Why Do African Elephants (Loxodonta africana) Simulate Oestrus? An Analysis of Longitudinal Data

Female African elephants signal oestrus via chemicals in their urine, but they also exhibit characteristic changes to their posture, gait and behaviour when sexually receptive. Free-ranging females visually signal receptivity by holding their heads and tails high, walking with an exaggerated gait, and displaying increased tactile behaviour towards males. Parous females occasionally exhibit these visual signals at times when they are thought not to be cycling and without attracting interest from musth males. Using demographic and behavioural records spanning a continuous 28-year period, we investigated the occurrence of this “simulated” oestrus behaviour. We show that parous females in the Amboseli elephant population do simulate receptive oestrus behaviours, and this false oestrus occurs disproportionately in the presence of naïve female kin who are observed coming into oestrus for the first time. We compare several alternative hypotheses for the occurrence of this simulation: 1) false oestrus has no functional purpose (e.g., it merely results from abnormal hormonal changes); 2) false oestrus increases the reproductive success of the simulating female, by inducing sexual receptivity; and 3) false oestrus increases the inclusive fitness of the simulating female, either by increasing the access of related females to suitable males, or by encouraging appropriate oestrus behaviours from female relatives who are not responding correctly to males. Although the observed data do not fully conform to the predictions of any of these hypotheses, we rule out the first two, and tentatively suggest that parous females most likely exhibit false oestrus behaviours in order to demonstrate to naïve relatives at whom to direct their behaviour.     

Age at maturity in wild baboons: genetic, environmental and demographic influences

The timing of early life-history events, such as sexual maturation and first reproduction, can greatly influence variation in individual fitness. In this study, we analysed possible sources of variation underlying different measures of age at social and physical maturation in wild baboons in the Amboseli basin, Kenya. The Amboseli baboons are a natural population primarily comprised of yellow baboons (Papio cynocephalus) that occasionally hybridize with anubis baboons (Papio anubis) from outside the basin. We found that males and females differed in the extent to which various factors influenced their maturation. Surprisingly, we found that male maturation was most strongly related to the proportion of anubis ancestry revealed by their microsatellite genotypes: hybrid males matured earlier than yellow males. In contrast, although hybrid females reached menarche slightly earlier than yellow females, maternal rank and the presence of maternal relatives had the largest effects on female maturation, followed by more modest effects of group size and rainfall. Our results indicate that a complex combination of demographic, genetic, environmental, and maternal effects contribute to variation in the timing of these life-history milestones.     

Life-history characteristics of a wild population of vervets (Cercopithecus aethiops sabaeus) in Barbados,West Indies

Life history data are presented for a population of vervets, Cercopithecusaethiops sabaeus, in Barbados, West Indies. The data were obtained from two habituated troops and from vervets captured during a large-scale trapping program. Individuals of known age from one troop were weighed periodically, and separate growth curves generated for males and females. The mean weight of captured adult females was 3.3 kg; that of adult males, 5.3 kg. The average age at sexual maturity is estimated at 34 months for females and 60 months for males. Vervets give birth throughout the year, but most infants are born between April and July. The average interbirth interval following a surviving infant is 11.8 months. The mortality of juveniles is heaviest between birth and 2 years of age and decreases thereafter. Males emigrate from their natal troops at sexual maturity and one incident of a juvenile female emigrating is reported.     

Coping with a challenging environment: effects of seasonal variability and reproductive status on glucocorticoid concentrations of female baboons (Papio cynocephalus)

Environmental stressors impact physiology and behavior in many species of animals. These effects are partly mediated through changing concentrations of glucocorticoids, which also vary with reproductive state and social conditions. Prior research has focused largely on seasonal breeders, but the close temporal linkage between season and reproductive state in these species hinders ability to disentangle environmental effects from those of the animal's reproductive status. Here we assessed the effects of environmental challenges on the fecal glucocorticoid (fGC) levels of non-seasonal breeders, female baboons (Papio cynocephalus) of Amboseli, Kenya. Amboseli is characterized by a long dry season, during which food and water become scarce, and by extreme temperatures above 40 °C in the shade during some months of the year. We found that after accounting for female reproductive status and individual variability, females exhibited higher fGC levels during the dry season than during the wet season. Further, during the wet season, fGC levels were higher in months of high average daily maximum temperatures. During the dry season, fGC levels were elevated both in hotter months and in months during which the baboons spent a relatively high proportion of time feeding. In spite of these stressors, female baboons reproduce during all months of the year in Amboseli, unlike most other mammals in this environment. This may be attributable to their extreme adaptability, specifically their diversified diet, and their ability to modify their behavior, including their activity profiles.     

Female Reproductive Parameters of Tana River Yellow Baboons

We describe the female reproductive parameters of yellow baboons at the Tana River National Primate Reserve, Kenya. We present data on menarche, cycle length, pregnancy, birth, postpartum amenorrhea, interbirth interval, and infant survival. We also briefly compare our data to those reported for yellow baboon females at the Amboseli Reserve, Kenya. Our results indicate statistically significant differences in some of these reproductive parameters between the two sites.     

Age determination by back length for African savanna elephants: extending age assessment techniques for aerial-based surveys

Determining the age of individuals in a population can lead to a better understanding of population dynamics through age structure analysis and estimation of age-specific fecundity and survival rates. Shoulder height has been used to accurately assign age to free-ranging African savanna elephants. However, back length may provide an analog measurable in aerial-based surveys. We assessed the relationship between back length and age for known-age elephants in Amboseli National Park, Kenya, and Addo Elephant National Park, South Africa. We also compared age- and sex-specific back lengths between these populations and compared adult female back lengths across 11 widely dispersed populations in five African countries. Sex-specific Von Bertalanffy growth curves provided a good fit to the back length data of known-age individuals. Based on back length, accurate ages could be assigned relatively precisely for females up to 23 years of age and males up to 17. The female back length curve allowed more precise age assignment to older females than the curve for shoulder height does, probably because of divergence between the respective growth curves. However, this did not appear to be the case for males, but the sample of known-age males was limited to ≤27 years. Age- and sex-specific back lengths were similar in Amboseli National Park and Addo Elephant National Park. Furthermore, while adult female back lengths in the three Zambian populations were generally shorter than in other populations, back lengths in the remaining eight populations did not differ significantly, in support of claims that growth patterns of African savanna elephants are similar over wide geographic regions. Thus, the growth curves presented here should allow researchers to use aerial-based surveys to assign ages to elephants with greater precision than previously possible and, therefore, to estimate population variables.     

Social affiliation matters: both same-sex and opposite-sex relationships predict survival in wild female baboons

Social integration and support can have profound effects on human survival. The extent of this phenomenon in non-human animals is largely unknown, but such knowledge is important to understanding the evolution of both lifespan and sociality. Here, we report evidence that levels of affiliative social behaviour (i.e. ‘social connectedness’) with both same-sex and opposite-sex conspecifics predict adult survival in wild female baboons. In the Amboseli ecosystem in Kenya, adult female baboons that were socially connected to either adult males or adult females lived longer than females who were socially isolated from both sexes—females with strong connectedness to individuals of both sexes lived the longest. Female social connectedness to males was predicted by high dominance rank, indicating that males are a limited resource for females, and females compete for access to male social partners. To date, only a handful of animal studies have found that social relationships may affect survival. This study extends those findings by examining relationships to both sexes in by far the largest dataset yet examined for any animal. Our results support the idea that social effects on survival are evolutionarily conserved in social mammals.     

Excreted urine,bladder urine,and the delay of sexual maturation in female house mice

A series of experiments tested effects of excreted and bladder urine of group- and singly-caged female house mice (Mus musculus) on sexual maturation of young female mice caged alone or in groups of eight. Puberty was determined by vaginal smears. A maturation-delaying pherompne was present in bladder urine of all female mice and excreted urine of group-caged females, but not in excreted urine from singly-caged females. Test mice treated with bladder urine homogenized with urethras from singly-caged females matured at the same age as controls. Apparently the urethras or associated glands of singly-caged females produce a substance which deactivates the maturation-delaying pheromone contained in bladder urine.     

Life history of yellow baboons: Physical development,reproductive parameters,and infant mortality

Longitudinal data from a population of yellow baboons,Papio cynocephalus, in the Amboseli National Park, Kenya, provide life history parameter estimates. Females reached menarche at approximately four-and-a-half years of age and then cycled for approximately a year before first conception. Postpartum anestrum averaged 12 months but ranged from six to 16 months. In cases of still births or infant death during postpartum amenorrhea, females commenced cycling after approximately one month. In mature females the time spent cycling before conception was five months on the average with a range from one to over 18 months. Only half of all full-term pregnancies resulted in infants who survived the first year of life; only a third, in infants who survived until the birth of their mother’s next infant. In comparison with data from laboratory colonies, our data indicate that female baboons in Amboseli are older at birth of first infant. They have, on the average, a somewhat shorter interbirth interval than was estimated from earlier crossectional field data, and therefore spend a larger portion of their adult life pregnant, but have a much longer interval—at least three years on the average—between the birth of an infant and the birth of that infant’s next older surviving sibling. A number of morphological changes in immature baboons are described.     

Patterns of coalition formation by adult female baboons in Amboseli, Kenya

Coalitionary support in agonistic interactions is generally thought to be costly to the actor and beneficial to the recipient. Explanations for such cooperative interactions usually invoke kin selection, reciprocal altruism or mutualism. We evaluated the role of these factors and individual benefits in shaping the pattern of coalitionary activity among adult female savannah baboons, Papio cynocephalus, in Amboseli, Kenya. There is a broad consensus that, when ecological conditions favour collective defence of resources, selection favours investment in social relationships with those likely to provide coalitionary support. The primary features of social organization in female-bonded groups, including female philopatry, linear dominance hierarchies, acquisition of maternal rank and well-differentiated female relationships, are thought to be functionally linked to the existence of alliances between females. Female savannah baboons display these characteristics, but the frequency and function of their coalitionary aggression is disputed. In our five study groups, 4-6% of all disputes between females led to intervention by third parties. Adult females selectively supported close maternal kin. There was no evidence that females traded grooming for support or reciprocated support with nonkin. High-ranking females participated in coalitionary aggression most frequently, perhaps because they derived more benefits from group membership than other females did or could provide support at lower cost. Females typically supported the higher ranking of two contestants when they intervened in disputes between subordinates, so most coalitions reinforced the existing dominance hierarchy. Results indicate that female baboons participate in coalitionary aggression in a manner strongly influenced by nepotism and individual benefits.     

Eating for two: Behavioral and environmental correlates of gestation length among free-ranging baboons (Papio cynocephalus)

Variation in environmental conditions during pregnancy and differences in the feeding behavior of females during pregnancy were consistently associated with variation in gestation length among free-ranging yellow baboons, Papio cynocephalus,in Amboseli National Park, Kenya. Females whose pregnancies ended during the wet season gave birth after shorter pregnancies than females whose pregnancies ended during the dry season. When rainfall is held constant, another source of variation in gestation length emerges. Females that spent progressively less time feeding over the course of their pregnancies gave birth after longer pregnancies than females that spent progressively more time feeding over the course of their pregnancies. These two factors, which provide rough indices of maternal nutritional status, accounted for a substantial fraction of the observed variation in gestation length among these female baboons.     

Group extinction and fusion in free-ranging vervet monkeys

Three cases of group fusion were observed among vervet monkeys in Amboseli National Park, Kenya, during the 1984 drought. In all cases, group fusion involved the migration of at least two females into a small neighboring group. Following migration, the females' natal groups ceased to exist. Although migrants initially assumed low ranks, none received high rates of aggression, and two rose in rank 6 months after their arrival. Migrant females appeared to employ a number of “strategies,” including grooming, allomothering, participation in intergroup encounters, and the formation of coalitions against males and each other to hasten integration. Results suggest that the causes of male and female migration differ fundamentally, and that group fusion occurs primarily in response to the inability of small groups of females to compete successfully with other groups for resources.     

Urine-marking activity of female mice under the influence of male urine odors

We conducted a study of the urine-marking activity of female mice when simultaneously exposed to urine odors from 2 kinds of males. Females, deposited a greater number of urine spots when presented with urine of normal, rather than castrated, males. Two androgen-dependent compounds known to be present in normal male urine enhanced female urine-marking when mixed with castrated male urine. These results are consistent with previous results concerning odor preference of females. However, females showed no marking preference between normal male urine and preputialectomized male urine, in spite of their preference for the former in our previous odor-preference study. Further experiments with various combinations of male urine revealed that females showed no marking preference when 1 of the 2 presented urine samples was from preputialectomized males, regardless of the other presented stimulus. We concluded that female urine marking is not a simple reflection of sexual preference, but possibly a phenomenon of a complex motivational system.     

Perimenstrual behavior changes among female yellow baboons: Some similarities to premenstrual syndrome (PMS) in women

Although numerous studies have documented changes in the behavior of nonhuman primate females around the time of ovulation, very little attention has been paid to behavior changes around the time of menstruation. Yet such information is obviously relevant to under standing the origins and etiology of the adverse mood and behavior changes experienced premenstrually by many women. Analysis of data obtained during 115 hours of observation on 13 female Amboseli (Kenya) baboons (Papio cynocephalus) during 24 menstrual periods showed that prior to and during the time of menstrual onset, these individuals exhibited distinct changes in their activity budgets, nearest-neighbor distances, and patterns of social interaction. Furthermore, in comparison to females around the time of ovulation, perimenstrual females showed increased rates of agonistic interaction and decreased (but nonzero) rates of sexual interaction with adult males. These premenstrual and perimenstrual behavior changes among female yellow baboons thus show some intriguing similarities to several commonly reported behavioral symptoms of premenstrual syndrome (PMS) in women.     

Pheromone-induced reproductive inhibition in young female Peromyscus leucopus

Soiled bedding and urine from adult female white-footed mice (Peromyscus leucopus) were tested for their capacity to inhibit reproduction of young females. Test animals were given either physical or airborne contact with soiled bedding from adult females, adult female urine, clean bedding, or water from 21 to 150 days of age. Results indicate that reproductive inhibition is due to an airborne pheromone emitted by the adult females as a component of their urine. In the second experiment, young female mice were exposed to an adult female for 0, 1, 3, 6, 12, 18, or 24 h/day from 21 to 150 days of age. Results from this experiment show that exposure to adult females of as little as 3 h/day was sufficient to cause reproductive inhibition to occur. This phenomenon has important implications in terms of both female-female reproductive competition and socially mediated population regulation.     

A releaser pheromone that attracts males in the urine of mature female masu salmon

A releaser pheromone of masu salmon Oncorhynchus masou was investigated using Y-maze behavioural experiments. During the reproductive season, urine of mature females contains a releaser pheromone which acts as a sex attractant for spermiating mature male parr. The releaser pheromone in mature female urine is one or more low molecular weight substances (less than 10 000) soluble in ether under basic conditions. The attractant was not present in either the coelomic fluid of ovulated females nor in neutral or acidic extracts of female urine which contain free steroids and F-type prostaglandins, respectively.     

Secondary sex ratio and maternal dominance rank among wild yellow baboons (Papio cynocephalus) of Mikumi National Park,Tanzania

The Trivers and Willard model predicts that offspring of dominant mothers will be biased toward males and offspring of subordinate mothers towards females, whereas a local-resource-competition hypothesis predicts the reverse. Available data bearing upon these alternative predictions are inconsistent. It has been suggested that the local-resource-competition hypothesis will predominate when resources are scarce, and the Trivers-Willard hypothesis when resources are abundant. The relationship between maternal dominance and secondary sex ratio for 214 offspring of 61 females was examined using four troops of wild yellow baboons living in Mikumi National Park, where the population was increasing in a resource-rich habitat. For all troops combined, no significant relationship was found between offspring sex ratio and maternal rank. The four troops separately showed inconsistent trends, and in no case did the relationship reach conventional levels of statistical significance. In contrast, the local resource-competition hypothesis was supported by a study of yellow baboons done with one troop and 80 offspring in Amboseli National Park, where a massive population decline had occurred. The contrasting Amboseli and Mikumi results may be due to differences in resource-competition at the two study sites, or to stochastic variation. © 1992 Wiley-Liss, Inc.     

Acceleration of puberty in female mice by a chemosignal from pregnant and lactating females: circadian rhythm effects

Two experiments were designed to test whether the urinary chemosignal excreted by pregnant and lactating female mice that accelerates puberty in young females is affected by circadian rhythms. The experiments also measured the possible influence of circadian rhythms on the response of the young recipient females. For urine from both pregnant and lactating females there was no difference in the effectiveness for accelerating puberty in urine collected during all 24 h. However, pregnancy urine used for treatment at 1800 and 0000 h, and lactation urine used for treatment at 1800, 0000 and 0600 h, all resulted in significantly earlier mean ages for puberty than pregnancy urine treatment at 0600 or 1200 h, or lactation urine treatment at 1200 h. There was also a significant interaction between the time of urine collection and the time of urine treatment for each urine source; urine was generally more effective in accelerating sexual development when used for treating young females at the same hour at which it had been collected, or at the time interval(s) just before or after the time at which it had been collected.     

Oestrous state dynamics in chemical communication by captive female Asian elephants

In many mammals, reproductive status is revealed through chemical cues in urine. The reproductive status of receivers may influence their interest in such signals. For social mammals that live in matrilineal groups, females may benefit by detecting the reproductive condition of herdmates. Responses to urine during oestrous cycles of senders and receivers are potential indicators of signal functions. We examined the chemosensory responses, first by four captive female Asian elephants,Elephas maximus , over their oestrous cycles to familiar follicular and luteal phase urine and second by 14 different female Asian elephants to unfamiliar conspecific follicular and luteal phase urine. We asked whether females could distinguish the reproductive state of another female as measured by their differential response to luteal- and follicular-phase urine. We further examined the influence of the receiver's reproductive status on response levels. Females responded more with specific tactolfactory trunk behaviours to follicular- than to luteal-phase urine, but only when the receiving female was in her follicular phase. Like their male conspecifics, Asian elephant females can detect changes in the reproductive state of conspecifics. The functional significance of this ability has yet to be determined but may be related more to the resource holding power of females in follicular phase than to a means for females to synchronize oestrous cycles. Such female-female communication may have important effects on social group dynamics. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Life history of hamadryas baboons: Physical development,infant mortality,reproductive parameters and family relationships

Demographic and life history parameters were estimated for a band of free-ranging hamadryas baboons, observed for 5.5 years in Ethiopia. Age-related changes in body weight and dentition were found to be delayed relative to laboratory-reared baboons. On the average, females reached menarche at 4.3 years of age and had their first infant at the age of 6.1 years. The mean interbirth interval was 24 months if the infant survived this period. The survival of infants and juveniles was higher compared to Amboseli yellow baboons, but somewhat lower than in gelada baboons in the Simen Mountains. Males acquired their first juvenile or adult female at the age of 8.5 to 11 years. Male-female pair-bonds lasted several years in most cases. The Cone Rock baboons were organized in a four-level social structure. The troop could split into bands, bands were divided into clans, and clans into one-male units with bachelor followers. The exchange of individuals between social units predominantly occurred within the band. All males of known origin became adult members of their presumed natal clan. Most females transferred also within clans, and juvenile females tended to remain in their natal clan. Females lost by one male to several rivals tended to reassemble in the same new one-male units later on.