Jeanne Altmann

Observational study of behavior: Sampling methods.

Seven major types of sampling for observational studies of social behavior have been found in the literature. These methods differ considerably in their suitability for providing unbiased data of various kinds. Below is a summary of the major recommended uses of each technique: In this paper, I have tried to point out the major strengths and weaknesses of each sampling method. Some methods are intrinsically biased with respect to many variables, others to fewer. In choosing a sampling method the main question is whether the procedure results in a biased sample of the variables under study. A method can produce a biased sample directly, as a result of intrinsic bias with respect to a study variable, or secondarily due to some degree of dependence (correlation) between the study variable and a directly-biased variable. In order to choose a sampling technique, the observer needs to consider carefully the characteristics of behavior and social interactions that are relevant to the study population and the research questions at hand. In most studies one will not have adequate empirical knowledge of the dependencies between relevant variables. Under the circumstances, the observer should avoid intrinsic biases to whatever extent possible, in particular those that direcly affect the variables under study. Finally, it will often be possible to use more than one sampling method in a study. Such samples can be taken successively or, under favorable conditions, even concurrently. For example, we have found it possible to take Instantaneous Samples of the identities and distances of nearest neighbors of a focal individual at five or ten minute intervals during Focal-Animal (behavior) Samples on that individual. Often during Focal-Animal Sampling one can also record All Occurrences of Some Behaviors, for the whole social group, for categories of conspicuous behavior, such as predation, intergroup contact, drinking, and so on. The extent to which concurrent multiple sampling is feasible will depend very much on the behavior categories and rate of occurrence, the observational conditions, etc. Where feasible, such multiple sampling can greatly aid in the efficient use of research time.

Social relationships among adult female baboons (papio cynocephalus) I. Variation in the strength of social bonds

Sociality has positive effects on female fitness in many mammalian species. Among female baboons, those who are most socially integrated reproduce most successfully. Here we test a number of predictions derived from kin selection theory about the strength of social bonds among adult female baboons. Our analyses are based on systematic observations of grooming and association patterns among 118 females living in seven different social groups in the Amboseli Basin of Kenya over a 16-year period. Females in these groups formed the strongest bonds with close kin, including their mothers, daughters, and maternal and paternal sisters. Females were also strongly attracted toward females who were close to their own age, perhaps because peers were often paternal sisters. Females’ bonds with their maternal sisters were strengthened after their mother’s deaths, whereas their relationships with their maternal aunts were weakened after their mother’s death. In addition, females formed stronger bonds with their paternal sisters when no close maternal kin were available, and they compensated for the absence of any close kin by forming strong bonds with nonrelatives. Taken together, these data suggest that social bonds play a vital role in females’ lives, and the ability to establish and maintain strong social bonds may have important fitness consequences for females.

True paternal care in a multi-male primate society

Although male parental care is rare among mammals, adult males of many cercopithecine primate species provide care for infants and juveniles. This care is often in the form of grooming, carrying, support in agonistic interactions, and protection against infanticide. For these behaviours to be interpreted as true parental care, males must selectively direct care towards their own offspring and this care must result in fitness benefits. With the exception of males defending probable offspring from infanticide, male primates living in multi-male, multi-female social groups have not been shown to selectively direct care towards their own offspring. We determined paternity for 75 juveniles in a population of wild savannah baboons (Papio cynocephalus) and collected data on interventions in agonistic disputes by adult males on behalf of juveniles as a form of male care. Here we show that adult males differentiate their offspring from unrelated juveniles and selectively support their offspring in agonistic disputes. As support in agonistic disputes is likely to contribute to rank acquisition and protect juveniles from injury and stress, this can be considered true parental care.

Balancing Costs and Opportunities: Dispersal in Male Baboons

Young male baboons typically disperse from their group of birth as they near adult size and may continue to migrate between social groups throughout their lives. Long-term data on dispersal and residence patterns of male baboons in Amboseli National Park, Kenya, were available for 110 males in the population, including 43 that were monitored during their natal dispersal. These data enabled us to provide not only a detailed evaluation of the effects of reproductive competition on dispersal but also the first direct estimates of the costs of dispersal in male primates and one of the few direct estimates of fitness costs associated with breeding in the natal group. Males underwent natal dispersal at a median age of 8.5 yr (range, 6.8-13.4 yr) and subsequently remained in nonnatal groups for a median tenure of 24 mo (range, 1-138 mo). Half of the males in the study engaged in moderate to extensive reproductive activity before natal dispersal. Reproductive costs associated with breeding in the natal group were suggested by the high mortality of offspring for whom natal males were their likely fathers, even though maternal relatives avoided mating with each other. Dispersal involved considerable time spent alone, and therefore the costs of dispersal were substantial, because of mortality risks and missed reproductive opportunities during dispersal. Female availability and male mating success apparently affected both natal and secondary dispersal patterns. We present a model of dispersal tendency in order to explicate the ways in which differences in population density, predation risk, and the distribution of mating opportunities among groups might result in complex dispersal patterns that are consistent with both the results of the present study and the disparate empirical reports in the literature.

Queuing and queue-jumping: long-term patterns of reproductive skew in male savannah baboons, Papio cynocephalus

In many animals, variance in male mating success is strongly correlated with male dominance rank or some other measure of fighting ability. Studies in primates, however, have varied greatly in whether they detect a relationship between male dominance rank and mating success. This variability has led to debate about the nature of the relation between rank and mating success in male primates. We contribute to the resolution of this debate by presenting an analysis of the relationship between dominance rank and male mating success over 32 group-years in a population of wild savannah baboons. When data were pooled over the entire period, higher-ranking males had greater access to fertile females. However, when we examined successive 6-month blocks, we found variance in the extent to which rank predicted mating success. In some periods, the dominance hierarchy functioned as a queue in which males waited for mating opportunities, so that rank predicted mating success. In other periods, the queuing system broke down, and rank failed to predict mating success when many adult males were in the group, when males in the group differed greatly in age, and when the highest-ranking male maintained his rank for only short periods. The variance within this single population is similar to the variance observed between populations of baboons and between species of primates. Our long-term results provide strong support for the proposition that this variance is not an artefact of methodological differences between short-term studies, but is due to true variance in the extent to which high-ranking males are able to monopolize access to females.  2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.

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.

Long-Term Consistency of Dominance Relations Among Female Baboons (Papio cynocephalus).

At maturity, female baboons in the Amboseli National Park of Kenya generally attain a rank position among adults near to that of their mothers. However, the age of a females mother and the difference in ages between sisters also influence the rank acquisition process. These latter demographic variables, which are sensitive to changes in resource availability, may account for the close association both within and among primate species of specific patterns of rank organization and specific environmental conditions.

Social relationships among adult female baboons (Papio cynocephalus) II. Variation in the quality and stability of social bonds

A growing body of evidence suggests that social bonds have adaptive value for animals that live in social groups. Although these findings suggest that natural selection may favor the ability to cultivate and sustain social bonds, we know very little about the factors that influence the quality or stability of social bonds. Here, we draw on data derived from a 16-year study of baboons living in seven different social groups in the Amboseli basin of Kenya to evaluate the quality and stability of social bonds among females. Our results extend previous analyses, which demonstrate that females form the strongest bonds with close maternal and paternal kin, age mates (who may be paternal kin), and females who occupy similar ranks but are not maternal relatives. Here we show that the same factors influence the quality and strength of social bonds. Moreover, the results demonstrate that the quality of social bonds directly affects their stability.

A matter of time: evaluating the storage of fecal samples for steroid analysis.

The extraction and immunoassay of fecal steroids is an increasingly common technique, used in both captive and field studies to provide an approximation of an animals circulating concentration of hormones through non-invasive methods. Storage of fecal samples is of critical concern because fecal bacteria metabolize fecal steroids within hours after deposit. Ethanol is often used as a preservative for fecal samples stored for several hours at room temperature. We examined the stability of fecal estrogen (fE) and glucocorticoid (fGC) metabolites from baboon (Papio cynocephalus) samples in a 95% ethanol solution at ambient temperature and at -20 degrees C over the course of six months, to determine the effect of storage on steroid concentrations. As measured by radioimmunoassay, fE metabolite concentrations increased by 122% at 90 days and fGC metabolite concentrations increased by 92% at 120 days. After peaking, both hormones declined to near initial concentrations by 180 days in ambient temperature samples. In samples stored at sub-zero temperatures, fGC metabolite concentrations showed a similar but dampened pattern, while fE metabolite concentrations exhibited small and variable changes with no consistent trend. We discuss explanations for the dynamic pattern of changing fecal metabolite concentrations and offer practical and analytical guidance to field workers for situations in which ideal conditions for stabilizing hormones are not available.

Life at the top: rank and stress in wild male baboons

Alpha males have higher stress levels than beta males in a wild baboon society. In social hierarchies, dominant individuals experience reproductive and health benefits, but the costs of social dominance remain a topic of debate. Prevailing hypotheses predict that higher-ranking males experience higher testosterone and glucocorticoid (stress hormone) levels than lower-ranking males when hierarchies are unstable but not otherwise. In this long-term study of rank-related stress in a natural population of savannah baboons (Papio cynocephalus), high-ranking males had higher testosterone and lower glucocorticoid levels than other males, regardless of hierarchy stability. The singular exception was for the highest-ranking (alpha) males, who exhibited both high testosterone and high glucocorticoid levels. In particular, alpha males exhibited much higher stress hormone levels than second-ranking (beta) males, suggesting that being at the very top may be more costly than previously thought.