Influence of the mother's reproductive state on the hormonal status of daughters in marmosets (Callithrix kuhlii)

Behavioral and endocrine suppression of reproduction in subordinate females produces the high reproductive skew that characterizes callitrichid primate mating systems. Snowdon et al. [American Journal of Primatology 31:11-21, 1993] reported that the eldest daughters in tamarin families exhibit further endocrinological suppression immediately following the birth of siblings, and suggested that dominant females exert greater control over subordinate endocrinology during this energetically challenging phase of reproduction. We monitored the endocrine status of five Wied's black tufted-ear marmoset daughters before and after their mother delivered infants by measuring concentrations of urinary estradiol (E(2)), pregnanediol glucuronide (PdG), testosterone (T), and cortisol (CORT). Samples were collected from marmoset daughters 4 weeks prior to and 9 weeks following three consecutive sibling-litter births when the daughters were prepubertal (M=6.1 months of age), peripubertal (M=11.9 months), and postpubertal (M=17.6 months). The birth of infants was associated with reduced ovarian steroid excretion only in the prepubertal daughters. In contrast, ovarian steroid levels tended to increase in the postpubertal daughters. Urinary E(2) and T levels in the postpubertal daughters were 73.8% and 37.6% higher, respectively, in the 3 weeks following the birth of infants, relative to prepartum levels. In addition, peak urinary PdG concentrations in peri- and postpubertal daughters were equivalent to luteal phase concentrations in nonpregnant, breeding adult females, and all of the peri- and postpubertal daughters showed clear ovulatory cycles. Cortisol excretion did not change in response to the reproductive status of the mother, nor did the concentrations change across age. Our data suggest that marmoset daughters of potential breeding age are not hormonally suppressed during the mother's peripartum period or her return to fertility. These findings provide an additional example of species diversity in the social regulation of reproduction in callitrichid primates.     

Post-conception reproductive competition in cooperatively breeding common marmosets

Common marmosets are cooperatively breeding monkeys that exhibit high female reproductive skew. Subordinate females usually fail to breed as a consequence of ovulation suppression and inhibition of sexual behavior, and, even when they do breed, typically rear fewer infants than dominants. We evaluated possible mechanisms of post-conception reproductive competition by comparing hormonal profiles across pregnancy, pregnancy outcomes, infant survivorship, and behavior in laboratory-housed families containing one (N=9) or two (N=7) breeding females. Breeding females in plurally breeding groups did not exhibit well-defined dominance relationships and rarely engaged in escalated aggression with one another. No significant differences were found among singly breeding mothers, plurally breeding mothers, and plurally breeding daughters in urinary chorionic gonadotropin or estradiol sulfate concentrations during pregnancy, fetal biparietal diameter, frequency of spontaneous abortion, frequency of stillbirths, number of live-born infants per litter, or infant mortality rates. When females gave birth while another female in the family was pregnant, however, their infants were highly likely to be killed. The perpetrator was definitively identified in only one family, in which a pregnant female killed her daughter's infant. These results are consistent with observations of free-living common marmosets and suggest that breeding females do not regularly influence one another's pregnancy outcomes, but that they may commonly kill each other's infants, especially during their own pregnancy. Our findings further suggest that infanticide by breeding females may have selected for the evolution of reproductive restraint in subordinate female marmosets.     

The reproductive status of nonbreeding group members in captive golden lion tamarin social groups

Reproductive activity is limited to only one female in many species of callitrichid primates (marmosets and tamarins): daughters and subordinate females do not produce offspring. A suppression of ovulatory cyclicity is responsible for the lack of reproductive activity in three species of callitrichids studied to date. This study evaluated the endocrine status of golden lion tamarins (Leontopithecus rosalia) housed as daughters or sons in family groups and of individuals housed in isosexual peer groups. Daughters 17 months of age and older and a subordinate female had high levels of estrogen excretion. Mean levels of estrogen excretion in these females were similar to those of nonpregnant, breeding adult females (17.14 ± 6.82 versus 11.93 ± 6.33 μg/mg creatinine, respectively). Estrogen profiles were similar to those of breeding adult females, with sinusoidal cycles in estrogen excretion. Younger daughters in family groups (10 and 12 months old) showed markedly lower levels of estrogen excretion (0.84 ± 0.58 μg/mg creatinine). Estrogen profiles lacked the sinusoidal nature of cycles in older daughters and breeding females, and elevations in estrogen excretion occurred frequently and remained elevated for 1 or 2 days. Plasma testosterone levels in males varied widely, but mean concentrations did not differ among males housed in different social conditions. These results suggest that older daughters and subordinate females may be capable of expressing normal ovarian function in the presence of a breeding adult female. This finding may account for two unusual observations in the lion tamarin: the high level of female-female aggression and the presence of groups in the wild with more than one actively breeding female.     

Behavioral and Hormonal Analysis of Social Relationships Between Oldest Females in a Wild Monogamous Group of Common Marmosets (Callithrix jacchus)

Social organization in common marmosets (Callithrix jacchus) appears to be based on a hierarchy of dominance, in which dominant females limit the reproductive functioning of subordinates. Reduced fertility was documented for captive but not for wild females. We investigated social dynamics and hormonal profiles of females living in a wild group. We collected behavioral and hormonal data during 7 mo from the reproductive and two nonreproductive adult females (probably daughters), and their interactions with the reproductive male. The subjects lived in a monogamous group at Nísia Floresta field station in northeastern Brazil. We collected behavioral data as all day follows, once a week and fecal material twice a week. We analyzed fecal samples for cortisol and progesterone using enzyme immuno assay (EIA) techniques. During the first three months we monitored the reproductive female (GRE) and the oldest nonreproductive female, (GIO). After the disappearence of GIO, we monitored the next oldest subordinate adult female, (GRA). GRE had just given birth at the beginning of the study, and she gave birth 6 mo later. During the last 2 weeks of GRE's gestation, GRA showed elevated progesterone levels, indicating ovarian cycling. Cortisol levels from GRE and GRA reached very high levels then. During GRA's cycling period, her interactions with the reproductive male (GT) were lower than when she was not cycling. We saw sexual interactions only between GT and GRE. The results provide evidence that ovarian function in mature subordinate females might be affected by the reproductive condition of the dominant female.     

Ovarian function of pygmy marmoset daughters (Cebuella pygmaea) in intact and motherless families

Reproductive inhibition of subordinate Callitrichid group members has been shown to vary with genus; whereas female Leontopithecus subordinates have normal ovarian cycles and occasionally breed within groups, subordinate Saguinus females almost never do so, with Callithrix species showing intermediate levels of reproductive inhibition. No information has been available on patterns of reproduction or inhibition in subordinate females in the genus Cebuella. We assessed fertility in Cebuella pygmaea daughters to allow comparison with the remaining Callitrichid genera. Specifically, the project had two goals: 1) to determine if there was evidence of reproductive inhibition of daughters living in family groups, and 2) to compare the ovarian function of daughters living in intact family groups with that of daughters residing in motherless families (i.e., without the breeding female). We collected daily urine samples for 6–8 weeks from eight pygmy marmoset daughters living in intact family groups or motherless families. Determination of ovulatory cycling or reproductive quiescence depended on the hormonal profiles generated from urinary luteinizing hormone and pregnanediol-3-glucuronide concentrations. All females in the motherless condition (aged 13–30 months) (n = 4) ovulated during the study. In contrast, only one of four daughters (aged 13–20 months) residing in intact families was found to be cycling. Data from motherless groups indicate ovarian cycling may begin between 15 and 17 months of age. Reproductive inhibition occurs in pygmy marmosets, although in a pattern more similar to Callithrix than to other Callitrichid genera. Am. J. Primatol. 43:347–355, 1997. © 1997 Wiley-Liss, Inc.     

Why do cotton‐top tamarin female helpers carry infants? A preliminary study

Shortly after giving birth, cotton-top tamarin mothers frequently attack the eldest female helpers [Snowdon et al., American Journal of Primatology 31:11-21, 1993]. Sometimes this aggression leads to the eviction of the eldest daughters from their natal groups 3-4 months after the birth of infants [Price & McGrew, Folia Primatologica 57:16-27, 1991]. We propose that daughters, during the act of carrying infants, may receive less aggression from mothers than when they are not carrying. On the other hand, given that mothers benefit from having others carry their infants, overall aggression received by female helpers from their mothers might be lower in those female helpers with a larger relative contribution to this activity. Four groups were observed during the first 9 weeks following the birth of infants, and aggression received was recorded for symmetrical as well as nonsymmetrical interactions. We found a positive correlation between contribution to infant carrying of female helpers and the overall aggression received from their mothers. Furthermore, the two eldest daughters with highest values of carrying contribution were evicted from their natal groups 3-4 months after the birth of infants. Although mothers do not appear to be more tolerant of female helpers that contribute the most to infant carrying, daughters do benefit from reduced aggression during the act of carrying, and remain in their natal group during the time period when infants must be carried.     

High rates of pregnancy loss by subordinates leads to high reproductive skew in wild golden lion tamarins (Leontopithecus rosalia)

Across taxa, cooperative breeding has been associated with high reproductive skew. Cooperatively breeding golden lion tamarins (Leontopithecus rosalia) were long thought to have a monogynous mating system in which reproduction was limited to a single dominant female. Subordinates with few reproductive opportunities delayed dispersal and remained in the natal group to provide alloparental care to siblings, thus allowing dominant reproductive females to meet the energetic needs associated with high rates of reproduction and successful infant rearing. The goal of this study was to re-assess monogyny in wild golden lion tamarin groups based upon pregnancy diagnoses that used non-invasive enzyme immunoassay for progesterone and cortisol, combined with weekly data on individual weight gain, bi-annual physical examinations noting pregnancy and lactation status and daily behavioral observations. We established quantitative and qualitative criteria to detect and determine the timing of pregnancies that did not result in the birth of infants. Pregnancy polygyny occurred in 83% of golden lion tamarin groups studied. The loss of 64% of subordinate pregnancies compared to only 15% by dominant females limited reproductive success mainly to dominant females, thus maintaining high reproductive skew in female golden lion tamarins. Pregnancy loss by subordinate adults did not appear to result from dominant interference in subordinate hormonal mechanisms, but more likely resulted from subordinate abandonment of newborn infants to mitigate dominant aggression.     

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.     

Fertility status,dominance, and scent marking behavior of family-housed female cotton-top tamarins (Saguinus oedipus) in absence of their mothers

The effect of the mother's absence on the fertility status and expression of scent marking behavior was examined in 12 sexually mature female cotton-top tamarins (Saguinus oedipus) living in four motherless families. Behavioral data and urine samples were collected on an average of five times per week and levels of estrone-glucuronide (E1G) were quantified directly by radioimmunoassay to establish whether the females showed ovarian cyclicity. Of the 12 females investigated only the dominant female in each motherless group exhibited regular E1G cycles. By contrast, low and acyclic hormonal profiles were found for subordinate sisters. In addition to differences in the patterns and levels of E1G, scent gland morphology and rates of scent marking differed between females: dominant, cycling females had elaborate glands and significantly higher rates of scent marking than subordinates. These results indicate that the presence of the mother (breeding female) is one important factor regulating ovarian function in cotton-top tamarin daughters. The findings also show that dominance status is a vital determinant of each female's subsequent physiological reproductive competence, with only the dominant female obtaining complete fertility and probably inhibiting ovulatory activity in her subordinate sisters. Scent marking behavior seems to be involved in the regulation of this phenomenon of intrasexual reproductive competition.     

Social influences on estrogen excretion and ovarian cyclicity in saddle back tamarins (Saguinus fuscicollis)

The effects of different social living conditions on estrogen excretion and on the ovarian cycle of saddle back tamarins were investigated. Urinary estrogens were monitored as indices of ovarian cyclicity in groups of adult females living under the following experimental conditions: (1) five parous females, each living with an adult castrated male (in one of the females plasma estradiol and progesterone were also measured); (2) five adult daughters living with their families. Each daughter was then removed from her family and paired with a vasectomized male. After pairing, scent marks collected from the family were introduced daily to one of the daughters and her mate. Scent exposure was then discontinued and hormone excretion measured for several weeks. The remaining four daughters were not exposed to family scent. The females living with castrated males showed urinary estrogen cycles of an average length of 17.5 ± 1.0 days. The plasma estrogen cycle was of the same length. The females studied under condition 2 showed low, noncycling estrogen levels while living in their families. They responded to pairing with an increase in the level of urinary estrogens, and four out of five showed regular estrogen cyclicity. The fifth female exposed to family scent marks after pairing also showed an increase in urinary estrogens. However, as long as scent transfer was maintained, no cycle was observed. Estrogen excretion increased again, and cyclicity commenced when scent transfer was discontinued. It is concluded that ovarian estrogen production is suppressed and cyclicity does not occur as long as daughters live in their families. Release from suppression and perhaps stimulation by the male cause a rapid increase in estrogen levels and the onset of cyclicity. Chemical stimuli produced by the family, perhaps particularly by the mother, may be involved in reproductive suppression.     

Behavioral strategies and hormonal profiles of dominant and subordinate common marmoset (Callithrix jacchus) females in wild monogamous groups

New insights into the mating systems of common marmosets suggest that they are mainly monogamous, although polygyny and polyandry occasionally occur. Long-term monitoring of wild common marmosets has shown that some reports of polygynous groups (i.e., groups that contain more than one reproducing female) in fact indicate an unbalanced reproductive output associated with extragroup copulation. In this study we describe the behavioral and hormonal profiles of common marmoset (Callithrix jacchus) females living in three wild monogamous groups (Q, PBf, and T), varying from five to 11 individuals, at Nísia Floresta field station, RN, Brazil. The mating system of the groups was previously characterized in terms of affiliative, sexual, and mate-guarding behaviors. Behavioral data were collected once a week, and fecal samples were collected at least twice a week for 10-16 months, depending on the group. A preferential allogrooming relationship was recorded between dominant males and females. Under field conditions the reproductive inhibition of subordinate females appears to be more behavioral than hormonal, since subordinate females of the three groups ovulated and two conceived during the study. In these cases, the subordinate and dominant females reproduced 1 month apart, and infanticide (one case confirmed and one suspected) appeared to be part of the reproductive strategy of dominant females. Following the infanticide, ovarian inhibition (group T) or emigration and return to the natal group (group PBf) were observed. In the third group (Q) the subordinate female showed hormonal profiles compatible with pregnancy, but no infants were seen. These findings reflect the different alternatives that wild subordinate common marmoset females use to reproduce.     

Onset of plural cooperative breeding in common marmoset families following replacement of the breeding male

Common marmosets, Callithrix jacchus, are usually characterized as singular cooperative breeders, with only a single, dominant female reproducing in each group. Anecdotal reports, however, have described two females breeding concurrently when an unrelated male joins their group. We tested the hypothesis that incorporation of an unrelated adult male into a family systematically leads to the onset of reproductive activity in a daughter and investigated the underlying mechanisms. We collected hormonal and behavioural data from mothers and their eldest daughters before and after the father was removed from the family and either replaced by an unrelated male (N=11) or immediately returned to the family (N=7). Variation between daughters in the occurrence of ovulatory cyclicity was not associated with the presence of an unrelated male but was closely linked to daughters' relationships with their mothers: only anovulatory daughters behaved submissively towards their mothers. Daughters never engaged in sexual behaviour with their fathers, but most did so with unrelated males. Similarly, daughters never conceived in intact natal families but did so in eight of 11 families containing an unrelated male. In six of these families, the mother and daughter bred concurrently. Thus, incorporation of an unrelated adult male into a marmoset family may frequently lead to the onset of plural breeding by activating sexual behaviour in a daughter; however, daughters ovulate only if they are not behaviourally subordinate to their mothers. Therefore, both inbreeding avoidance and rank-related reproductive suppression may constrain reproduction in marmoset daughters and contribute to maintenance of singular breeding.     

Differential reproductive success and facultative adjustment of sex ratios among captive female bonnet macaques (Macaca radiata)

In a group of captive bonnet macaques (Macaca radiata) housed at the California Primate Research Center, variance in reproductive success among females is primarily due to differences in infant survival. The infants of low-ranking females have a smaller probability of surviving to 6 months of age than do the infants of other females. In addition, the juvenile daughters of low-ranking females are more vulnerable to behaviourally induced mortality than are other immature animals. Observational evidence indicates that this mortality is the direct result of aggression by unrelated, higherranking adult females. Although infants' sex is not consistently related to survival, yearly fluctuations in the survival of male and female infants are reflected in the extent and direction of the skew in the sex ratio of offspring produced the following year. Years in which the highest proportion of male infants survive are followed by years in which the largest proportions of the birth cohorts are composed of males, and years in which the largest proportions of females survive are followed by years in which the largest proportions of birth cohorts are composed of females. For infant females the probability of surviving is reduced when a substantial proportion of the birth cohort is composed of females. The same pattern is evident among the sons of low-ranking females. The adaptive significance of behaviourally induced variation in reproductive success among females is considered in relation to these data.     

The effects of dominance rank and group size on female lifetime reproductive success in wild long-tailed macaques,Macaca fascicularis

Demographic changes were recorded throughout a 12-year period for three social groups ofMacaca fascicularis in a natural population at Ketambe (Sumatra, Indonesia). We examined the prediction that females' lifetime reproductive success depended on dominance rank and group size. Average birth rate was 0.53 (184 infants born during 349 female years). For mature females (aged 8–20 yr) birth rate reflected physical condition, being higher in years with high food availability and lower in the year following the production of a surviving infant. High-ranking females were significantly more likely than low-ranking ones to give birth again when they did have a surviving offspring born the year before (0.50 vs 0.26), especially in years with relatively low food availability (0.37 vs 0.10). Controlled comparisons of groups at different sizes indicate a decline in birth rate with rroup size only once a group has exceeded a certain size. The dominance effect on birth rate tended to be strongest in large groups. Survival of infants was rank-dependent, but the survival of juveniles was not. There was a trend for offspring survival to be lower in large groups than in mid-sized or small groups. However, rank and group size interacted, in that rank effects on offspring survival were strongest in large groups. High-ranking females were less likely to die themselves during their top-reproductive years, and thus on average had longer reproductive careers. We estimated female lifetime reproductive success based on calculated age-specific birth rates and survival rates. The effects of rank and group size (contest and scramble) on birth rate, offspring survival, age of first reproduction for daughters, and length of reproductive career, while not each consistently statistically significant, added up to substantial effects on estimated lifetime reproductive success. The group size effects explain why large groups tend to split permanently. Since females are philopatric in this species, and daughters achieve dominance rank positions similar to their mother, a close correlation is suggested between the lifetime reproductive success of mothers and daughters. For sons, too, maternal dominance affected their reproductive success: high-born males were more likely to become top-dominant (in another group). These data support the idea that natural selection has favored the evolution of a nepotistic rank system in this species, even if the annual benefits of dominance are small.     

Factors Influencing the Reproductive Success of Female Black Howlers (<Emphasis Type="Italic">Alouatta pigra</Emphasis>)

Measuring reproductive success is necessary to determine an organism's fitness, and thus, to address evolutionary processes. In this study we determined the effects of social, ecological, and individual attributes on the reproductive success of black howlers (Alouatta pigra) measured through infant survival and interbirth intervals (IBIs). From 2006 to 2012 we studied 29 black howler (A. pigra) females living in 11 groups in Campeche (Mexico), and recorded 82 births. We recorded group size and composition and the sex of infants during weekly surveys. We calculated a food availability index based on rainfall levels and on the size of preferred food resources. Daughters had a 30 % higher probability of surviving than sons, and the survival of the latter was positively related to food availability. IBI decreased when the first infant in the interval died, and when considering only IBIs in which the first infant survived, IBIs were longer following the birth of a daughter. These results suggest that, even if the production of daughters reduces reproductive output due to longer IBIs, female black howlers may still accrue higher reproductive success through their daughters due to differences between sexes in survival.     

Reproductive status of adult daughters in family groups of common marmosets (Callithrix jacchus jacchus)

Urine samples were collected from 6 pubertal female common marmosets at approximately 3-day intervals over a period of 3-7 months. All 6 females were the eldest daughters in their family groups. 3 of these females were subsequently removed from their families and sampled in isolation for another 2-3 months. Levels of urinary pregnanediol-3 alpha-glucuronide were measured by radioimmunoassay in order to establish whether these females showed ovarian cyclicity. No cyclical fluctuations in the levels of pregnanediol were observed in the females housed with their families even though the females were of a reproductively mature age. Cyclicity was quickly established (within 18-22 days) when the females were removed from the family. These results are discussed in the light of similar recent investigations on callitrichid reproduction and the behavioural implications are considered.     

Parent-daughter relationships and social controls on fertility in female common marmosets,Callithrix jacchus

This study examined the reproductive status of mothers and daughters to determine if parent-eldest daughter interactions would influence the daughter's fertility in the cooperatively breeding common marmoset, Callithrix jacchus. Twice weekly fecal samples were collected for hormonal analyses from seven daughters and their mothers and analyzed for cortisol, estradiol, and progesterone by EIA. Behavioral data were collected three times weekly while the daughters were under three conditions: natal family, after removing from the family and paring with a novel male, and after removing from the paired condition and placed back with the family (renatal). Under the natal condition, five of the seven daughters exhibited ovarian cycles while their mother was pregnant or displaying ovarian cycling. The five cycling daughters spent significantly more time initiating affilative behaviors with their mothers than with their fathers and showed significant changes in their behavior over the length of the ovarian cycle. However, aggression, submission, and sexual behaviors were very low for daughters in the natal phase. No differences between cortisol levels were found for a daughter and her mother. Cortisol levels showed a significant and sustained increase upon pairing within the first 20 days. All females but one cycled while paired. Six of the seven females were accepted back into the family after pairing and five of the seven females were ovulating. These results suggest that marmoset daughters are not necessarily reproductively suppressed while living with the family. Additionally, these data suggest that female marmosets that leave their family to explore mating opportunities with other groups may return to their family without reproductive or aggressive consequences.     

Synchronization of ovarian cycles within and between social groups in golden lion tamarins (Leontopithecus rosalia)

Synchronization of ovarian events has been reported in a number of primate species, with the temporal resolution of synchrony ranging from the occurrence of seasonal breeding within the annual cycle to a close matching of ovarian events within a single ovarian cycle. However, ovarian synchrony has not been reported in a New World primate. The temporal association of ovarian events was examined in female golden lion tamarins (Leontopithecus rosalia) living in the same or different social groups. Ovarian cyclicity was monitored by measuring the excretion of urinary estrogen metabolites. There was a high degree of synchronization in the occurrence of urinary estrogen peaks for females in different social groups (mean peak discrepancy = 2.1 days) and in females housed in the same social group (mean peak discrepancy = 1.3 days). Contrary to previous reports on callitrichid primates, daughters housed in their natal family group exhibited cyclic patterns of urinary estrogen excretion. These findings represent the first explicit demonstration of ovarian synchrony in a New World primate, and the tight coupling of ovarian cycles in female tamarins resembles the nature of menstrual synchrony in human females.     

Predicting primate responses to “Stochastic” demographic events

Comparative approaches in contemporary primate behavioral ecology have tended to emphasize the deterministic properties of stochastic ecological variables. Yet, primate responses to ecological fluctuations may be mediated by the interactions among demographic processes at the levels of individuals, groups, and populations. In this paper I examine long-term data collected from June 1982–July 1998 on one expanding group of muriquis (Brachyteles arachnoides) at the Estação Biologica de Caratinga, Minas Gerais, Brazil to explore the demographic and life history correlates of reproductive seasonality and skewed infant sex ratios. Variation in the size of annual birth cohorts (≥2 infants) was positively related to variation in the annual distribution of births (r _s=0.96,n=10,p<0.01), indicating the importance of considering the effects that the number of reproductive females may have on interpretations of reproductive seasonality. The female-biased infants sex ratio documented from 59 births was attributed exclusively to multiparous mothers. Primiparous mothers produced comparable numbers of sons (n=6) and daughters (n=7), and were increasingly likely to produce daughters with each subsequent reproductive event. Seven of the 11 females that have produced≥3 infants to date exhibited biases in favor of daughters whereas only 1 was biased in favor of sons. Variation in female sensitivity to local resource competition at different stages of their life histories may account for the female-biased infant sex ration in this population.     

Effects of social status, age, and season on androgen and cortisol levels in wild male golden lion tamarins (Leontopithecus rosalia)

This study examines factors affecting androgen and cortisol levels in wild, male golden lion tamarins (Leontopithecus rosalia). Golden lion tamarins are a cooperatively breeding species in which groups often contain two potentially breeding males. Brothers frequently emigrate together and develop a clear dominance hierarchy, but interactions between them are primarily affiliative. Duos in which the males are not related are less stable. In addition, reproductive skew theory predicts that dominant males will be less likely to share reproduction with related subordinates. As such, we predicted that both androgens and cortisol would be higher in subordinate males unrelated to the dominant male. We also predicted that androgens in breeding males would be higher during the mating season than the birth/infant care season, as per Wingfield's "challenge hypothesis" (1990). Fecal samples were collected from 24 males in 14 social groups and assayed by enzyme immunoassay. Androgen levels were higher in breeding males during the mating season, thus supporting the challenge hypothesis. However, while subordinate males unrelated to the dominant male had significantly lower androgens than any other group, cortisol levels were not correspondingly higher. These results suggest that unrelated subordinate males show measurable reproductive suppression and may use strategies such as infantilization to avert aggression from dominant males.