Variation in the resumption of cycling and conception by fecal androgen and estradiol levels in female Northern Muriquis (Brachyteles hypoxanthus)

We measured fecal androgen (T+DHT) and estradiol (E2) levels in female northern muriquis (Brachyteles hypoxanthus) at the Esta??o Biológica de Caratinga/RPPN Feliciano Miguel Abdala, Minas Gerais, Brazil, to evaluate the hormonal bases underlying individual variation in the resumption of cycling and conception. We found that androgen levels were significantly lower in females than in males, and that there were no consistent patterns in female androgen levels across precycling or cycling conditions. Females that resumed cycling earlier in the study (weeks 4-8) had higher precycling E2 levels and correspondingly lower precycling androgen/E2 ratios than females that resumed cycling later (weeks 12-16). There were no differences in female precycling androgen levels, but cycling females that conceived during or immediately after the study period had lower androgen levels and threefold higher E2 peaks than the one cycling female that failed to conceive. These results suggest that minimum E2 thresholds are necessary for both the resumption of ovarian cycling and conception. Individual variation in these components of fertility may be regulated by differences in E2 levels, which affect androgen/E2 ratios, rather than by androgen levels per se. Further research into the relative concentrations of T vs. DHT will be necessary to fully evaluate whether androgens affect cycling and conception in this species.     

Hormonal changes during the mating and conception seasons of wild northern muriquis (Brachyteles arachnoides hypoxanthus)

We investigated hormonal and behavioral changes in wild male and female northern muriquis (Brachyteles arachnoides hypoxanthus) at the Estação Biológica de Caratinga, Minas Gerais, Brazil, during a 6‐mo period that encompassed the onset of the 1998–1999 mating and conception seasons. Individual females resumed mating with the resumption of ovarian cycling, which was not synchronized among them or related to their cortisol levels. Females experienced two to seven cycles prior to conceiving, and the first conception occurred 2 mo after the onset of the group's mating season. There were no differences in female cortisol levels across their premating, mating, and conception conditions. Cortisol levels were significantly higher in females than in males prior to the conception season, consistent with the prediction that energy reserves may be associated with breeding readiness in females, but not males, in this species. The sustained elevation in male cortisol occurred after the peak in their sexual activity, which resulted in the first conception of the year. Male cortisol levels were positively correlated between years that were similar in rainfall, but differed in the timing of sexual and reproductive events. The timing of cortisol elevations in males appears to be generally regulated by environmental cues, but is responsive to fine‐tuning by social and behavioral cues related to the unpredictable timing of reproductive opportunities within their extended mating season. Am. J. Primatol. 61:85–99, 2003. © 2003 Wiley‐Liss, Inc.     

Socio-sexual behavior of female northern muriquis (Brachyteles hypoxanthus)

Female northern muriquis (Brachyteles hypoxanthus) are known to engage in frequent copulations with multiple partners, a pattern that in other primates has been attributed to various functions such as confusing paternity, reducing male aggression, or ensuring fertilization. However, in some female primates, promiscuity is restricted to times when conceptions are unlikely. We investigated whether female northern muriquis might exhibit a similarly mixed strategy by examining their mating, social, and activity patterns during their conception cycles versus other times. Systematic behavioral data were collected during an 18-month period between August 2001 and February 2003 on 13 adult females in a well-studied group at the RPPN-Feliciano Miguel Abdala, Minas Gerais, Brazil. Females mated on an average of 12.5+/-7.9 days during the study period, and spent significantly less time resting and engaging in non-sexual social behaviors, and significantly more time in sexual behaviors on days that they copulated than on days they did not. Three of the eight females for which sufficient data were available copulated significantly more often with their spatially closest non-kin associates, and four of five females that could be analyzed copulated significantly more often with their most frequent non-kin embrace partners. Comparisons between conception and non-conception periods revealed no differences in female activity budgets or in either the number of copulations or the number of different mating partners per female. Our results suggest that some females mate preferentially with close associates and social partners, but there is no indication that females alter their behavior during the cycles in which they conceive.     

An atypical reproductive cycle in a common viviparous Asia Agamid Phrynocephalus vlangalii

Viviparous lizards living in cold climate of high altitude often exhibit atypical reproductive cycles, in which mating and fertilization occur synchronously and annually with parturition occurring at the end of the year. Nevertheless, detailed case studies on atypical reproductive cycles are few. Using anatomical data combined with behavioral observations, we examined the reproductive cycle of a common Asian agamid, Phrynocephalus vlangalii, from a high‐elevation area in Sichuan, China. Male spermiation of P. vlangalii occurred in May, and spermatogenesis began in June and reached a maximum in October. For females, ovulation and fertilization occurred in May, and females developed gestation and pregnancy in 3 months from June to August, without vitellogenesis during this period. Females gave birth synchronously in late August, then vitellogenesis began and lasted until May of the next year. All adult males and females were synchronized in the same reproductive condition each month. The synchronous and annual reproductive cycle of P. vlangalii clearly represents an atypical cycle. The male courtship and mating behaviors were concordant with gonadal cycle and mainly happened in May and June. Despite the short growth period for neonates, they had a high over‐winter survival rate of 84.4%, suggesting that autumn parturition did not generate high costs to this reproductive cycle. We propose that the high over‐winter survival rate of neonates is likely linked with female delayed sexual maturity, female asynchronous vitellogenesis and gestation, large relative clutch mass (RCM), and adult‐offspring burrow sharing behavior during hibernation.     

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.     

Steroid excretion during the ovarian cycle in captive and wild muriquis,Brachyteles arachnoides

Urine, feces, and copulation frequency were collected from two captive muriqui females, Brachyteles arachnoides, at the Centro de Primatologia do Rio de Janeiro following the resumption of postpartum ovarian cycles. Fecal steroid profiles from seven wild muriqui females at the Estação Biologica de Caratinga, Minas Gerais, Brazil, were compared to the captive females to determine the approximate patterns of steroid excretion relative to the urinary LH peak. Hormonal profiles from one of the captive female muriquis revealed a discrete urinary LH peak. For this female, fecal progesterone increased on the same day as the urinary LH peak, while fecal estradiol increased 6 days later and urinary steroids increased 5 days later. For both captive females, the onset of fecal progesterone increase was preceded by the onset of copulations, which occurred during at least a 5-day period. The complete fecal hormonal profiles of the one captive female for which continuous data were available were similar to those found in wild muriqui monkeys, with the onset of an increase in sustained progesterone levels occurring several days prior to the onset of sustained estradiol increase. These patterns suggest that fecal progesterone may be excreted rapidly in this species. The onset of sustained increase in fecal progesterone levels, together with the consistent delay in the onset of the sustained increase in estradiol, may provide the best indicators of the periovulatory period for muriqui females. Am. J. Primatol. 42:311–321, 1997. © 1997 Wiley-Liss, Inc.     

Suppressed copulatory behavior and ovarian function in lactating Japanese monkeys (Macaca fuscata fuscata) during the mating season

The influence of lactation on copulatory behaviors and ovarian functions was studied in Japanese monkeys (Macaca fuscata fuscata) during the mating season. Three lactating females were housed in an outdoor group cage with their infants, and three nonlactating females were housed in an adjacent outdoor cage. They were mated by introduction of one of four rotationally chosen males into the females' cage, for two hours three times a week; the occurrence of ejaculatory copulations was recorded. Blood samples were collected on each observation day, and plasma levels of estradiol, progesterone, and luteinizing hormone (LH) were measured by specific radioimmunoassays. In nonlactating females, plasma estradiol increased during the transition into the mating season, and rose to levels over 90 pg/ml for the first time on about 50 days before the first ovulation. Shortly after plasma estradiol exceeded 90 pg/ml in the nonlactating females, the onset of ejaculatory copulations occurred. They received ejaculations continuously up to the early ovarian luteal phase. On the other hand, in lactating females, there were lower levels of plasma estradiol (below 90 pg/ml) during the transition into the mating season, and they received no ejaculation during that period. Two of the three lactating females ovulated only once, and they received ejaculations only during the periovulatory period, coinciding with the rise of their plasma estradiol levels over 90 pg/ml. The remaining lactating female remained anovulatory and received no ejaculation throughout the entire mating season. These results have demonstrated that the low sexual activity of lactating females is clearly correlated with low levels of plasma estradiol due to suppressed ovarian function.     

Individual copulatory preference and the “strange female effect” in a captive group-living male chimpanzee (Pan troglodytes)

The common chimpanzee has been considered to be a promiscuous species, although transient consort relationships and male possessive behavior have been described byTutin (1975, 1979). A prolific adult male chimpanzee was housed with from four to seven adult female chimpanzees (depending on the females' maternal status) and copulations were recorded from August 7, 1978 until February 16, 1979, during morning feeding periods. Ten females composed the fluctuating available partners. Sixty-four observed copulations involved six females. One female was clearly preferred, including occasions when she was not maximally tumescent (e.g., completely detumescent, pregnant) and other available females were maximally tumescent. The remainder of the observed copulations, with one exception, involved females who had recently been reintroduced into the one-male breeding group. Such copulations took place on and continued temporarily after the day of reintroduction for two females, and after the resumption of menstrual cycling for two females who had been reintroduced while still lactating. Nonpreferred females were impregnated during the period of data collection, even though copulations with them were not observed. These data suggest that the male chimpanzee can form an individual mating preference regardless of the hormonal status of his available partners without lessening his reproductive success with nonpreferred females, but also tends toward maximization of his reproductive success by copulating with novel females.     

New World primates,new frontiers: Insights from the woolly spider monkey,or muriqui (Brachyteles arachnoides)

Despite over 25 years of intensive research, much of our knowledge of primate behavior has been limited to a small number of Old World, semiterrestrial species. With the exception of chimpanzees, these species share consistent patterns of behavior, including aggressive competition between males, male dominance over females, male dispersal and female kin groups, that have come to characterize the Order Primates. However, as research has expanded to include a number of newly studied species, such generalizations about primate behavior are no longer appropriate. Data presented on the wooly spider monkey, or muriqui (Brachyteles arachnoides), provide examples of the ways in which traditional views of primate behavior are being reconsidered. In muriquis, and in several other New World monkeys, female kin groups do not occur because females disperse from their natal groups. In addition, aggressive competition between males is virtually absent; and, female muriquis appear to be codominant with males. Constraints on sexual dimorphism may help to explain why muriquis (and other New World primates) differ so fundamentally from the better known semi-terrestrial Old World monkeys.Logo of the 1988 International Primatological Society.     

Hormonal and behavioral changes at puberty in the squirrel monkey

Developmental changes in the reproductive behavior and physiology of 9 male and 15 female juvenile squirrel monkeys were evaluated in a 20-month study. Plasma levels of gonadal steroids remained relatively low for this species until most animals reached puberty between 2.5 and 3 years of age. Longitudinal assessment of plasma progesterone levels indicated that the onset of ovarian cycles tended to be synchronized between females although the 5 heaviest females began to cycle earlier than the rest. The heavier females reached puberty at a time which was appropriate to their birth in the wild, whereas most of the remaining females conceived 6 months later during a second period of reproductive activity that coincided with the laboratory mating season. Pubescent males underwent their first seasonal elevation in plasma testosterone levels during the second period and its onset was synchronized across all males. Thus, even in the absence of adults, pubertal processes in the squirrel monkey were strongly influenced by the seasonal breeding pattern. In addition, behavioral observations revealed that social maturation closely parallels reproductive ability in females, whereas males enter a protracted subadult stage after puberty.     

Timing of births in sympatric brown howler monkeys (Alouatta fusca clamitans) and northern muriquis (Brachyteles arachnoides hypoxanthus).

We monitored the birth patterns of sympatric brown howler monkeys (Alouatta fusca clamitans) and northern muriquis (Brachyteles arachnoides hypoxanthus) during a 4‐yr period from October 1996 to August 2000 at the Estação Biológica de Caratinga, Minas Gerais, Brazil. Brown howler monkey births (n = 34) occurred throughout the year, and birth frequencies did not differ between rainy and dry season months. The aseasonal birth patterns of the howler monkeys differed significantly from the dry season concentration and dry month peak in muriqui births (n = 23). We found no effects of infant sex or the number of females on interbirth intervals (IBIs) in our 10 howler monkey study troops. IBIs of brown howler monkeys averaged 21.2 ± 2.5 mo (n = 8, median = 21.0 mo), and were significantly shorter following dry season births than rainy season births. Their IBIs and yearling survivorship (74%) were similar to those reported for other species of howler monkeys, but yearling survivorship was much lower than that of muriquis (94%), whose IBIs were more than 12 mo longer than those of the howler monkeys. Our study extends comparative knowledge of birth patterns in Alouatta to a poorly known species, and provides insights into the different ways in which diet and life history may affect the timing of births in large‐bodied platyrrhines under the same seasonal ecological conditions. Am. J. Primatol. 55:87–100, 2001. © 2001 Wiley‐Liss, Inc.     

The stimulatory effect of males on the initiation but not the maintenance of ovarian cycling in cotton-top tamarins (Saguinus oedipus)

Prior studies have shown that female cotton-top tamarins usually do not ovulate while living with natal groups, and most females do not ovulate until they are paired with an unfamiliar adult male. To examine the role of unfamiliar adult males on stimulating ovarian function, four cotton-top females were studied during three conditions: females living with their natal group for six weeks, living alone but exposed to a single unfamiliar adult male located 15 cm away from the female's cage for four weeks, and living with an unfamiliar adult male for six weeks. Behavior and urinary hormonal concentrations were measured during the three conditions. Exposure to the male consisted of visual, auditory, and airborne olfactory contact. First ovulation occurred during exposure to the unfamiliar male in three of the four females indicating that direct physical or sexual contact with the male is not required for onset of ovarian cycling. The fourth female did not ovulate even during six weeks of direct contact with the unfamiliar adult male. In addition, four parous females in either family groups (3) or singly caged (1) were examined for ovarian function 4–6 months after the death or removal of their mates. All femals continued to cycle in the absence of the male indicating that the male was not needed to continue ovarian cycling. In fact, two of the females were pregnant at the time their males died and both delivered normal infants and resumed cycling. The results of this study indicate that an unfamiliar adult male may facilitate the onset of ovarian cycling without being in direct contact with the female and visual, auditory, or airborne olfactory cues may be involved. Once repeated ovarian cycling occurs the male is not required to maintain ovarian function.     

Food consumption and growth of California sea lions (Zalophus californianus californianus)

The daily food consumption of 26 California sea lions at the Harderwijk Marine Mammal Park was recorded. Average annual food consumption of males increased with age to stabilize at approximately 4,000 kg/year by the age of 10 years. Females showed a rapid increase in average annual food consumption until they were 3 years old. Thereafter, females housed outdoors averaged 1,800 kg/year, whereas those housed indoors ate approximately 1,400 kg/year. Between the ages of 4 and 7 years, the food intake of males began to fluctuate seasonally, decreasing between May and August. The low food intake in summer was associated with an increase in aggressive behavior. Seasonal fluctuation in the food intake of non‐reproductive females was negligible. Between the ages of 6 and 8 years, many females began to reproduce successfully. Pups were born in May and June. The females' food intake decreased approximately 3 days before birth and ceased the next day. Feeding resumed the day after birth, and by 2 days after birth, it had usually returned to normal. On average, female intake increased in the year of conception, the year of birth, during which the pup was suckled for 6 months, and the following calendar year, during which the pup was weaned. Pups began to eat fish at approximately 11 months of age. When forcefed, they were fully weaned within 2 to 23 days. Male weight and body length increased until approximately 20 years of age. Females increased in body length until 6 years and in weight until approximately 13 years of age. The relationship between standard body length and body weight is given. The heavier an animal is, the lower is its food intake as a percentage of body weight. Zoo Biol 19:143–159, 2000. © 2000 Wiley‐Liss, Inc.     

Behavioral Correlates of Dispersal in Female Muriquis (Brachyteles arachnoides)

We documented four adolescent female transfers—two emigrations, two immigrations—during a 12-month study from August 1994 to July 1995 on one group of muriquis (Brachyteles arachnoides) at Estação Biológica de Caratinga in Minas Gerais, Brazil. Intergroup transfers occurred throughout the year independent of season. A total of 613 focal samples of 10-min duration conducted on six adolescent females (5–9 years of age) revealed significant differences in the behavior of migrant versus natal resident females. During the seasons encompassing their respective group transfers, both emigrants and immigrants devoted more of their time to resting than resident females belonging to the same age cohorts did. Time spent feeding on mature fruit was higher for one emigrant and lower for both immigrants compared to the two resident females. Emigrant females had fewer neighbors within a 1-m radius than resident females did, whereas immigrant females were within a 1-m radius of adult females and within a 5-m radius of adult males more often than resident adolescent females were. Adolescent females were displaced on 26 occasions. Displacements occurred mainly during the dry seasons (n = 21) and mainly at food sources (n = 21). Using the number of focal samples conducted on each female as an estimate of observation time, immigrant females were displaced at twice the rate of residents. However, like other behavioral differences detected between resident and migrant adolescent females, differences in individual displacement rates were evident only during the season in which each of the immigrants joined the group. Collectively, our findings imply that female intergroup transfer in this population involves relatively mild, short-term costs.     

Fecal cortisol levels in free-ranging female chacma baboons: relationship to dominance, reproductive state and environmental factors

Savannah baboons are one of the few mammalian species that do not exhibit seasonal reproduction patterns and are therefore ideally suited to study the effect of female reproductive states (cycling, pregnant, lactating) on cortisol levels independent of seasonal factors. Fecal samples from 10 free-ranging female chacma baboons (Papio hamadryas ursinus), collected during a period of 17 months, were analyzed using a steroid-extraction method. Reproductive state had a significant effect on fecal cortisol, with lowest levels found in estrous females. Fertility was not related to fecal cortisol levels; we found no significant differences between samples collected on conceptive and nonconceptive cycles. Environmental factors explained most of the variance of fecal cortisol levels. Cortisol measures were strongly correlated with seasonal differences such as daylight duration, temperature and the amount of time that baboons spent resting. We measured higher cortisol levels during winter months and suggest that this could be related to shorter resting periods and to the cold minimum ambient temperatures at this study site. Finally, we found no relationship between social rank nor the rate of agonistic interactions with basal fecal cortisol levels.     

Fecal progesterone metabolites and ovarian activity in cycling and pregnant mountain gazelles (Gazella gazella)

Fecal reproductive progestagen monitoring in the mountain gazelle (Gazella gazella) provided a non-invasive method for tracking reproductive cycling, estimating age of sexual maturity and diagnosing pregnancy in this species of gazelle. Fresh fecal samples were collected from eight female mountain gazelle (Gazella gazella) for a period of two months. Two of the animals were pregnant while the other six were not. Using the progestagen profile the luteal phase, interluteal (follicular) phase and estrous cycle in adult female gazelles were determined to be 12.5 ± 1.2, 5.9 ± 0.59 and 18.8 ± 0.98 days respectively. Significant inter-animal differences in fecal progestagen concentration were observed in both the luteal and follicular phases. Significant differences were observed in the levels of fecal progestagen between cycling females and females in late pregnancy. Low concentrations of fecal progestagen in females aged less than 18 months old indicated that sexual maturity in captivity is not attained before that age.     

Lack of pubertal influences on female dispersal in muriqui monkeys, Brachyteles arachnoides

The hormonal mediation of dispersal in female mammals is poorly understood, in part because of the difficulties of detecting the onset of ovarian cycling and puberty in dispersing individuals. We used noninvasive methods of faecal steroid assays to determine the timing of dispersal relative to puberty and ovarian cycling in wild female muriqui monkeys, a species in which males are philopatric and nearly all females transfer from their natal groups. Natal females had a mean+/-SE age of 73.4+/-7.2 months (N=18) at the time of their transfers. Intergroup transfers occurred when one or more sexually active adult females were present, but did not show any seasonal patterns. Faecal progesterone and oestradiol profiles from nine natal females prior to transfer and four non-natal females that transferred into our study group demonstrate unequivocally that dispersal occurs prior to puberty in this species. All females showed baseline oestradiol levels and low progesterone levels compared with cycling adult females. Immigrants were first observed to copulate at 11.2+/-2.2 months of age (N=4), prior to the onset of normal ovarian cycles, and gave birth to their first offspring at 33.8+/-7.3 months (N=4) after transferring. Mean cortisol levels did not differ between natal emigrants or recent immigrants, and were within the range of those of adult males during the nonbreeding season in 10 of the 11 prepubertal females sampled. These results indicate that female dispersal is not triggered by activational hormones associated with puberty or escape from reproductive suppression in this species. Copyright 2000 The Association for the Study of Animal Behaviour.     

Social and hormonal mechanisms underlying male reproductive strategies in black howler monkeys (Alouatta pigra)

We investigated the social and hormonal mechanisms underlying male reproductive strategies in two multimale-multifemale groups of black howler monkeys (Alouatta pigra) during a 14-month study in Palenque National Park, Mexico. Fecal glucocorticoid (fGC) and androgen (fA) levels were analyzed for 343 fecal samples collected from 14 males during their presence in the study groups. Neither immigrating males nor resident males that remained in the group had elevated fGC and fA levels during 11 observed male migration events, suggesting that competition over group membership was not associated with variation in the fecal hormonal levels of males. Instead, fGC and fA levels were significantly higher in males who maintained a central position in the group and had almost exclusive access to fertile females than in other resident males. These “central” males were responsible for maintaining close spatial associations and cultivating strong affiliative relationships with cycling, sexually active females but not with noncycling, sexually inactive females. “Noncentral” males did not form strong social relationships with either cycling or noncycling females and had no or very few mating opportunities. Our findings suggest that male black howler monkeys compete nonaggressively by fostering relationships with cycling females and that the elevated fGC levels of central males may be indicative of the social challenges involved in their indirect competition.     

Plasma cortisol responses to stress in lactating and nonlactating female rhesus macaques

Lactating female rats without their pups exhibit lower HPA responsiveness to stress than nonlactating females. However, responsiveness to stress is similar when lactating females are tested with their pups and the stressor involves a potential threat to the offspring. This study constitutes the first comparison of stress responsiveness in lactating and nonlactating female nonhuman primates. Subjects were 53 multiparous female free-ranging rhesus macaques. Approximately half of the females were lactating and half were nonpregnant/nonlactating. Blood samples were obtained after capture and after overnight housing in an individual cage. Lactating females were tested with their infants. Lactating females had significantly higher plasma cortisol levels than nonlactating females on both days. Having or not having an infant was also a better predictor of plasma cortisol levels among all females than their age, dominance rank, group of origin, time of day at which the sample was obtained, and time elapsed since beginning of the sampling procedure or since anesthesia. Plasma cortisol levels of lactating females were not significantly correlated with post-partum stage or with the cortisol levels of their infants. Capture, handling, and individual housing in a cage are powerful psychological stressors for free-ranging primates. We suggest that the higher plasma cortisol levels exhibited by lactating females reflect greater responsiveness to stress associated with perception of risks to infants. Hyporesponsiveness to stress may not be a general characteristic of lactation in all mammalian species, but a short-term effect of infant suckling that is most apparent with stressors unrelated to the offspring.     

Group Composition Affects Seasonal Birth Timing in Captive Japanese Macaques

We analyzed birth dates recorded during an 18-year period in a group of Japanese macaques housed in the Rome zoo to assess the influence of environmental, physiological, and social factors on birth seasonality. Birth timing differed significantly among years. Birth timing was affected by reproductive condition of females—ones that had given birth in the previous year delivered significantly later than those that had not—but not by their age or dominance rank. We conducted further analyses separately on females that had or had not given birth in the previous year. In both subgroups of females mean birth date was not influenced either by environmental temperature and rainfall during the previous mating season or by group size. On the contrary, among females that had not given birth in the previous year, socionomic sex ratio—ratio of sexually mature males to sexually mature females—is positively correlated with both mean birth date and date of the first birth, but not with date of the last birth. Contrarily, among females that had given birth in the previous year, there is no significant relationship between these variables. We hypothesize that the effects of socionomic sex ratio on birth timing might depend on competition among males for access to fertile females. When the number of males per female was higher, mutual disruption of consort pairs may have led to a delay in the onset of mating.