41% of records) despite being uncommon (Ivlev electivity ratios were 0.76, and 0.84, respectively), though the third most commonly used species (11%), Amanoa oblongifolia (Euphorbiaceae), was selected at near parity. All three species have broad, open canopies with large horizontal limbs and uncluttered interiors. Compared with random trees, sleeping trees had above average diameter at breast height (DBH) and height, lacked lianas and wasp nests, and were more frequently within 5 m of open water. Uacaris generally slept one adult per tree or widely separated in the same canopy and on the outer third of the branch. These behaviours are interpreted as maximising detection of both aerial and arboreal predators.

Seasonal versatility in the feeding ecology of a group of titis (Callicebus coimbrai) in the northern Brazilian Atlantic Forest.

The feeding behavior of a group of titis (Callicebus coimbrai) was monitored over an annual cycle at a site in northeastern Brazil. Behavioral data were collected in scan samples (1-min scan at 5-min intervals), and complementary data on fruit availability and new leaf cover were collected. Feeding time accounted for 28.9% of daily activity. Fruit was the principal item of the diet (61.2% of records) and the primary category in all months except September, when it was surpassed by leaves. Young leaves were the second most important category (20.0%). The consumption of seeds and insects was prominent in November and December. Fifty-two plant species were exploited, and the Elaeocarpaceae, Myrtaceae, Sapotaceae, and Passifloraceae provided the vast majority (86.0%) of plant feeding records. The phenological record did not provide a good measure of fruit availability, but a strong correlation (r(s) =0.902, P<0.0001, n=12) was found between the consumption of leaves and the exploitation of lianas each month. Lianas accounted for 28.2% of plant feeding records, and predominated between August and December. This suggests that lianas may represent a key factor in the ability of the species to tolerate the intense habitat fragmentation found throughout its geographic range.

Within-group social bonds in white-faced saki monkeys (Pithecia pithecia) display male-female pair preference.

White-faced saki monkeys (Pithecia pithecia) lack most of the behavioral and physical traits typical of primate monogamy [Fuentes, 1999]. In order to determine if social bonds in this species reflect patterns displayed by pair-bonded groups or larger multimale-multifemale groups, we draw on 17 months of data collected on wild white-faced sakis at Brownsberg Nature Park, Suriname. We analyzed within-group social bonds for three habituated groups (one two-adult and two multiadult groups) by measuring grooming, proximity, and approach/leave patterns between adult and subadult group members. We found that both two-adult and multiadult groups showed significantly stronger social bonds between a single male-female dyad within each group (deemed "primary dyads"). In all three groups, primary dyads were composed of the oldest adult male and a breeding female. These pairs had significantly higher levels of grooming than other within-group dyads and were also in close proximity (<1 m) more often than nonprimary dyads. Grooming in primary dyads was nonreciprocal, and consistently biased toward female investment. Grooming patterns in nonprimary dyads varied, but were often more reciprocal. Grooming and proximity of the primary dyad also changed in relation to infant development. Our results suggest that while white-faced sakis do not show behavioral and physical traits typical of monogamy or pair-bonding, social bonds are strongest between a single male-female pair. Pitheciine social systems range from small group monogamy in Callicebus to large multimale-multifemale groups in Chiropotes and Cacajao. As the middle taxon in this platyrrhine radiation, behavioral strategies of white-faced sakis provide a model for how social bonds and affiliation could be influenced by and affect the evolution of larger group size in primates.

Use of sleeping sites by a titi group (Callicebus coimbrai) in the Brazilian Atlantic Forest.

Predation pressure affects most aspects of primate behaviour, and is especially pronounced in the context of the use of sleeping sites, given the vulnerability of the animal at this time. Most small-bodied platyrrhines have highly systematic patterns of sleeping site choice and use. This study analyses the use of sleeping sites by a free-ranging group of titis (Callicebus coimbrai) monitored at a site in Sergipe, Brazil, between July, 2009 and June, 2010. When the subjects approached a sleeping tree their behaviour was typically cautious, including slow and silent movement, early retirement (20-162 min before sunset on 52 dry afternoons), and sleeping in a tight huddle with their tails entwined. Despite this behaviour, which has an obvious anti-predator function, the group slept in only three different trees during the course of the study, and returned to the same tree used on the previous night on a quarter of evenings (n = 56). This was despite the availability within the group's home range of a large number of trees with similar structural characteristics (i.e. tall, open crown in the upper canopy). Surprisingly, the three trees were all members of the same species, Licania littoralis (Chrysobalanaceae). The choice of this species, which was not an important source of dietary resources, and the repeated use of a small number of sites, did not seem to be related to factors such as ranging or foraging patterns, but may have a been a response to the specific threat from capuchins, Cebus xanthosternos.

Predation and predation attempts on red titi monkeys (Callicebus discolor) and equatorial sakis (Pithecia aequatorialis) in Amazonian Ecuador.

Anecdotal reports of predation as well as observed predation attempts and rates of animal disappearance provide some of the most relevant data for evaluating the influence that predation risk may have on primate behavioural ecology. Here, we report rates of disappearance from six groups of red titi monkeys (Callicebus discolor) and two groups of equatorial sakis (Pithecia aequatorialis) followed over a period of four and a half years at a lowland site in Amazonian Ecuador. We also describe the first direct observation of a harpy eagle (Harpia harpyja) preying upon a titi monkey, as well as 3 unsuccessful attacks by tayras (Eira barbara) on titi monkeys and 4 unsuccessful attacks by various raptors on sakis. Our data indicate that pitheciid primates may face a wider array of possible predators than previously recognized, and that titi monkeys and sakis are susceptible to different major classes of predators. Our observations also suggest differences in the sex role during predator defence that could be related to the evolution and maintenance of monogamous systems.

Relationship between age and activity varies by sex in titi monkeys (Callicebus cupreus).

Motor activity increases during development and typically peaks before reproductive maturity. Males may become more active than do females, and this sex difference in activity may be a mechanism contributing to sex-biased dispersal. The generality of the link between sex differences in activity and sex differences in dispersal would be strengthened with evidence of greater female, as compared to male, activity during development in a species with a female bias in dispersal. Titi monkeys are a good model for such an investigation because subadult females leave their natal group at an earlier age than do subadult males. This study examines whether developing female titi monkeys become more active with age than do developing male titi monkeys. Locomotor activity of 13 female and 8 male captive titi monkeys was measured as they matured from initial ambulatory independence (6 months of age) to reproductive maturity (24 months). Increased activity of females was associated most strongly with increasing age. Increased activity of males, however, was associated most strongly with repeated testing. Compared to maturing males, greater activity of maturing females may underlie the earlier dispersal of female titi monkeys that has been reported.

Behavioral effects of introducing pied tamarin (Saguinus bicolor) to black howler monkey (Alouatta caraya) and white-faced saki (Pithecia pithecia) in a zoological park.

Mixed-species primate exhibits are becoming more common in zoological parks as a means to display a diverse array of animals both more naturalistically and with more economy of space. Here, we describe behavioral changes during the introduction process of a pair of pied tamarins (Saguinus bicolor) to an established group of black howler monkeys (Allouatta caraya) and white-faced saki monkeys (Pithecia pithecia). Data were collected during six phases, representing introductions among the various species and to exhibit space and off-exhibit holding. The pied tamarins were consistently the most active of the three species. Although activity levels of the howler and saki monkeys remained constant throughout, that of the tamarins declined as the introduction progressed. Several episodes of aggression between the tamarins and the sakis were observed, but did not coincide with patterns predicted by previous intra-specific introductions. The three-species mix remained stable for several months; however, escalating aggression ultimately led to the removal of the sakis from the mixed-species exhibit. Despite our mixed results, we contend that only through continued trials, coupled with careful and systematic monitoring, can we ultimately identify stable mixes of species.