Comparative chromosome morphology in three Callitrichid genera: Cebuella, Callithrix, and Leontopithecus

A G-band karyotypic analysis was carried out in individual species groups of three Callitrichid primate genera: Cebuella, Callithrix, and Leontopithecus. Within Callithrix, the karyotypes of the morphologically distinct and geographically isolated morphotypes C. jacchus jacchus and C. jacchus penicillata were identical. Within the lion tamarin genus, Leontopithecus, the karyotypes of the three morphotypes (L. rosalia rosalia, L. rosalia chrysomelas and L. rosalia chrysopygus) were also indistinguishable from one another. These results are consistent with the taxonomic designation of subspecies rank to the different morphotypes. A comparison of type specimens among the three Callitrichid genera showed that their phyletic radiation has been paralleled by a limited number of chromosome rearrangements and a relatively high amount of karyotypic invariance. A fusion/fission event has been postulated to account for the difference in diploid number between Cebuella (2n = 44) and the other species (2n = 46). The karyotype of Callithrix jacchus was found to be more directly derived from Cebuella than was that of Leontopithecus. These findings differ from the previous proposition that Leontopithecus might have diverged from a common Callitrichid ancestor before the emergence of the genus Callithrix.     

Stereology of the myocardium in Leontopithecus (Lesson, 1840) callitrichidae - primates

Rare morphological features of the Leontopithecus cardiovascular system have been reported in the literature. The samples analyzed in this study came from 33 specimens of Leontopithecus from the collection of the Center of Primatology of Rio de Janeiro-FEEMA (CPRJ-FEEMA). Morphometry and stereological data were obtained from all animals. Adult body weights of L. rosalia were the lowest, the greatest being those of L. chrysopygus caissara; body weights of L. chrysomelas and L. c. chrysopygus were similar and in between those of the two former species. Cardiomyocytes (left ventricular myocardium) were bigger in adults than in infants. The myocardium of L. rosalia showed focal fibrosis, fatty vacuoles, and hyalinization. In L. chrysomelas the myocardium showed areas of fibrosis and presence of mononuclear cells. Fibrosis and areas of congestion were observed in L. c. chrysopygus; areas of disorganization and vascular congestion were found in L. c. caissara. In L. rosalia infants, a greater density of vessels per myocardial area and a greater length density of vessels were observed as compared with those of L. chrysomelas. In adults, L. chrysomelas showed greater density of connective tissue in the myocardium than L. c. chrysopygus and L. c. caissara did. In L. rosalia, cardiomyocyte nuclei had a greater area density than those of the other forms of Leontopithecus. These characteristics may explain the faster development of L. rosalia infants as compared with that of L. chrysomelas and L. c. chrysopygus kept under the same handling conditions at the CPRJ-FEEMA.     

Morphological and morphometric age-related changes of the upper thoracic aorta in Leontopithecus (Lesson, 1840) (callitrichidae--primates)

Few morphologic aspects of Leontopithecus are listed in the literature, which does not provide sufficient knowledge of those simians. However, such results are important to evaluate the potential of those species as a model for studying comparative pathology. In this study, the material came from the collection of the Museu de Primatologia do Centro de Primatologia do Rio de Janeiro, fixed in 10% formalin pH 7.2. The samples were 33 individuals (five infant and five adult L. rosalia, L. chrysomelas, L. chrysopygus chrysopygus, and three adult L. chrysopygus caissara). The sections of the thoracic aorta were prepared for histological study and measurements were obtained. In the infant Leontopithecus, the aorta presents endothelial line and subendothelial space, the tunica media is not comparatively very thick. In the adults, the aorta showed enlarged tunica intima with consequent thick arterial wall, which increased with the aging process. The atheromatous plates were seen in the aortic wall of all adult L. chrysopygus chrysopygus, in two adult L. rosalia and only one adult L. chrysomelas. Differences in the aortic internal and external diameters were not significant in infants. In the adults, no significant difference was found among L. rosalia, L. chrysomelas and L. c. caissara, although the average of the aortic internal diameter in L. chrysomelas was larger than in L. c. caissara. In infant and adults L. c. chrysopygus, the thickness of the thoracic aorta is larger than in the other forms of Leontopithecus. These findings suggest a morphometric and morphologic aortic difference among the species of Leontopithecus, which could have biological significance that needs major investigation in the future.     

Ontogenetic characteristics of the vomeronasal organ in Saguinus geoffroyi and Leontopithecus rosalia,with comparisons to other primates

It has been suggested that the variability of the primate vomeronasal organ (VNO) may be greater than previously thought, especially among New World monkeys. It is not clear to what extent VNO variation reflects ontogenetic, functional, or phylogenetic differences among primates. The present study investigated VNO anatomy in an ontogenetic series of two genera of callitrichid primates, in order to assess recent attempts to develop VNO character states and to examine the evidence for VNO functionality at different life stages. A sample of six Leontopithecus rosalia, one L. chrysomelas, and six Saguinus geoffroyi was serially sectioned and stained using various methods. Two adult Callithrix jacchus were also sectioned for comparative purposes. The VNO of each primate was examined by light microscopy along its entire rostrocaudal extent. VNOs of the tamarins were described to determine whether they fit into 1 of 3 character states recently attributed to various New World monkeys. At birth, the two species of tamarins differed in the nature of communication between the VNO and nasopalatine duct (NPD). Two of 3 neonatal S. geoffroyi exhibited a fused VNO duct in a more dorsal position (adjacent to the nasal cavity) compared to that of L. rosalia. The VNO duct communicated with the NPD and was patent in neonatal L. rosalia. Both species appeared to have an age-related increase in the amount of sensory epithelium in the VNO. Subadult L. rosalia had caudal regions of the VNO that were exceptionally well-developed, similar to those of strepsirhine primates. Compared to subadults, all adult callitrichids appeared to have more ventral communications of the VNO duct directly into the NPD. Adult S. geoffroyi and L. chrysomelas both had VNO sensory epithelium separated by multiple patches of nonsensory epithelium. This contrasted with the VNOs of C. jacchus, which had a nearly continuous distribution of receptors on all surfaces of the VNO. The findings indicate that tamarins have delayed maturation of the VNO epithelium, and that some species have little or no perinatal function. These results also suggest that ontogenetic changes in craniofacial form may alter the position of the VNO in tamarins. The present study supports the use of at least two character states to categorize the VNO of various callitrichids, but it is suggested that one of these, previously called "reduced sensory epithelium" should be instead termed "interrupted sensory epithelium." The distribution of VNO sensory epithelium does not appear to reflect phylogenetic influences; it is more likely a functional characteristic that varies throughout postnatal life. Therefore, this chemosensory system has a high degree of plasticity relating to age and function, which in some instances can confound the use of characteristics as phylogenetic traits. Further study is needed to quantify VNO receptors in various species to determine if functional differences exist and if some species have more precocious VNO function than others.     

Circadian and Ultradian Time Patterns in Human Behavior: Part 2: Social Synchronisation During the Development of the Infant's Diurnal Activity-Rest Pattern

Individually developing patterns of activity-rest rhythms in infants and the influence of environmental factors in the tuning and synchronisation of parent-infant pairs have important implications for the health of both infant and parents. After discharge from the hospital newborn infants are exposed to patterned influences of maternal and environmental regularities of a family's daily life resulting in varying degrees of social synchronisation. Actigraphic monitoring was used in this longitudinal study to examine how activity patterns of the entire family agree or disagree with each other, and how the infant entrains to the environment. Activity data of 12 families (father, mother and infant) were continuously recorded using non-invasive Actiwatch units. Recordings of parental activity started at the beginning of the 37th week of gestation, and were continued in parallel with the infants' recordings in three series of 21 days each until four months after birth: 1st to 3rd week, 7th to 9th week and 13th to 15th week of life. Fast Fourier transformation and cross correlation techniques were used to determine frequencies of each family member and to quantify the synchronisation of activity between parents and infants. To elucidate differences in social synchronisation between human cultures, synchronisation of a Melanesian family was additionally compared. Results showed the existence of corresponding ultradian frequencies in the activity patterns of mother-infant pairs at 1, 2 and 4 months. Increases in the synchronisation of parental activity were found from prenatal to postnatal and for mother-infant pairs from the first to the second month. Synchronisation between mother and infant always exceeded that of father and infant. Transient mono-, bi- or polyphasic activity patterns emerged in the infants immediately after birth. Good correspondence of mother-infant activity patterns during the early postnatal period was correlated with a rapid development of an entrained daily pattern in the infant.     

Interactions between parents and infants,and infants-father separation in the common marmoset (Callithrix jacchus)

The New World monkey,Callithrix jacchus, diversely from most Old World primates, shares infant care among all family members. This paper intends to describe a pattern of infant care in the common marmoset, and to verify if the infant-father separation induces a depression reaction, such as found in other separation studies. In Experiment 1, 11 families (mother, father, and twin newborn infants) ofCallithrix jacchus were observed from 1st to 36th days of infants' life. It was noted that infants were carried most of the time in the beginning of life, and that from the 4th day on this care came mostly from the father. In Experiment 2, nine families were divided in three separation groups (15, 30, and 45 days of infants' life). Each group had a control and two experimental families. Removal of the father was done for three days, beginning at the indicated infants' ages. Results showed that upon removal of the father, the mother assumes the infants' care, and no depression reaction appeared.     

Attachment and social preferences in cooperatively-reared cotton-top tamarins

In many primate species a close attachment between mother and infant provides a secure base for the infant when the infant is frightened or under stress. In cooperatively breeding primates infant carrying is divided among several individuals in the group, with the mother often doing little more than nursing. In these species it is not clear which individual would best serve as a secure base for the infant. We studied eight infant cotton-top tamarins from birth through 20 weeks of age, noting who carried the infant during the first 100 days, who transferred food with the infants, and, as infants became independent, with whom they associated during social play and affiliative behavior. From week 9 to week 20, when infants were independent of carriers most of the time, we presented families with six trials (once every 2 weeks) with a threatening stimulus (a human dressed in a lab coat and wearing an animal mask). Infants played primarily with their twin or youngest sibling and had affiliative interactions with many family members. However, in fearful situations, infants ran to those who had carried them and transferred food with them the most-their father or oldest brother (never to the mother). Although adults increased rates of mobbing calls in response to the threat, infants significantly reduced their vocalization rate. For these cooperatively breeding monkeys, the attachment object for infants is the family member that invested the most effort in carrying the infant and transferring food with the infant. These results parallel and extend results from bi-parental infant care species in which the father serves as the primary attachment figure.     

Exposure to infant scent lowers serum testosterone in father common marmosets (Callithrix jacchus)

Common marmoset (Callithrix jacchus) males are bi-parental non-human primates that show extensive paternal behaviour. Fathers are in direct sensory contact with their infants during the natal period. We found that fathers exposed to isolated scents of their infant displayed a significant drop in serum testosterone levels within 20min after exposure, whereas parentally naive males did not. These data suggest that infant's scent may have a causal role in regulating paternal testosterone in their fathers. This is the first study to demonstrate that olfactory cues have an acute effect on paternal care.     

Phylogeny of lion tamarins (Leontopithecus spp) based on interphotoreceptor retinol binding protein intron sequences

The evolutionary relationships of the lion tamarins (Leontopithecus) were investigated using nuclear interphotoreceptor retinol binding protein (IRBP) intron sequences. Phylogenetic reconstructions strongly support the monophyly of the genus, and a sister relationship between the golden lion tamarin, Leontopithecus rosalia, and the black lion tamarin, L. chrysopygus, to the exclusion of the golden-headed lion tamarin, L. chrysomelas. The most parsimonious evolutionary reconstruction suggests that the ancestral lion tamarin and the common ancestor of L. rosalia and L. chrysopygus had predominantly black coats. This reconstruction is not consistent with a theory of orthogenetic evolution of coat color that was based on coat color evolution in marmosets and tamarins. An alternative reconstruction that is consistent with metachromism requires that ancestral lion tamarins had agouti hairs.     

The Role of Non-reproductive Helpers in Infant Care in Captive Callithrix jacchus

Callitrichids have been proposed as communal breeders, with non-reproductive animals acting as helpers. Reports in the literature attest to the high variability in the behaviour of helpers. The present study assessed four proximate factors involved in helping in captive Callithrix jacchus: age, gender and experience of helpers and composition of their groups. Forty-eight helpers from 15 families from Natal (Brazil) and Reading (UK) colonies were observed during the first 4 weeks of life of new-born twins. Infant carrying, transfers and retrieval, and duration of carrying bouts were registered. Results showed that the presence of helpers decreased the costs of infant carrying, with a negatively significant correlation between group size and mean percentage of carrying. Age, but not gender, was a significant factor for infant carrying, transfers and retrieval, and mean duration of carrying bouts, with older helpers contributing more than younger ones. Singletons and oldest non-reproductive offspring carried infants more, and longer, than twins and offspring with older siblings present in the group. Experience decreased motivation and actual carrying time, but not to a significant level. Age, experience and group composition appeared important factors in the display of infant care by helpers in captive Callithrix jacchus.     

Genetic variation within and among lion tamarins

The golden lion tamarin Leontopithecus rosalia rosalia, one of the rarest and most endangered of New World primates, has been the focus of an intensive research and conservation effort for two decades. During that period, managed breeding from 44 founders has brought the captive population to over 400 individuals, a number that equals or exceeds the estimated number of free-ranging golden lion tamarins. The extent of genetic variation among golden lion tamarins was estimated with an electrophoretic survey of 47 allozyme loci from 67 captive and 73 free-ranging individuals. The amount of variation was low, compared to 15 other primate species, with 4% of the loci being polymorphic (P), and with an average heterozygosity H estimate of 0.01 in these callitrichids. Electrophoretic analyses of captive and free-ranging animals (N = 31) of two allopatric morphotypes, Leontopithecus rosalia chrysopygus and L. r. chrysomelas, were similar to the L. r. rosalia findings insofar as they also revealed limited genetic polymorphism. Computation of the Nei-genetic distance measurements showed that the three morphotypes were genetically very similar, although discernible differentiation had occurred at two loci. These data are consistent with the occurrence of recent reproductive isolations of these subspecies.     

Longitudinal observations of care and development of infant titi monkeys (Callicebus moloch)

The purpose of this study was to describe patterns of infant care and development in family groups of the monogamous titi (Callicebus moloch). Three infant titis were observed with their natal family groups over the first six months of life. Field observations of extensive male involvement with infants were confirmed. Adult males were clearly the infants' primary social companions, and infants spent more time in contact with adult males at all ages than with mothers or older siblings. However, mothers and siblings also carried infants at times and interacted with them in affiliative ways (e.g., grooming, nuzzling, play). Mothers often invited nursing, but otherwise it was infants who primarily initiated transfers between carriers and approaches to parents, reminding us that the infants' own activities and their effects on caregivers should not be overlooked in considerations of the patterns of infant care. This is particularly important for those species exhibiting extensive parental involvement by males. It is suggested that substantial male care of young titi infants leads to several important consequences for the infant's social development, including the development of a stronger attachment to the father than the mother.     

Diet, foraging, and use of space in wild golden-headed lion tamarins

Lion tamarins (Callitrichidae: Leontopithecus) are small frugi-faunivores that defend large home ranges. We describe results from the first long-term investigation of wild golden-headed lion tamarins (L. chrysomelas; GHLTs). We present data about activity budgets, daily activity cycles, diet, daily path length, home range size, home range overlap, and territorial encounters for three groups of GHLTs that were studied for 1.5-2.5 years in Una Biological Reserve, Bahia State, Brazil, an area characterized by aseasonal rainfall. We compare our results to those from other studies of lion tamarins to identify factors that may influence foraging and ranging patterns in this genus. Ripe fruit, nectar, insects, and small vertebrates were the primary components of the GHLT diet, and gums were rarely eaten. Fruit comprised the majority of plant feeding bouts, and the GHLTs ate at least 79 different species of plants from 32 families. The most common foraging sites for animal prey were epiphytic bromeliads. The GHLTs defended large home ranges averaging 123 ha, but showed strong affinities for core areas, spending 50% of their time in approximately 11% of their home range. Encounters with neighboring groups averaged two encounters every 9 days, and they were always aggressive. Data about time budgets and daily activity cycles reveal that the GHLTs spent most of their time foraging for resources or traveling between foraging sites distributed throughout their home ranges. The GHLTs spent much less time consuming exudates compared to lion tamarins in more seasonal environments. Additionally, the GHLTs had much larger home ranges than golden lion tamarins (L. rosalia), and did not engage in territorial encounters as frequently as L. rosalia. GHLT ranging patterns appear to be strongly influenced by resource acquisition and, to a lesser extent, by resource defense.     

Early mother-infant relationships ofMacaca thibetana at Mt. Emei China

Quantitative data on early mother-infant relationships in the Tibetan macaque was collected during the first 23 weeks of infant life in spring, 1987, at Mt. Emei, China. During the first week of life, infants spent 98.3% of their time in ventroventral contact with their mothers. This contact rapidly decreased to 33.8% by the 4th week and thereafter to 0.85% by the 23rd week. Nipple contact decreased relatively slowly from 89.7% to 62.9% within the first 4 weeks of infant life and to 19.8% by the 23rd week. Ventrolateral and ventrodorsal contact appeared by the 2nd week, meanwhile, maternal restraining behavior appeared, and reached a peak by the 3rd week. The mother neither encouraged nor discouraged her infant's independence during 4–8th weeks. Maternal rejection of the infant was first observed when the infant was 11 weeks old and continued thereafter.     

Maternal care and infant development in <Emphasis Type="Italic">Callimico goeldii</Emphasis> and <Emphasis Type="Italic">Callithrix jacchus</Emphasis>

Callimico goeldii gives birth to single offspring, whereas other callitrichids, including Callithrix jacchus, twin. This study compares maternal effort and infant development in C. goeldii and C. jacchus; it is the first study to look at nursing frequency. Infants were observed from birth for 7 weeks in two captive groups each of C. goeldii and C. jacchus. C. goeldii mothers physiologically invested the same or less than C. jacchus mothers. C. goeldii mothers gained the same amount of weight during pregnancy in absolute terms as did the smaller C. jacchus. This results in a smaller gain in proportion to maternal weight but an equivalent proportional gain on a per fetus basis. C. goeldii mothers nursed their infants less based on duration of nursing bouts compared with C. jacchus mothers. C. goeldii mothers transported their infants exclusively through the first 2 weeks of life, which is longer than C. jacchus mothers, who exclusively transported infants only during the first week of life. As maternal infant carriage declined, other group members transported offspring in both species. C. goeldii infants engaged in independent locomotive sequences later in development and tasted solid foods less frequently than C. jacchus infants when compared at equivalent ages. A single, opportunistic milk sample obtained from a C. goeldii mother when her infant was 48 days old indicates that C. goeldii milk contains gross energy from crude protein within the range of variation observed in Callithrix milk. Despite the similarities in milk quality and prenatal effort in individual fetuses, C. goeldii infants gain weight faster from 0 to 18 months than do C. jacchus infants. A reduction in litter size allows C. goeldii mothers to spend more time carrying their infant and to delay weaning, thereby allowing accelerated infant and juvenile growth rates compared with C. jacchus.     

Observations on the interaction between adult group members, group members without rearing experience and infants in the common marmoset, Callithrix jacchus

This study describes the reactions of adult, rearing-experienced members of three Callithrix jacchus families to interactions between animals without rearing experience and infants. The direct involvement of adult group members in situations in which inexperienced animals interact with infants decreases as the infants become older. Inexperienced animals will approach infants more seldom, if adult, rearing-experienced animals are nearby. Adult, rearing-experienced members of the group display primarily noninteractive behavior. This is especially true when no physical contact has taken place between an inexperienced animal and the infant, or if contact was initiated by the infant. Most of the dissociative reactions were observed in the context of competition for objects and play activity. However, there was a similarly high percentage of associative behavior towards inexperienced animals during play activity. In the episodes observed, the adult, rearing-experienced animals displayed mainly taking care behavior towards the infant but never any dissociative behavior. The following points appear to be of significance for the way adult, rearing-experienced members of the group behave: spatial distance between inexperienced animal and infant; which party was the initiator of the interaction; the level of experience already acquired by the animals without previous rearing experience, and the level of maturation of the infant.     

Relationship of early infant state measures to behavior over the first year of life in the tufted capuchin monkey (Cebus apella)

Data on activity states were collected from 29 group-housed capuchin monkey (Cebus apella) infants for 3 h each week from birth to 11 weeks of age. The amounts of time spent in sleeping/drowsy, alert–quiet, and alert–active states were measured in these subjects. Videotaped observations of these infants were recorded 3 times/week in the home cage over the first year of life and were scored for a number of social and exploratory behaviors. The extent to which early infant activity state scores predicted later behavior in the home cage was examined. Infant state measures correlated significantly with home cage behavior during months 2–6 in that infants that had been more active in early infancy spent more time alone, with other animals, and in exploration and play and less time with mothers than did quieter infants. Early state measures were less successful in predicting home cage scores beyond 8 months of age, whereas differences in behavior attributable to housing variables became more salient in the latter part of the first year. There was also a negative correlation between mother and infant activity in months 2 and 3, in that more sedentary mothers tended to have more active infants. Am. J. Primatol. 44:43–56, 1998. © 1998 Wiley-Liss, Inc. This article is a U.S. Government work, and, as such, is in the public domain in the United States of America.     

Multiple breeding females in captive groups of golden-headed lion tamarins (Leontopithecus chrysomelas): causes and consequences

In Callitrichidae, reproduction in subordinate females is generally inhibited but occurs in rare cases, possibly in association with the presence of an unrelated male, important food resources or low dispersal opportunities. This study investigates the occurrence of groups with multiple breeding females in captive golden-headed lion tamarins (Leontopithecus chrysomelas), the factors leading to their formation and the consequences for the group. Information obtained from studbook data on the world captive population during 1984-1998 revealed that polygynous groups in captivity are very rare: only 7 cases were discovered. Family groups in which daughters started breeding with a related male were larger than average, had a high number of sexually mature sons and eldest offspring that were well past the age of sexual maturity. Following a breeding attempt, severe aggression frequently occurred, especially if the infants were liveborn. Polygynous groups composed of two related females and an unrelated male tended to remain stable for a longer period than families with breeding daughters. Competition for infant care is probably an important factor determining whether the polygynous situation can persist and for how long.     

Case-control study of sudden infant death syndrome in Scotland, 1992-5.

OBJECTIVE: To investigate the relation between routine infant care practices and the sudden infant death syndrome in Scotland. METHODS: National study of 201 infants dying of the sudden infant death syndrome (cases) and 276 controls by means of home interviews comparing methods of infant care and socioeconomic factors. RESULTS: Sleeping prone (odds ratio 6.96 (95% confidence interval 1.51 to 31.97) and drug treatment in the previous week (odds ratio 2.33 (1.10 to 4.94)) were more common in the cases than controls on multivariate analysis. Smoking was confirmed as a significant risk factor (odds ratio for mother and father both smoking 5.19 (2.26 to 11.91)). The risk increased with the number of parents smoking (P < 0.0001), with the number of cigarettes smoked by mother or father (P = 0.0001), and with bed sharing (P < 0.005). A new finding was an increased risk of dying of the syndrome for infants who slept at night on a mattress previously used by another infant or adult (odds ratio 2.51 (1.39 to 4.52)). However, this increased risk was not established for mattresses totally covered by polyvinyl chloride. CONCLUSIONS: Sleeping prone and parental smoking are confirmed as modifiable risk factors for the sudden infant death syndrome. Sleeping on an old mattress may be important but needs confirmation before recommendations can be made.     

Social dynamics and individual plasticity of infant care behavior in cooperatively breeding cotton‐top tamarins

Individual variation in infant caretaking behavior is prevalent among marmoset and tamarin monkeys. Although most group members participate in infant care, the timing and amount provided differs greatly. In this study, we quantified general trends in infant carrying behavior by using a longitudinal database that included 11 years of instantaneous scan observations following 80 births of cotton‐top tamarins. Using detailed focal observations on a subset of the same families (10 births) we identified influences that affected expression of infant care at the group and individual levels. Fathers were the primary carriers and paternal carry time gradually decreased with increasing infant age. Paternal carry time also decreased significantly with an increasing number of older sibling helpers. Most fathers began to carry on the first day postpartum. However, we report circumstances in which fathers delayed carrying until almost a month postpartum. Fathers retrieved infants the most, although adult brothers' rates of retrievals peaked and surpassed fathers' rates during week 4 postpartum. Fathers delayed rejection of infants until week 4, whereas mothers rejected infants immediately and throughout the eight weeks. Nonetheless, infants climbed onto their mothers more than onto any other family member. Mothers showed a high initial investment in carrying during the first two weeks, decreasing quickly thereafter. Maternal contributions to infant carrying remained low and relatively consistent regardless of group size. However, mothers dramatically increased their infant carrying behavior in families in which fathers were absent. Older siblings cared for infants more than did younger siblings, and brothers retrieved and carried infants more than did sisters. Individual expression of infant care changed to accommodate infant needs and changed according to varying social dynamics and circumstances across litters. Am. J. Primatol. 72:296–306, 2010. © 2009 Wiley‐Liss, Inc.