Comments On: Baboon Mothers and Infants

Baboon Mothers and Infants by Jeanne Altman excerpts part 1
ONE OF THE FIRST things that strikes anyone who follows a group of baboons across the African savannahs is how much time and energy they spend just making a living: feeding, walking, avoiding predation. The surprise is particularly great for those who have watched captive monkeys, or even have seen films of wild primates, for these films are inevitably constructed from the 10 percent (or less) of the daytime that the animals spend socializing, not the 65 to 70 percent of the time they spend walking and feeding. One can easily overlook these activities of maintenance or production. In captivity such activities occupy only 10 to 20 percent of the day even in large enclosures or small islands with provisioning and typically involve nothing more than picking prepared food pellets out of a hopper. In the baboons' natural habitat, however, it is impossible to ignore these activities when animals walk several kilometers a day in the hot sun, when they laboriously dig bulbs or grass corms from the ground for much of the day, and when their rest periods seem necessary for recovery from fatigue, rather than just a response to boredom. Thus, in our studies of yellow baboons, Papio cynocephalus, in Amboseli National Park, Kenya, my colleagues and I have been led--no, virtually forced--by our animals to consider their lives as an integrated whole and not as composed of in dependent social and nonsocial pieces. In all likelihood, the fact that they spend three-quarters of their day making a living in itself affects their other activities; and not just in how much time they have avail able for those other activities, which is immediately obvious, but in much more complex ways as well. It is some of these complexities that I shall explore in the following analyses.
Two particularly crucial life stages for any primate are those of motherhood and infancy. Most field studies of wild primates have included qualitative descriptions of mothers and infants, and some provide quantitative results for a few behaviors or for a few typical individuals. In addition, quantitative studies of captive nonhuman primate mothers and infants have been conducted on a variety of species and have included a wide range of social settings, from isolation studies to those conducted with small groups of mixed ages and both sexes. Most of these studies have focused on infants rather than mothers and have emphasized predictions of adult behavior and effects of mother-infant separation as part of an attempt to provide a monkey model for human mother-infant separation. They have included both "purely" observational and more experimental investigations. However, with but rare exceptions, systematic quantitative studies of mothers and infants in the complexity of natural settings have been absent from research on both human and nonhuman primates, and studies of environmental influences have been limited primarily to a few elements in the social environment.
This study is an attempt to assess the nature and extent of external influences on baboon mothers and their young infants. My general goal was to identify and measure factors affecting survival and behavior during motherhood and infancy, and to identify likely ontogenetic origins of differences in adult behavior and life history patterns. Toward this end I collected systematic ecological, demographic, and behavioral data on all baboon mothers in a group in Amboseli National Park, Kenya, for which long-term life history data were available on individual members.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 2
The project had several specific goals generated by the overall aims. First, I sought an understanding of the demography of motherhood and infancy. Are these high-risk periods for baboons, as they are for other animals? To answer this, I turned to the demographic data that my colleagues and I collected from 1971 through 1978. Such information accumulates slowly, and David Post, Jane Scott, Sue Ann McCuskey, Francis Saigilo, Jeffrey Walters, David Stein, and especially Stuart Altmann and Glenn Hausfater have all contributed to its collection. From these data I was also able to examine the effects of maternal age, dominance rank, and infant gender on mortality and the effects of maternal dominance rank on infant gender.
Next, I sought a quantitative description of the nonsocial milieu or ecology as it affected mothers and infants and a test of the extent to which this ecology limits infant survival and the timing of births and of weaning. Further, I asked whether mothers' time budgets were affected by their dominance rank and, if so, whether these effects were reflected in differential survival or in the time or attention available for infant care.
Another set of related questions involved the social milieu. In what ways does a female's social life change when she gives birth? Which changes are beneficial and which harmful? What are the nature and sources of individual differences in social milieu among mothers? In the literature of behavioral biology, much attention has been directed to the relationship between male dominance rank and reproductive success yet for a number of primates female dominance ranks have been shown to be more stable both within and between generations. Few field studies identify either the onto genetic origins of female dominance relationships or their potential evolutionary consequences. Can we identify the origins of dominance rank "inheritance" in these early social interactions? Who are the individuals who constitute the social world of mothers and infants? Are they members of particular age classes? Are they particular individuals? Are the individual preferences in choice of associates observed in sexual consortships also manifested at this time?
With an understanding of the ecological and social milieu and the demographic (e.g., age, parity) and sociological (e.g., dominance rank) characteristics of the mothers, I could determine not only whether these variables affect each other, affect mothers directly, and affect infant mortality, but also whether they affect maternal care, the mother infant relationship, and infant development.
Recently, models of the evolution of parent-offspring relations, have been developed to explain phenomena such as weaning conflict and to make predictions about aspects of parental care that evolve through natural selection. Models of the evolution of behavior are economic models; their major parameters include costs and benefits of behavior, usually measured in the "currency" of fitness, of survival and reproduction, and of relative representation of genes in the next generation. Assuming a degree of heritability of the behavior in question, one is then concerned with attempts to estimate these costs and benefits. Does the behavior appear to be selectively advantageous or disadvantageous? The answer, of course, will not tell us whether any behavior evolved through natural selection. However, given certain assumptions about the population, the mating system, and the heritability of the traits involved, the answer will suggest whether the behavior could have spread at least partially through natural selection.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 3
This book is about motherhood and infancy more than about "mothers" and "infants": motherhood and infancy are only two stages, albeit particularly important ones, in the life histories of individuals whose lives extend before and, one hopes, after these periods. Females enter motherhood with their pasts and they and their infants carry into their futures the marks of their experiences during the life stage they so intimately share. Even during this period females' lives remain more complex and multifaceted than the label "mother" or "motherhood" implies. It is the pasts and the futures and the concurrent facets of their lives that I shall examine and relate to the experiences of motherhood and infancy.
In all aspects of the present study, one fact recurs: baboon mothers, like most primate mothers, including humans, are dual-career mothers in a complex ecological and social setting. They do not take care of their infants while isolated in small houses or cages as the rest of baboon life goes on. They are an integral part of that life and must continue to function within it. The baboon world affects them, and they it, throughout their lifetime. Determining the consequences of limited available time and the effects of maternity, social milieu, and physical environment on individuals' time budgets constitutes a major thrust of this investigation.
For short-lived animals the life history approach to the study of behavior is common. In studies of primates, including humans, it is much rarer, partially because life histories are so long and each stage is sufficiently complex to keep researchers themselves occupied for whole lifetimes. The work that follows is one, necessarily incomplete, attempt to combine a life history approach with the quantitative study of primate behavior in the field.
The general problems that face most human and nonhuman primate mothers are in many ways similar. High rates of infant mortality and appreciable maternal risk have probably been characteristic of most human and other primate populations. Rates of birth and death will determine the relative ages of siblings, affect the size of age cohorts and other aspects of the social group in which mothers and infants find themselves. Likewise, ecological and social factors that affect mortality will be important aspects of motherhood and infancy.
The complex ways in which members of society provide support and also transmit existing social structure to infants are important for any primate mother. If we can begin to understand the origins of normal differences in maternal behavior, we shall be better able to predict the consequences of changes in these factors.
By examining the problems of motherhood and infancy in detail for one primate species we may learn questions to ask, parameters that should be measured, and possible strategies for studying complex ecological and social relationships in related species. Just as the study of another human culture sometimes provides a different perspective, new insights, occasionally new answers, and more often new questions, I hope that this attempt to deal with the complexity of experience in a study of baboon motherhood and infancy will strike some familiar chords or stimulate some new ideas in those whose main interest is in another species, perhaps even our own.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 4
BABOONS (Papio spp.) are large, group-living, Old World monkeys, full-grown males weighing about 23 kilograms, females 11 to 12 kilograms. There are several species of baboons, exhibiting various social structures and occupying a wide variety of African habitats. Baboons are among the most terrestrial of the monkeys, a feature that makes them more observable than most other primates and has led also to attempts to regard them as models for hominid evolution.
Despite the widespread use of baboons in medical research, few behavioral studies have been conducted on members of the genus in captivity. Major early exceptions were Hans Kummer's study of hamadryas baboon (Papio hamadryas) social organization in the Basel Zoo and Thelma Rowell's investigations of captive olive baboons (P. anubis), particularly of female social relations and infant development. Terrence Anthoney (1968) and Gilbert Boese (1975) studied the Brookfield, Illinois, colony of guinea baboons (P. papio), and more recently Anthony Coelho, Claud Bramblett and colleagues have begun long-term studies of several Papio species and hybrids at the Southwest Foundation for Research and Education. In contrast, rhesus monkeys (Macaca mulatta), other macaque species, and squirrel monkeys (Saimiri sciureus) have been major subjects of behavioral investigations for many years, especially at the Japanese Primate Centre and in laboratories such as those of Bernstein, Harlow, Hinde, Jensen, Kaufman, Mason, Mitchell, Ploog, Rosenblum, and Sackett, where research has often focused on infant development and the ontogeny of behavior.
Field studies of the various Papio species have had a very different history. At the turn of the century Eugene Marais lived for several years with chacma baboons (P. ursinus) in South Africa. Only recently, many years after his death, was his unfinished manuscript found and published, providing a sometimes strange but intriguing view of baboon life. In the 1950s field studies began in earnest; research on chacma baboons by Bolwig and by Hall and on olive baboons by DeVore were the first of what have become a growing variety of investigations of the several baboon species. Kummer's field study of social organization of hamadryas baboons, Stolz and Saayman's of the behavior of chacma baboons, Thelma Rowell's field research on olive baboons, and the Altmanns' study of the ecology of yellow baboons all followed shortly after, in the 1960s. Most of these studies were general investigations producing qualitative or quantitative descriptions of aspects of ecology or behavior. An annotated bibliography of the early field studies has been published by Baldwin and Teleki (1972). The less closely related gelada baboon, Theropithecus gelada, has been the subject of research for a number of years in the Semien mountains of Ethiopia, particularly by Crook, the Dunbars, Kawai, Osawa, and Mori.
The 1970s have seen two major changes in the conduct of baboon field studies. First, research is more focused and problem oriented and is often concerned with hypothesis testing: following up on questions raised by the more general studies, by theoretical considerations (e.g., foraging and kin selection theory), or by laboratory investigations of other species. This change was foreshadowed in the second wave of studies in the 1960s and is partially paralleled by changes in field studies of other species. Second, long-term studies were initiated at several sites. These two changes, combined, are beginning to result in longitudinal, life history studies of known individuals and in-depth investigations that have involved researchers from a wide variety of disciplines. The sites with appreciable continuity include those of Kummer, Abbeglen, and colleagues with hamadryas baboons in Ethiopia; Ransom, Packer, and others with the olive baboons at Gombe in Tanzania; the Altmanns, Hausfater, and colleagues with the yellow baboons in Amboseli, Kenya; Harding, Strum, and others in Gilgil, and more recently Popp and DeVore at Mara, all with olive baboons in Kenya; W. J. Hamilton with chacmas in Botswana; and the Rasmussens and Rhine with yellow baboons in Mikumi, Tanzania.
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My daughter has to do a report on the Golden Monkey(not any kind of group but the "Golden Monkey". We are having a hard time finding any information. Is their another name for this type of monkey, or do you have an idea where we can go to find this information. Thanks
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Baboon Mothers and Infants by Jeanne Altman excerpts part 5
General Natural History
Savannah baboons are born into semiclosed groups of about 40 animals, but ranging from less than 10 to almost 200. In the wild, baboon females reach menarche at four-and-a-half to five years of age, and then before their first conception they experience a series of menstrual cycles for about a year ("adolescent sterility"). Their infants are born after a gestation period of six months (177 days). Postpartum amenorrhea follows, such that successive infants are produced at intervals of one-and-a-half to two years unless the process is accelerated by the death of the previous infant. Thus, after a female reaches full maturity, that is, experiences her first pregnancy, she spends approximately half her life with a dependent infant, almost a third of her life pregnant, and the remaining time undergoing menstrual cycling. This is true throughout adulthood; menopause has not been demonstrated for any nonhuman primate.
The Amboseli Population
The cynocephalus, or yellow, baboons include those we studied in the short-grass savannah region of southern Kenya just a few miles north of Mt. Kilimanjaro. There they inhabit Amboseli National Park along with a wide variety of other animals. Some of these, such as leopards, lions, and most recently hyenas, prey on baboons. The baboons themselves prey on others. These prey are usually grasshoppers and other invertebrates, but, they also include African hares, gazelles, and vervet monkeys, especially during the dry season. Most other large mammals in the area, such as elephants, wildebeest, and zebras, have an essentially neutral relationship with baboons, or an indirect one, through plants.
The Amboseli baboons have two sources of contact with humans other than the observers. First, Maasai pastoralists live within the animals' home range during the dry seasons. Although the Maasai seldom overtly interfere with most wildlife, their cattle constitute a large proportion of the Amboseli biomass during the dry seasons and compete with other grazers. Baboons, along with other wildlife, are displaced by cattle herds. Additionally, Maasai dogs and children occasionally chase baboons. Since late 1976, the Maasai have been excluded from Amboseli National Park in exchange for services and alternative resources, and have been provided with financial compensation for allowing wildlife to graze on Maasai land outside the park. Contact between baboon groups and Maasai and the effect of competition with Maasai cattle are now restricted to portions of the baboons' home range that are outside the park.
Second, tourists visit Amboseli. In recent years the number of tourists has greatly increased; but in contrast to the tourists of the early 1960s, these visitors are usually non-Kenyans and often on commercial tours. The result is that they spend little time in the park, almost all of it within a small area near the lodges, several miles from our main study area. Thus, members of our main study group currently seem to come in contact with tourists even less than study groups did when we first worked in Amboseli in the early 1960s.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 6
Western and van Praet (1973) have described the recent transformation of the Amboseli habitat from an acacia woodland to a dry habitat with salt-loving plants. Several years of heavy rainfall during the 1960s raised the water table to unusually high levels. The rising water led to the death of most of the trees in the acacia woodland. Some trees died as a result of salts, which, through "wicking action," form a layer in the soil just above the water table and are carried up into the root zone of the trees. The roots of other trees were completely drowned. Overgrazing by large herds of Maasai livestock reduced available grazing material, and this depletion, in turn, probably resulted in a shift in diet of animals that are both browsers and grazers, most notably elephants, and led to further damage to trees. Overgrazing was aggravated by poor rainfall starting with a drought in 1969.
As a result, major declines occurred in populations of many species of wildlife that depend on woodland habitats, including the vervet monkeys. Over two thousand baboons were censused in the Amboseli waterhole area during 1963-64, but by 1971 the baboon population in the same area was only about 10 percent that size. The percentage decline in the baboon population was considerably greater than that of the vervets (90 percent versus 33 percent), at least partially owing to the trapping of approximately 230 baboons in 1965 by the Southwest Foundation for Research and Education; also probably partially owing to the presence of a paralytic virus in the population in 1964.
In 1971 we began a general program of periodic monitoring of the baboon population as a whole and longitudinal monitoring of demographic events in a single study group. A detailed analysis of the baboon population since 1971 is under way but the most relevant demographic data for the subjects of the longitudinal study, Alto's Group, are described here.
Alto's Group
The subjects of this study and of most of our detailed longitudinal observations are members of Alto's Group, one of five groups now occupying the central waterhole area of Amboseli. Longitudinal data on individuals in Alto's Group have been collected since 1971 by a succession of field workers. In the fall of 1972, a fusion was completed between the original Alto's Group and a one-male group, High Tail's Group.
Of the 35 members of Alto's Group in July 1971 and the 15 members of High Tail's Group when they were first censused in November 1971, 23 from Alto's Group and 8 from High Tail's Group remained in the combined Alto's Group of 47 members four years later, at the onset of this study in July of 1975. Thus there was approximately a 40 percent turnover of group members from 1971 to 1975. This turnover was due to births, deaths, and migrations. Of course, even more such demographic events occurred during the intervening years than are indicated by these figures, since many individuals were born and died, joined the group and then either died or emigrated, and so on. Consider Dogo, an infant born to female Preg in August 1971. He has grown up in an ever-changing social group and has known approximately 68 animals, while staying in his natal group of about 45 animals. None of his cohort from 1971-72 remained in Alto's Group in 1975; yet almost all of his younger sister's 1973 cohort did. Most of those who were juveniles when he was born were still in the group in 1975, though some males had temporarily emigrated and then returned. Of the eight males who were fully adult when Dogo was born, only three, Peter, Max, and Stubby, remained. In January 1979, Dogo was just reaching adulthood when he was killed by two hyenas. At the time of his death, he was the oldest animal that we knew from birth. His mother and two younger sisters remained in the group; a younger brother, Pedro, one of the subjects of the current mother-infant study, died when eight months old.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 7
In the results of this study I document a more complicated picture of motherhood and infancy than has usually been considered. Yet I still deal primarily with only one year and one reproductive stage in the lives of adult females: their last month of a pregnancy and the first year of their infants' lives. Occasionally I shall hint at the longer view: relationships established before infants' birth, bonds that persist for infants as they mature, lifetime dominance relations, and long-term ecological patterns. Many of these are currently topics of Amboseli research conducted by Stuart Altmann, Glenn Hausfater, David Post, David Stein, and Jeffrey Walters. However, the continuities and the long-term bonds will also be set against the ever-changing demographic panorama.
Each year approximately ten infants are born into the group of about 45 animals. Usually six or seven infants survive the first year of life. In recent years, Alto's Group has remained at a fairly stable population level, and on the basis of age-specific birth and death rates, we predict that this stability will continue. These were essentially the demographic conditions during the study of mothers and their newborn infants that I conducted from July 1975 through July 1976, during October 1976, and during a short pilot study in 1974.
The Mother-Infant Study
All females in Alto's Group who had infants under a year of age were included in the study. Except for one adult female, Lulu, who has not conceived since at least 1971, probably not since 1969, all adult females in Alto's Group had at least one infant during this study, including four primiparous females and two females who experienced their first viable delivery during this period. Infant maturation and mortality factors resulted in three females being included with two successive infants. Five infants who were included were born before July 1975. Thirteen others were born during the study.
When I arrived in Amboseli at the end of June 1975, several infants were already in the group. Information about them was kindly provided by David Post and Jane Scott. The oldest was Pooh, a scrawny eight-month-old female with locomotor impairment who was Plum's first-born. Several other infants were born about the same time as Pooh and would have been included in the study if they had survived. Slinky's first infant was stillborn in November. Brush's first infant was born in October with congenital deformities; it lived only a few weeks. A high-ranking older female, T.T., had given birth to a healthy male infant, Tom, within a week of Pooh's birth. I was quite disappointed when we learned of T.T.'s and Tom's sudden disappearance in May.
Four females gave birth within two weeks at the end of 1974, when Pooh was about two months old. Alice was born to high-ranking, elderly Alto, who had two other known surviving daughters in the group, Spot and Dotty. Another high-ranking older female, Mom, also gave birth to a daughter, Softy at that time, but Softy died of unknown causes after only a few weeks. Mid-ranking Oval and Fem both gave birth to sons, Ozzie and Fred. Oval's juvenile daughter, Fanny, was in the group. Being Fem's first infant, Fred had only one "known" relative, his mother's putative sister, Gin.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 8
In addition to Pooh, Alice, Ozzie, and Fred, two-month-old Eno was already in the group when I arrived. Early one evening in late April she was born to Este, an elderly low-ranking female who had a five-year-old son, Toto, in the group when Eno was born. We had expected Eno to have a close peer, because Judy, another elderly low-ranking female, was due to give birth in March. However, Judy temporarily disappeared from the group at about that time, and was wounded and apparently experienced a stillbirth before her return. At the end of June Slinky gave birth to her second infant. Although it apparently was well formed externally, it was rigid at birth and died two days later.
In July Pooh, Alice, Ozzie, Fred, and Eno were joined by Summer and Pedro, offspring of mid-ranking Scar and Preg, who already had surviving offspring in the group. In August, Misty and Bristle were born, within a few days of each other. Misty was the daughter of high-ranking Mom. Bristle was Brush's son, and since Brush's first infant had died so early and had been malformed, it was almost as if Bristle was her first-born. The fact that Brush was very low-ranking made a striking contrast in the experiences of these two infants, one that had not been so evident with the more closely matched Summer and Pedro. Misty's death just after she turned two months old was a particularly unfortunate one.
Just before Misty's death, Handle, a low-ranking female, gave birth to her first infant, Hans. Hans's environment was quite similar to that of Bristle. Since their mothers were also rather frequent associates and since no other infants were born until late January, these two infants spent much time together.
Starting in late January, four infants were born within a month of each other. Gin gave birth to her first infant, Grendel. A few days later, Slinky finally gave birth to a viable infant, although Sesame appeared weak from birth and had bouts of illness periodically thereafter. At the end of February, Judy again gave birth, this time to a healthy son, Juma. Judy's daughter, Janet, was almost three years old by then. A few days later, Alto's daughter, Spot, also high-ranking, gave birth to female Safi. With two sisters and a grandmother in the group, Safi had more known relatives than any of the other infants.
Several months passed. One night during this period Judy and Juma disappeared and did not return. Then at the beginning of June, Mom gave birth to Moshi. I was struck with how similar Moshi's world was to that of his sister, Misty, and how different it was from that of infants such as Bristle who had been born at the same time as Misty. Similar contrasts were evident among the other infants. I shall explore the nature and origins of some of these differences in the chapters that follow. Often Bristle and Moshi will provide useful examples of contrast.
Just after I left Amboseli at the end of July, Ozzie's younger sister, Oreo, was born. When I returned to Amboseli for a month that October I was able to observe Oreo as well as Vicki and Peach, who were born at the beginning of October. Despite over a year of observations on new mothers, including the first day of life for most of the infants, had not actually been present for a birth. It was a particularly exciting surprise when the last birth, that of Plum's infant, Peach, occurred while I was making observations one afternoon, and when Plum, the most wary female in the group, tolerated my presence at a moderate distance, enabling me to watch most of the labor and birth.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 9
BECAUSE MOTHERS WITH NEONATES appear to be the class of individuals in our baboon population that are most sensitive to being observed, and because most of our previous research had been done from atop a vehicle, the first month of this study and to some extent the second were partially devoted to accommodating the females to observation on foot within 5 to 10 meters. Much closer distances were possible for most mothers, but consistent with the aim to minimize our effect on the system we were studying, I always tried to stay at a distance that would not discourage the mothers' interactions with the most sensitive individuals in the group. In addition, some types of detailed data were considered sufficiently reliable to use only after several weeks or more of systematic sampling. Consequently, for some analyses I used data obtained from July 1975 onward; for others, only data obtained later.
After various sampling schemes were tried, the following scheme was established by September 1975. Each female was sampled during the last month of pregnancy for two days at least a week apart, on the day of birth and the fifth day of infant life, on two days during the infant's second week, and one day per week thereafter until the infant was six months old. During the next six months samples were taken two or three days per month. Whenever impassable mud, illness, or other factors reduced available observation days, sampling mothers with older infants was sacrificed in favor of sampling those with younger ones.
In gathering behavioral data I sought samples that would provide unbiased estimates of rates of behavior, of time budgets and bout durations, and other measures that are derived from these.
Focal-animal (continuous) sampling and instantaneous (point) sampling are the most versatile and suitable of the common techniques used to provide unbiased estimates of time budgets; focal samples provide the data needed to estimate rates and bout durations. Unfortunately, until recently most field studies have utilized ad libitum sampling and many laboratory studies one-zero sampling, neither of which provides unbiased estimates of these parameters. Detailed comparisons of bouts, rates, time budgets, and derived measures within and between studies that utilize these techniques cannot be made unless the sampling scheme itself is considered as a possible source of the differences and similarities found in the results. Consequently, I shall primarily restrict detailed comparisons between the results of this study and others to those comparisons not likely to be confounded by sampling biases. More generally, I shall emphasize where possible, comparisons with those studies that differ in the fewest major variables, such as species and extremes of caging and rearing conditions, so that similarities or differences can be at least provisionally attributed to characteristics of the animals in their normal range of living conditions.
On each sample day, I sampled the behavior of two females. They were sampled alternately, each for 15 minutes in-sight time (or 20 minutes in the first months of the study) out of every hour from 0800 through the 1700 hour but excluding the 1200 hour. The females that were sampled on any one day were paired on the basis of proximity in age of their infants. Such pairing provided the possibility of control for variability in factors such as day journey length or unusual daily events and enabled me to sample females more frequently than if I had followed only one female per day. Alternating between two females did have the disadvantage that more time was devoted to searching for individuals and probably more of the 15-minute samples were missed altogether than would have been the case if I had stayed with just one female each day.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 9
BECAUSE MOTHERS WITH NEONATES appear to be the class of individuals in our baboon population that are most sensitive to being observed, and because most of our previous research had been done from atop a vehicle, the first month of this study and to some extent the second were partially devoted to accommodating the females to observation on foot within 5 to 10 meters. Much closer distances were possible for most mothers, but consistent with the aim to minimize our effect on the system we were studying, I always tried to stay at a distance that would not discourage the mothers' interactions with the most sensitive individuals in the group. In addition, some types of detailed data were considered sufficiently reliable to use only after several weeks or more of systematic sampling. Consequently, for some analyses I used data obtained from July 1975 onward; for others, only data obtained later.
After various sampling schemes were tried, the following scheme was established by September 1975. Each female was sampled during the last month of pregnancy for two days at least a week apart, on the day of birth and the fifth day of infant life, on two days during the infant's second week, and one day per week thereafter until the infant was six months old. During the next six months samples were taken two or three days per month. Whenever impassable mud, illness, or other factors reduced available observation days, sampling mothers with older infants was sacrificed in favor of sampling those with younger ones.
In gathering behavioral data I sought samples that would provide unbiased estimates of rates of behavior, of time budgets and bout durations, and other measures that are derived from these.
Focal-animal (continuous) sampling and instantaneous (point) sampling are the most versatile and suitable of the common techniques used to provide unbiased estimates of time budgets; focal samples provide the data needed to estimate rates and bout durations. Unfortunately, until recently most field studies have utilized ad libitum sampling and many laboratory studies one-zero sampling, neither of which provides unbiased estimates of these parameters. Detailed comparisons of bouts, rates, time budgets, and derived measures within and between studies that utilize these techniques cannot be made unless the sampling scheme itself is considered as a possible source of the differences and similarities found in the results. Consequently, I shall primarily restrict detailed comparisons between the results of this study and others to those comparisons not likely to be confounded by sampling biases. More generally, I shall emphasize where possible, comparisons with those studies that differ in the fewest major variables, such as species and extremes of caging and rearing conditions, so that similarities or differences can be at least provisionally attributed to characteristics of the animals in their normal range of living conditions.
On each sample day, I sampled the behavior of two females. They were sampled alternately, each for 15 minutes in-sight time (or 20 minutes in the first months of the study) out of every hour from 0800 through the 1700 hour but excluding the 1200 hour. The females that were sampled on any one day were paired on the basis of proximity in age of their infants. Such pairing provided the possibility of control for variability in factors such as day journey length or unusual daily events and enabled me to sample females more frequently than if I had followed only one female per day. Alternating between two females did have the disadvantage that more time was devoted to searching for individuals and probably more of the 15-minute samples were missed altogether than would have been the case if I had stayed with just one female each day.
Brad and Trouble
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Baboon Mothers and Infants by Jeanne Altman excerpts part 10
AGE DIFFERENCES BETWEEN SIBLINGS, the size and age-sex composition of available play groups, and other important features of the social milieu of mothers and infants are determined by the demographic patterns of a populationùage-specific rates of natality and mortality, migration, and so on. Mortality figures in particular provide clues to stressful life stages and are important in considering costs and benefits of various social behaviors within an evolutionary framework.
Survival of Infants, Juveniles, and Adult Females Of the 13 infants who were born during the study, 3 (Pedro, Misty, Vicki) died during the study after varying periods of illness, 1 (Juma) suddenly disappeared overnight with his mother during the study, 1 (Safi) became seriously ill and recovered, and 3 (Sesame, Moshi, and Peach) died after the study but during the first two years of life, that is, as infants (first 12 months) or in the beginning of the juvenile period. Six (Summer, Bristle, Hans, Grendel, Safi, and Oreo) survived the first two years of life, 2 of them (Bristle, Oreo) as orphans. Of the 5 infants who were included in the study but who were born before July 1975 (i.e., had already survived part of the first year when the mother-infant study began), 4 (Alice, Ozzie, Fred, Eno) survived through the second year of life and 1 (Pooh) died just before her second birthday. Alice was orphaned before her second birthday; Ozzie after his.
Only rarely can we determine either the immediate cause of death or an earlier precipitating cause. When an apparently healthy female or young juvenile disappears from the sleeping grove overnight we presume not only that it died but that it was probably taken by a. However, in the case of older juvenile, subadult, and adult males even overnight disappearances can be due to migration rather than death, a fact that makes it difficult even to construct life tables for these age-sex classes. Determination of an infant's death is somewhat easier because if it is survived by its mother, she usually carries the corpse for several days. Of the infants in the 1975-76 study Peach and Misty apparently died of disease; probably Pedro and Juma did as well, along with Juma's mother, Judy. Vicki's death was probably due to poor mothering, and Pooh's to the severe wounds inflicted by male Even just before her second birthday. Both Moshi and Sesame showed signs of possible nutritional deficiencies for at least a year before they died. Sesame disappeared just before her second birthday and soon after the birth of her mother's next infant. Moshi fell from a tree when his younger brother was a few months old The cause of his fall is unknown. Alice's mother, Alto, disappeared while we were away from the group for several days. When we first identified Alto in 1969, she appeared to be old, probably at least 15 years of age; during the last months of her life she seemed to become less interactive and to rest more.
Mortality patterns during the year of the mother-infant study were similar to those for the whole seven-year period of the longitudinal study; we can analyze the latter in greater detail because of the larger sample sizes. Sample sizes are now about twice those available in our earlier paper on infant survivorship, and data are now available for years two through five, taking females through the age of menarche and males almost to the beginning of subadulthood. In addition, the mortality data considered here come entirely from a period of fairly stable demographic parameters. No trends in survival could be detected when I examined infant mortality within Alto's Group from each year separately, and I have therefore combined the data since 1971 to provide more adequate sample sizes for the analyses that follow.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 11
Several features of the survival data are quite striking. Mortality is appreciable (.28) in the first year of life and only slightly less so (.25) in the second year. However, it is essentially zero in the next three years. (One juvenile male, Kub, either died or migrated in the fourth year.) Baboon mothers provide little care during an offspring's second year of life, often none during the latter part of that year. The low survival rates during the second year of life suggest that a prolongation of maternal I care might potentially be of considerable benefit to these offspring and I increase their rates of survival. By contrast, in the third, fourth, and fifth years, which are still prereproductive years, the survival rate is virtually 100 percent. Consequently, it seems unlikely that directing any additional care or "altruistic" behaviors of immediate consequence toward these older juveniles would result in a selective advantage: there is no potential survival difference that could provide the selective basis for the evolution of such behaviors. However, behaviors of long-term or delayed consequence, such as those affecting age of menarche, ability to obtain mating partners, or ability to learn parental skills, may be of advantage to this age group.
In early research , the period of infancy was considered to be the first year of life. However, from our longitudinal study we have since found that individuals identified as older infants by physical appearance and by the fact that they are still suckling and are associated with a single adult female, are up to 16 months of age. Baboons are not strictly seasonal breeders, interbirth intervals are individually variable, and the development of independence is a long, gradual process that extends into the second year of life. Thus it is difficult and somewhat arbitrary to define the age limits of infancy. All of the first year of life is clearly a period of dependence and infancy. This study focuses on that period and it is individuals of that age that I refer to as infants. However, considerations such as the mortality data above and mothers' future reproduction demonstrate the importance of the second year of life, and where possible, I extend my discussion to this period.
Consider next mortality risks for females during adulthood. In the seven years since mid-1971, we have data for approximately 78 female-years of exposure for fully mature females, defined as those who have conceived at least once. Nine fully adult females died in Alto's Group during the period from September 1971 to September 1978, for a mortality rate of .12 per annum. In most mammals mortality risk is low in the early years of adulthood, but rises, often rather sharply, in later years. Is mortality risk a function of age among adult female baboons? In particular, is the mortality risk to mothers age-related? Young adult females (taken here as those who have conceived at least once but who were within three years of first conception, that is, those who were approximately six to nine years old) had an annual death rate of .04; older ones (defined as those who were at least three years past first conception) a rate of .16. Adult females usually spend approximately one-half of their adult lives with a dependent infant under a year old the proportion is slightly lower if the infant dies, higher if the infant survives. Thus, one would expect mothers of infants to account for half of the deaths among adult females. In fact, six of the nine females who died had infants under a year of age at the time of death, for a rate of .15 deaths per annum for females with an infant. The rate for females without an infant was .08 per annum. Of these remaining three deaths, one was a young female (Brush) experiencing her third pregnancy. The other two were much older females who were experiencing abnormal reproductive cycles: in Jane's case, a particularly long cycling time without becoming pregnant, and in Alto's an unusually long postpartum amenorrhea. Perhaps their deaths resulted from or were made more likely by pathology or old age. The tentative conclusion is that in this population reproduction exacts a substantial cost in the form of increased mortality.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 12
It is reasonable to expect that two additional variables might affect the survival of mothers or infants: maternal dominance rank and infant gender. seven years, sex ratios at birth have been equal (26 males :26 females, plus two unsexed stillbirths and one unsexed case of neonatal death during late 1972). Infant (first-year) mortality, including stillbirths, was higher for male infants than for female infants. This difference does not approach statistical significance, and was due almost entirely to the fact that four of the male infants and only one of the female infants died at the same time that their mothers did. We cannot yet tell whether this latter result is an artifact or whether, for some reason, mothers of male infants are more vulnerable. Although these mothers were all older mothers, older mothers, like younger ones, produced infants at about a 1 :1 sex ratio. For humans, sex ratios at birth are slightly skewed toward males and become less so for older mothers or those of higher parity, but a sharp skewing in favor of males occurs for quite old mothers. We do not yet have sample sizes 18 for the baboons adequate to detect differences of the small magnitude found in humans.
It is generally assumed that female dominance rank is correlated with reproductive success. Although this has not yet been demonstrated for savannah baboons, there is some positive evidence for provisioned macaques and perhaps - for geladas. Female offspring assume their mother's relative dominance rank for life, whereas male offspring repeatedly change rank during adulthood and seem to be less affected, as adults, by their mother's rank, as has been reported for macaques.
As predicted on the basis of the behavioral life history results described above, in Alto's Group dominance rank is related to sex of off spring: low-ranking mothers produce more male than female offspring, high-ranking ones producing primarily female offspring. We can make a dichotomous characterization of these mothers as high- or low-ranking. Then we can also make a dichotomous characterization on the basis of whether a female had more male or more female offspring, considering each mother as the sampling unit, rather than each birth, in case there is a lack of independence of sex among offspring of the same female. The results (omitting ties) are that five or seven high-ranking females had more female than male infants, and five of seven low-ranking females had more male than female infants. Alternatively, if we do consider each birth as an independent sample and incorporate the two rank changes, we see that 10 of 29 infants born to high-ranking females were male, but 15 of 22 infants born to low-ranking females were male. Although I know of no literature on the possible mechanism leading to this result, physiological differences due to differences in stress levels could perhaps result in differential sperm survival in the two groups. The observed sex difference of offspring would appear to be the best strategy for each type of female, if female dominance rank is correlated with reproductive success, with female offspring of high-ranking mothers retaining their mother's rank and male offspring of low-ranking females being "freed" of their mother's rank. The relationship between maternal rank and sex of offspring has been modeled by Trivers and Willard (1973), and the model I am proposing for the observed baboon sex ratios is in the same spirit. However, this social and reproductive system requires a different set of assumptions than those used by Trivers and Willard; and therefore different specific predictions result.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 13
From the results that survival rates were as good or better for female (versus male) infants, and that high-ranking females tended to produce female offspring, one would expect to find that offspring of high-ranking females have lower mortality rates than those of lower ranking females. However, rates of infant mortality were slightly but not significantly lower for infants of low-ranking mothers (6 out of 20 = .30) than for infants of high-ranking mothers (9 out of 23 = .39). Thus despite the facts that low-ranking mothers produce more male than female offspring, high-ranking ones produce primarily female off spring, and female infants have survival rates as high as or higher than those of male infants, infants of high-ranking mothers have no higher, and perhaps somewhat lower, survival rates. This contra intuitive result for infant survival rates contradicts the limited available primate data on reproductive success from two provisioned rhesus colonies, as well as data on survival of human infants in different socioeconomic classes . However, even though infant survival does not appear to be positively correlated with dominance rank, other factors affecting the female's reproductive success, such as age of menarche and length of interbirth interval, may be rank related in our population. Only future study will clarify this puzzling situation.
When we combine the various pieces of information on maternal and infant mortality, the suggestion emerges that young adult females, essentially those with their first or second infant, are about as likely to have that infant survive and are more likely to survive themselves than are older females. Older females are more likely than are younger ones to die during the period of infant care, in which case the infants will also die. However, if these older mothers survive, their infants are also very likely to survive. Thus, their fate, more than that of young mothers, is closely linked to that of their infants during the first year.
Perhaps a cautionary reminder is necessary at this point. Some of the foregoing results have been obtained by partitioning a small data set, resulting of course in smaller subsets. Also there are several factors that may lead to differential exposure to the various reproductive stages, including the very results just discussed. There is at present no way to remove such confounding completely. Patterns will, one hopes, become clearer as more data accumulate. Sample sizes are not yet adequate to confirm statistically even appreciable differences.
Not only do infants affect their mothers' survival chances, but infants additionally affect demographic processes in a primate group in general and their mothers' reproductive success in particular through the direct effect that infants have on their mothers' future reproduction. Mothers of surviving Amboseli infants experience approximately 12 months of postpartum amenorrhea and then take an average of four cycles to conceive, whereas infant death results in resumption of cycles within one month of the death and conception after only one or two cycles on the average. It has been further suggested that mothers could improve their reproductive success if they reduced infant care, for example, by weaning their infants, thereby reducing the length of postpartum amenorrhea. While this has certainly been the case among humans in developing countries in recent years, such a result is dependent on (1 ) infant mortality rates not being appreciably increased by early weaning and (2) the mortality risk of childbirth and early stages of infant care being low. Weaning foods and even a semblance of modern medicine are probably sufficient to satisfy these conditions in developing countries. It is unlikely, however, that these conditions prevailed until recently for humans, or that they are satisfied in most animal habitats, including that of the Amboseli baboons. It is quite possible that in the absence of such advantages of early weaning females obtain higher reproductive success by engaging in fairly long periods of infant care than they would by reducing the period spent caring for a current infant in order to reduce the interbirth interval.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 14
A per annum mortality rate of almost .30 occurs during each of the first two years of life, with mortality dropping to virtually zero in the several years thereafter. Thus infancy and the following year are particularly crucial and difficult periods. The data also suggest that older adult females suffer higher mortality rates than do younger ones. This general pattern is the common mammalian one only the sharp discontinuity at age two may be unusual. Dittus' study of Macaca sinica (1975, 1977) provides the only comparable mortality data for a non-expending natural primate population. His data exhibit the same U-shaped distribution. Dittus' life-table data come from one cross-sectional census of many troops. Age was estimated by physical appearance, for immatures by making use of the short, discrete birth season in this species, and by comparisons with longitudinal changes in a smaller set of known individuals studied for several years. The life table data that Dittus presents (1975:132-138) agree in general with those reported here, with high rates of mortality for infants and for older adults, low rates for older juveniles and young adults. The one striking difference that merits comment here is the much higher mortality rate Dittus reports for young female versus male infants at his site in Polonnaruwa. However, this finding is based on the assumption that the 1970-71 sex ratio at birth was 1 :1. Although known births for four years averaged a 1 :1 ratio, the ratio reported for the Polonnaruwa region for 1970-71 is 21 :14, or 3:2, essentially the same ratio found in the infant-2 (older infant) class during the census. Thus it is not clear from the available data that differential female mortality occurs among the infants of that population. Subsequent data from the longitudinal project there should clarify this issue.
Additionally, in Amboseli there is a cost of reproduction. That is, females, especially older females, with dependent infants tend to suffer higher mortality rates than do younger ones or those in other stages of the reproductive cycle. I know of no published data on the cost of reproduction in other wild primate populations. Infants may impose an additional cost on their mothers' future reproduction due to an extended period of postpartum sterility, but this will depend on the risks mothers and infants would incur as a result of early weaning. Thus mortality risks are high for both mothers and infants. In the following chapters I shall attempt to identify those ecological and social factors that shape the first two years of life and in particular those that may enhance or hinder the survival of mothers, infants, and young juveniles and affect their interactions.*
* Deaths of adult females (four) and births (six to multipara and 5 to primipara) from 1 August 1978 through 31 December 1979 provide increased evidence for (1) the susceptibility of adult females to death when they have infants and (2) the relationship between maternal dominance rank and infant gender.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 15
IN RECENT YEARS time and energy have figured prominently in minimization or maximization models in behavioral ecology. However, the importance of time may be much more general than is suggested by minimization models. All of an animal's basic activities require time to perform; some of them require considerable amounts of time. Moreover, time is quite limited and all activities must be carried out within a fixed number of hours: baboons spend all 10 of the dark nighttime hours resting and sleeping in the trees, plus 3 to 4 additional late-evening and early-morning hours. Ten to 11 hours remain. I suggested previously that these daytime hours are filled to a considerable extent by maintenance activities. As a key to understanding the demographic results of the preceding chapter and the constraints within which mothers live, l shall now consider these daytime time budgets in detail and the ways in which the presence of a dependent infant influences a mother's productive activities.
Seasonality of Time Budgets
The Amboseli habitat is subject to considerable seasonal variability. As expected from its latitude (2 40'S), there is little variability in day length and temperature range, but seasonal variability in rainfall is considerable. The period from May or June through October is usually characterized as the long dry season, November and December as the short rains, January and perhaps February as the in terrains, and March through April or May as the long rains. Despite some variability from year to year, this basic pattern persists in Amboseli. During the dry season the baboons depend on the few permanent water holes in the central area of the park. When the rainy season arrives, however, temporary rain pools are available throughout their home range. At that time most large mammals in the area migrate out of the park, while others, such as the baboons, alter their habitat usage and movements within the same basic area. During 1975-76 the distance : traveled by the baboons was greatest (almost 6 kilometers daily) in December, January, and September. The values were lowest during the rainy season (4 to 4.5 kilometers). Time spent walking showed a some-what similar pattern, with highs of over of 17 to 19 percent in August, February, and March. Variability in feeding time was even greater, with 52 to 55 percent of the time spent feeding during June through November (the rains arrived in late November in 1975) and averages of only 40 to 42 percent during December, January, and May, with a gradual rise after January but a very sharp rise from May to June and for the rest of the dry season.
The effect of rainfall patterns on the group movements and on the percentage of time the baboons spend feeding and walking is a complicated one. Baboons are, and their various foods are distributed differently throughout their habitat and have different seasonal growth patterns. Particular food plants in particular parts of the animals' habitat leaf or fruit at various times after the onset of the rains. Then, too, some of these plants fruit annually, a month or two after the onset of one of the rains, whereas other plants, such as the grasses, develop lush new growth soon after the onset of each rainy season. Thus, in a linear regression, monthly rainfall accounted for less than 20 percent of the daily variance in distance traveled, and distance traveled or the time spent walking were no better as predictors of the time spent feeding. However, the average distance traveled by the group in a month accounted for 75 percent of the variance in the average time mothers spent walking that month. Also the sum of the rainfall in a month plus that for the previous month accounted for 50 percent of the variance in average monthly time spent feeding. That is, cumulative rainfall was a good predictor of the time an individual spent feeding, and the length of the group's day journey was a good predictor of the time an individual spent walking.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 16
The patterns of seasonal variability in day journey length and in the percentage of time spent feeding and walking are essentially the same as those found for 1963-64 for day journey length. Thus it seems reasonable to assume that these environmental conditions represent a pattern that is repeatedly encountered by the baboons from one year to the next, with relatively small annual variations, and that these conditions may be of considerable importance to the energetic requirements of lactating females, to the development of infant independence, and to the survival of both mothers and infants.
Synchrony of Activities within the Group
To what extent is the length of the group's day journey o; the amount of time an animal spends feeding determined by any individual and to what extent are an individual's activities determined by those of other group members or the group as a whole? We can assume that neither any individual nor the group as a whole affects short term rainfall patterns. Yet there remains some amount of flexibility in the amount of time spent feeding, and an individual may influence day journey lengths as well as vice versa. Survival differences may well depend on the extent to which environmental and social constraints combine with reproductive ones to place more or less burden on a female. Likewise, the degree of an individual's ability to affect these variables herself may well make a crucial difference to her survival.
Probably of particular importance to new mothers is the length of the day journey on the infant's first days of life. The infant often has trouble clinging during these first few days, sometimes requiring its mother to walk three-legged while pressing the infant against her ventrum with one hand, or to perform frequent repositioning during long marches. Especially on the infant's day of birth, the mother seems quite tired. This is reflected in her time budget for that day, during which she spends much more time resting than usual, and in the lack of concordance between her activities and those of other group members, discussed below.
Day journeys on days of birth were not likely to be shorter than average for the month, despite the mother's fatigue (6/10 lower than the mean, 5/9 lower than the median). Somewhat more of an effect is seen by comparing mean day journey length for the week before the day of birth with that of the week beginning with a birth day. The magnitude of the differences was small, but the direction of the effect was such that in six of eight cases for which had these data the distance traveled was shorter in the week following infant birth than in the previous week. It remains for further study to determine whether this slight effect is in some sense real (would reach statistical significance with somewhat larger samples) and if so, whether the magnitude is sufficient to be of any benefit to mothers. New mothers are sometimes seen dragging along at the rear of the group and thereby may be exerting a slight effect on the route, but there is no evidence that the group makes appreciable alterations in its activities to accommodate new mothers. If anything, during the early months of infant independence mothers probably spend more time walking than do other group members, because they spend time retrieving their infants.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 17
I expected that the percentage of time spent feeding, or feeding plus walking, would be a function of maternal dominance rank, with low-ranking females spending more time in these activities than do high-ranking ones. This was predicted on the assumption that low-ranking females would more frequently be displaced from high-quality foods and would therefore need more time to obtain an adequate amount of food. There was no such significant effect on feeding time, nor was there in Post's limited data on this topic. If a small but consistent effect does exist, the data are not yet adequate to detect it. Dominance effects may occur only during the dry season. The data from this study could not be analyzed by season owing to the confounding effects of infant age. This remains a problem for future research..
In approximately 90 percent of the samples the focal animal's predominant activity was the same as that of the group. Almost all deviation from that 90 percent figure occurred during the first month of infant life, when there was only 80 percent concordance. Most of this deviation, in turn, occurred on day one of infant life, when there was less than 60 percent agreement. This lack of concordance was due to the fact that mothers fed much less on day one and rested much more. I do not know whether the lower amount of time spent feeding on that day is the result of fatigue and of the difficulty in feeding due to the need to use one arm much of the time for infant support, or whether the afterbirth, which is eaten, supplies sufficient nutrients to allow less feeding that day. On the fifth day of infant life, residual discordance was still produced, partially by new mothers' doing more resting but also by their socializing more, as was true for the rest of month one. The rare instances of lack of concordance in later months were not consistently due to any particular activities.
Effects of Infants on Mothers' Feeding Patterns
In Amboseli the baboons feed while in either a seated or a quadrupedal standing position. Whereas catching insects and feeding on berries are probably more easily done from a standing position, two handed feeding is done primarily while sitting and may be advantageous for obtaining some foods, such as grass corms. Although it also appears that feeding while seated is more relaxing and energetically economical, the animals frequently have to move from one food patch to another. Some of these movements are caused by other group members, others by depletion of the present food patch. In either case, the seated animal must stand again with each move. However, during the first few months of an infant's life, if its mother is seated the infant can maintain contact and even doze without clinging and having to support its weight. After many hours of riding on the mother's ventrum, very young infants often tire and have trouble clinging. During the second month the infant can explore with considerable safety within its seated mother's ventral flexure. By the third month an active infant in her ventrum clearly hinders a mother's feeding attempts, and I often observed a mother embrace her infant to passive clinging, push it gently outside her ventrum (and just out of contact but nearby), or stand and feed with the infant exploring under her torso. Mothers of infants that were over five months old stood while feeding slightly more often than they sat; mothers of three- to five-month-olds were about as likely to sit as to stand, with considerable variety of direction from month to month and individual to individual. However, during the first month of infant life all nine mothers sat while feeding more than they stood and eight of nine mothers did so during month two. There were no differences on the basis of maternal dominance rank. Thus a mother's feeding positions seem to be an accommodation to her infant's needs during the first months, but gradually she prevents her infant from hindering her feeding by modifying her infant's behavior and sometimes modifying her own. By month four infants are disproportionately out of contact when their mothers feed.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 18
Maternal Time Budgets and Infant Gender
I can discern no significant effect of infant gender on the time mothers spend feeding or feeding plus walking. However, during all but one month mothers of female infants averaged slightly but insignificantly more feeding time than mothers of male infants. A gender effect resulting in more time spent feeding by mothers of male infants would be expected primarily if male infants grow faster than females. (Adult male baboons are about twice the weight of females.) A differential in the opposite direction would be predicted if males became nutritionally independent on their mothers earlier. The literature is contradictory on the former point for the first year of life for baboon and macaque infants kept in laboratories. Snow's data (1967) indicate no sex difference in growth rates until the end of the second year. Yet for rhesus macaques, Macaca mulatta, and another baboon colony there are indications that differences in growth rates may exist earlier, with male infants gaining weight more rapidly. During the first year of life, the infants in this study exhibited no consistent visible size differences: female Alice and male Ozzie were approximately the same size; same aged male Fred was smaller; two-month-older female Pooh was smaller; female Eno, who was four months younger than Alice, Ozzie, and Fred, was about Fred's size; and so on. With respect to earlier nutritional independence of one sex, there were no obvious qualitative differences, but the results of S. Altmann's nutritional study of baboon infants (in preparation) may clarify this issue. I shall turn now to a consideration of the factors that enable an infant to develop nutritional independence.
Maternal Time Budgets and Infant Maturation
We have seen that rainfall and group movements affect individuals' time budgets. How does the presence of an infant create additional demands that may restrict a mother's behavioral options and perhaps precipitate the higher mortality found at this period? In this section I shall consider the effects on a mother's time budget of just one variable, but an important one the energetic needs of her infant at various ages. The energetic requirements of pregnancy in humans and other mammals are well documented. With the birth of her infant a mother begins to provide nutrition through milk production, an energetically less efficient system than the plagcental one. As the infant grows, its nutritional requirements increase as energy is required to build new tissue, maintain existing tissue, and support higher activity levels as the infant begins to play, explore, and provide its own transportation. In the discussion of seasonal effects on time budgets I assumed that the differences between seasons in the percentage of time spent feeding result not from weight gain or loss or from changes in energetic requirements but rather from differences in feeding efficiency due to changes in food availability. In the present discussion I shall assume that within a season differences in feeding time result from differences in energetic demands. Although differences in feeding efficiency probably exist as a result of differences in dominance rank, it seems reasonable to assume that these are slight compared with differences caused by pregnancy and lactation.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 19
According to the model, at parturition a female would have to spend another 7 percent of her time (over her own 43 percent maintenance level) feeding just to provide energy for her newborn infant. A female would have to increase her feeding time from the 43 percent of her time spent feeding to maintain only her own body weight to 58-66 percent during month nine of infant life, if she were providing all of her infant's energetic needs and were doing so only through lactation (these percentage incremental values are consistent with those found for humans). She would still need to spend approximately 23 percent of her time walking for a total of 81 to 89 percent of her day occupied just by feeding and walking. However, at these levels she would no longer be able to rest for 17 percent of the day, even if she could totally eliminate time spent socializing. The result is that her overall activity level would in fact be greater, creating even greater energetic requirements for her own maintenance, and she would therefore need to spend even more time foraging.
The conclusion we must reach is that even with fairly conservative estimates of energetic demands, a mother could not provide all caloric requirements for herself and her infant beyond six to eight months of infant age and probably could do so up to that age only with difficulty and major restructuring of other aspects of her life. If the model were modified to incorporate the refinements I have indicated, would the apparent required feeding time be reduce. Most assumptions were conservative and refinements would lead to more intense time budget constraints, especially after the infant's first few months. Perhaps, then, mothers and infants have had to accommodate to these constraints as a reality. Perhaps mothers cannot always maintain their own weight and infants must provide some of their nutrition. If so, infants' maturation and learning and other factors that facilitate the transition to nutritional independence have probably been under considerable selective pressure .If we examine the observed values obtained during this study, it is clear that a female's time budget is affected by the fact that she has a dependent infant and by the age of that infant, but that the effect of infant age is less than that which was predicted above. Mothers are probably not maintaining their body weight. It is known that women who lactate successfully and whose caloric intake during lactation is 23 percent over their normal intake do not maintain their body weight, whereas at 32 percent, women maintain steady body weight. If such weight losses occur in baboons, in sufficient nutrition and weight loss may be a major source of maternal susceptibility to death and may place severe limits on the length and intensity of the lactational period unless females are able to store an appreciable supply of excess fat during pregnancy. Because reduced maternal health would directly affect the chances of survival of the mother's current infant, there will be immediate and evolutionary pressure for factors that enable infants to provide some of their own energetic requirements.
The time course of energetic demands seems to be as follows. During the first two months of infant life the additional amount of food that a mother requires to support a dependent infant over the amount required during pregnancy is primarily due to the lower efficiency of lactation but is also due to the energetic requirements of retrieving and attending to the infant when it is out of contact and to a small amount of infant growth. In the next three months the infant's continued growth and increased activity level place considerable strain on the mother's ability to maintain her own weight because the infant cannot contribute appreciably to its own nutrition at this stage; but also I suspect that mothers lose weight during this period. If milk supplies are reduced owing to maternal nutritional strain and weight loss, infants may have additional "motivation" to eat the many plant foods that they explore.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 20
By the time their infants are five or six months old, mothers are feeding all the time they can (60 percent) without sacrificing a considerable portion of social time and/or rest time. If they sacrifice rest time or time spent being groomed in favor of other activities, they will need to feed even more because any other activity would require more energy than these do. Thus it appears that mothers may have reached a maximum of feeding time by the time their infants are five to six months old, perhaps even with weight loss. It is surely necessary for their infants to provide considerably for some of their own nutritional needs. The infants can more readily do this at five or six 11 months of age if the right foods are available.
Weaning Foods
Recently, anthropologists have emphasized the importance of availability of so-called weaning foods as a major determinant of age of weaning, and consequently of the amount of lactational energetic demand on the mother and of the length of postpartum amenorrhea and interbirth intervals. What is a weaning food? For a human infant, it is a food that is both easily eaten (one that is soft and smooth, like porridge) and readily digested. For a baboon, whose mother neither collects nor prepares its food, the ease of obtaining the food must also rank among the major criteria of a weaning food.
A detailed analysis of feeding behavior and nutrition of weanling baboons in Amboseli is under way, as is a related study of vervet monkeys (Cercopithecus aethiops) in the same habitat. However, the extremes of food accessibilities are quite obvious even to the casual observer. Flowers, particularly those of umbrella trees (Acacia tortilis) and fever trees (A. xanthophloca), are abundant seasonally in Amboseli and easy to pluck. Any infant who can negotiate within the trees (i.e., any infant older than a few months) can readily eat large quantities of these sweet tidbits. In at least some years, umbrella trees flower more heavily than do fever trees. Their branches are more horizontal and have rougher bark, and therefore can be negotiated more readily by infants and at a younger age than can fever trees. Many mature blossoms come loose and fall to the ground as the baboons feed. Thus even Pedro, who could not climb at all and had no fine motor coordination in his hands, was able to eat umbrella tree blossoms, picked from the ground, as virtually his only food other than milk. It may not be coincidental that he lived two months (late December to early March) after his impairment became severe, when we thought that he would die any day. He died at the beginning of March, soon after the period of abundant acacia blossoms was over.
4 Fruits and berries (e.g., Tribulus terrestris, Azima tetracantha, Salvadora persica) and the freshest green grass blades and leaves are , probably the next best weaning foods; however, skill is particularly required to pluck A. tetracantha berries without being stuck by the abundant sharp thorns of this shrub and small infants can only reach the lowest branches. Gum of very small, shrub like fever trees is probably also a good weaning food--it is high in carbohydrates--but to obtain gum from the larger fever trees, such as are used by the baboons as nighttime roosts, requires an ability to negotiate the long, smooth, upright trunks and branches of these trees. The youngest green acacia pods are probably fairly accessible and manageable, but the youngsters apparently need to extract the seeds and discard the pods themselves, although adults (at least the males) eat virtually the whole pod. Consumption of the acacia seeds probably is limited by the presence of toxins, to which the young animals may have a lower tolerance than do adults.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 21
At the other extreme are grass corms, one of the few foods available by the end of the dry season (October), the harvesting and consumption of which account for the largest proportion of the adult baboons' time budget except in January, February, and August (feeding ; on berries and gum occupies almost as much time in December). Even older infants cannot dig anything other than the smallest corms. This is probably the major food that infants scavenge, sorting through the discarded scraps at a place where their mother or a male associate has been digging.
Is there a season when these weaning foods are most available, and is this time fairly circumscribed? For 1974-75, some relevant data are available from Post (1978). (For 1975-76 much comparable data will be available in S. Altmann in preparation and in D. Klein in preparation, but the timing of changes in the tree phenology in particular seems to be fairly invariant from year to year despite annual variations in rainfall patterns.) During 1974-75 over 50 percent of the umbrella trees had blossoms in December and January; about 15 percent in November, February, and March; none in other months. In December, January, and February over 10 percent of the trees had heavy blossom production, more than in any other months. During no month were as many as 10 percent of the fever trees in heavy flowering, but in October and November about 20 percent of the trees had some flowers, and in December about 5 percent. Ripe Azima berries were most abundant in January, and were very accessible in February and March.
Thus available data on plant phenologies indicate that December, January, and perhaps February are the months in which an infant could most readily start taking solid foods. From observations on development of both gross motor coordination for climbing in the trees and fine coordination for manipulating foods, I would estimate that a baboon infant in Amboseli would have to be at least three to four months of age even to begin to utilize much weaning food, and perhaps another two months older before attaining sufficient competence to contribute appreciably to its own nutrition. In the previous section it was also seen that mothers probably find it increasingly difficult to provide infants of this age with all their energetic needs. Infants that are conceived in December and January are born in June and July and would be at the right age to utilize weaning foods when these are most available. Therefore, one would predict the highest survival rates for infants born in June and July. Moreover, because food is apparently also most available to adults in December and January (these being the months during which the least time was spent feeding, during both 1974-75 and 1975-76), it is also reasonable to expect highest conception rates during these months.
Thus both factors suggest selection favoring a birth peak in June and July. I examined the seasonality of conception and the differential survival of infants born at different times of the year. The suggestion that food availability affects baboon reproduction in Amboseli is substantiated by the available data. Of 54 pregnancies for which we know the month of conception, 13, or 24 percent, of the conceptions occurred in the two months December and January, a result in the predicted direction but not statistically significant. However, conception by females with a surviving semi-independent offspring would be expected to be even more subject than other females to the effects of the seasonal availability of food. Of 34 such conceptions, 11, or 32 percent, occurred during December and January, about twice the number one would expect without seasonality.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 22
What of the survival chances of infants born at different times? The infants conceived in December and January also have higher chances of live birth and first-year survival (11/14 = .79) than do infants born at other times of the year (20/36 = .56). (The sample size is slightly smaller for this comparison than for the previous one owing to incomplete survival data for recent births.) Despite the large difference in these two values, sample sizes are not adequate to reach statistical significance.
Given the apparent advantages of seasonality, one might ask why infants are born any time other than June and July, and why mothers conceive any time other than December and January. Such extreme seasonality would entail interbirth intervals of either one year or two. That baboons are capable of one-year intervals is clear from the fact that these are the norm in some zoos (e.g., Lincoln Park, in Chicago); Thelma Rowell (1966b) gave the period of postpartum amenorrhea as five to six months and a subsequent cycling time as one to three cycles in her caged colony in Kampala, Uganda, and perhaps also in the rich riverine habitat of Ishasha, Uganda. Elsewhere, however, in the widely varied habitats of Gombe, Gilgil, and Amboseli, postpartum amenorrhea is of the order of ten to twelve months if an infant survives. A subsequent cycling time of about four to five months, then, does result in almost a two-year interval.
There are two situations in which one would expect "off-season" conceptions and infant survival to be higher than usual. The first is an occasional extended rainy season or other factor that leads to an extension of the season of good weaning foods and of generally higher food availability. When this occurs, females will be more likely to conceive during these off-months and infants who were born at nonoptimal times and are semi-independent when this extended good time occurs will be more likely to survive. Occasional good years may reduce the ecological pressures for birth seasonality. The second situation occurs when a female's previous infant dies. Then the female is faced with a "decision" conceive now or wait perhaps several months or more for the best conception time. If she has no dependent infant she will probably be in better physical condition and therefore more able to conceive and to produce a healthy infant with good survival chances. Also, by doing so she will not be endangering the survival of a present young offspring because she has none. Thus the variability in advantages of conception during different seasons is not as great for these females and they are more likely to have off-season conceptions. Since high infant mortality is characteristic of most nonexpanding populations, it will often tend to reduce seasonality of births in a habitat and species for which factors favoring birth seasonality are not already very strong.
These two situations probably account for the absence in the literature of a clear demonstration of birth seasonality for any baboon population. Kummer (1968) described two birth peaks in the hamadryas baboons but also noted that birth peaks occurred at different times in harems in the same area. More recently, Abegglen (1976) found a late spring birth peak in several bands of hamadryas over a period of several years. In Amboseli, the 1963-64 data and our less complete records on other groups since 1971 all indicate a seasonality similar to that documented here for Alto's Group.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 23
Seasonal variability in rainfall and food availability is reflected in the baboons' daily time budgets, the average distance they travel each day, and in the seasonality of births and infant survival. A model of a mother's feeding time, based on a mother's providing all her infant's nutrition, resulted in a prediction of severe time budget problems for mothers of older infants even at quite slow rates of infant growth. Observed feeding time was below predicted amounts for these mothers. I suggest that the compensating factors are (1) maternal weight loss to compensate for energetic requirements that cannot be met within a tolerable time budget and (2) eventually infants' providing some of their own food.
It is important to note that no category labeled "infant care" was included in mothers' time budgets. For baboon mothers as for human mothers, most infant care is done concurrently with other activities. This is true even when the infant is not in contact; mothers are probably "tuned" to their infants, alert to possible trouble. We do not yet know how infants affect mothers' abilities to attend to other stimuli or how efficiency of other activities is affected by concurrent infant care for any primate. It seems likely that the efficiency of activities such as feeding is lower than it would otherwise be for females when they have infants and that this might result in further strain on mothers.
Reproductive success and survival of Amboseli mothers and infants were seen to be subject to considerable environmental influences. It is not unreasonable to expect that their behavior and physiology have been molded through natural selection to respond to these conditions and that they also respond facultatively (flexibly) to make the best use of alternatives in the particular situations in which they find themselves at any given time.
In the next chapters I shall consider social and other behavioral interactions, between mothers and infants and between each of them and other group members; the developmental course of these interactions; the meshing of these behaviors with the less social aspects of their world that have been considered thus far; and the extent to which they relieve or intensify pressures on mothers and infants.
Social Milieu
JUST AS THE MONTH of birth and other aspects of seasonality may result in individual differences in the ecological pressures on mothers, so may individual differences in social experiences affect the mother and her infant, both directly and indirectly through effects on the mother. What are these social experiences? How do they affect mothers' time budgets and create attentional demands, and in what other ways do they appear to improve or to reduce maternal and infant survival? An understanding of the sources of these differences is the first step toward determining potential lifetime and intergenerational continuity in such experiences. At parturition the social life of a baboon female changes dramatically. She must not only nurse, carry, and protect the neonate while still providing her own food, transport, and protection, as before; but in addition, she and her infant become a major focus of interest within the group, a common characteristic of primates.
Several field studies have resulted in qualitative or normative descriptions of the social world of savannah baboon mothers and their infants: those of DeVore (1963) and Rowell and Ransom (Ransom and Ransom 1971, Ransom and Rowell 1972), all for olive baboons, Papio anubis. In addition, some quantitative data are provided by Nash (1978) primarily for 6-to 18-month-old anubis baboons and their mothers. Cheney (1978) and Seyfarth (1976) present some relevant quantitative data from a general field study on a small group of chacma baboons, P. ursinus. Rowell et al. (1968) studied mothers and infants in a small captive colony of anubis baboons.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 24
Most general descriptions of the social relations of baboon mothers are consistent across studies. In particular, all authors report that mothers of young infants receive little overt aggression, more grooming, and more approaches than do females in other reproductive stages. This general pattern of interactions within a group has also been described from field and laboratory studies of most other primates and probably comes closer than any other to being a primate "universal." One might be tempted to assume that the social group provides support for and reduction of the stress of motherhood. However, as I discuss below, a baboon mother's social world may not be as benign as these descriptions imply, and occasional extreme violence toward primate mothers or infants provides rare but chilling exceptions in some species.
The social group has an impact in both active and passive ways. The active ways are fairly obvious: individuals groom each other, play, fight, and so on. Most of the time, however, individuals, even close neighbors, are not interacting. But the mere presence of conspecifics can have a social effect even when these animals are not engaging in a social interaction. The presence of individuals who are alert to danger or who locate possible food sources may be beneficial to nearby group members. Spatial arrangements within groups also provide the broad outlines for social opportunities that may be used or not--a large adult male, just by his presence, may discourage others from approaching a mother; the mere presence of an individual who has harmed a mother previously may disrupt her activities or result in her restricting her young infant's movements. Thus consistent spatial patterning can be a clue to rarely observed interactions. In the sections that follow, before analyzing the more obviously beneficial or harmful interactions, I shall first examine the spatial patterning, that is, neighbor relationships. Then I shall turn to approaches: the interface between the statics of spatial relationships and the dynamics of interaction. Finally, I shall examine two major forms of social interactions, apparently beneficial social grooming and stressful agonistic interactions, including spatial displacements, or supplantations.
Sleeping Grove Subgroups
Most of our observations of baboons are made during the daytime. At night the animals are high in the sleeping trees. Their location and the lack of available light make observations most difficult. Fortunately for us, it appears that little activity usually goes on at night; this is suggested by two all-night watches that we have made and by the fact that individual animals can usually be relocated in the morning in the same place that they were seen to settle into the night before. Yet spatial relationships at night may be quite important. Early-morning grooming sessions often occur in the trees between individuals who sleep near each other. Also, predation seems to occur primarily at night, a fact that perhaps renders nighttime spatial relationships particularly important; for example, smaller animals may be less subject to predation if they sleep next to an adult male. Then, too, close neighbors may provide each other with warmth during the cool, dry savannah nights. Finally, the nighttime relationships, by their static nature, may elucidate patterns that exist during the day but are overlaid with foraging and with general activity levels that produce a more fluid social world. Therefore, it seems appropriate to consider first the sleeping grove subgroupings that constitute the spatial milieu during 13 to 14 hours each day.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 25
During 1975-76, although Alto's Group used 15 to 20 groves of trees that were scattered throughout the group's home range, one was used most often, over 40 percent of the time. This "Favorite Grove," as we dubbed it, consisted of two trees with continuous crown, and a third tree that was sufficiently separate that animals moving from one subgrove to the other went down one trunk and up the other rather than attempt the long jump across the branches.
It appeared that the same individuals consistently used each of these two parts of the grove: in fact, individuals were very consistent in where and near whom they slept even within their respective trees. Although I could not keep consistent records of the latter, I did record which tree subgroup each group member was in whenever the group slept in Favorite Grove and members were still settled in their sleeping postures when I arrived in the morning. All members were so located and identified on each such sample day unless, as occasionally happened, descent occurred too rapidly or too soon after my arrival. This sampling was begun in February 1976, when I could identify with confidence all individuals in virtually any position while they were huddled high in the tree, something that I could not do in the early months. I have data for 44 days that the group used Favorite Grove, from February through July and in October. For those baboons who neither died nor migrated during this period I have data from about 35 to 40 days in which subgrouping occurred (none occurred on 2 nights).
Ad libitum observations of spatial patterning in other sleeping groves were consistent with those in Favorite Grove, especially at the level of the three or four closest animals in small subgroups. However, comparable quantitative data could not be gathered for these groves owing to a lack of clear physical division within the groves, infrequency of grove use, or poorer visibility within some of the groves.
The subgroup data yield information about the affiliative patterns within age classes and the maturational trends in relationships of mothers, infants, maternal siblings, and cohorts.
Almost all individuals were at least 90 percent consistent (no more than 4 switches out of 40 samples) in their subgrove choice, most of them making no switches at all despite births of infants and other changes in reproductive state. Of the fully adult baboons, only male High Tail with 5 switches in 13 samples taken before his death and female Plum with 12 switches in 33 were less consistent. Adults consistently in the left subgroup were males Slim and Peter and females Alto, Spot, Vee, Preg, Scar, Oval, and Judy. Consistently in the right subgroup were females Mom, Lulu, Fem, Gin, Slinky, Handle, Este, and Brush and males Even, Ben, and Max.
Of the young adult males, Russ and Stu gradually made the transition from one subgrove in February to the other by October, moving in opposite directions. Among the older juveniles and subadults, Toto made 7 switches in 37 samples. The two- to three-year-olds tended to sleep in the same subgrove as their mothers but not uniformly.
All animals less than two years old slept in the same subgrove as their mothers except Pooh, who had severe locomotor impairment. She slept with or near adult Max in the rightmost tree, which she could climb more easily than the others. Fred slept with his mother; Alice did so until Alto's death, at which time Alice began to sleep with adult male Peter or her sister Dotty (Alice was 17 months old at the time). Ozzie slept with his mother, Oval, until the latter part of Oval's next pregnancy, when she overtly rejected him; he was then about 18 to 19 months of age. After that he sometimes slept with his older sister, Fanny.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 26
Several factors probably led to the average relatedness within sleeping subgroups being higher than the relatedness in the group as a whole. As noted above, both infants and their next older siblings tended to sleep in the same subgroup as their mothers. Moreover, the adult males and females of a subgroup tended to be each other's preferred mating partners. Therefore, the fathers of a subgroup's infants were likely to be the males of that subgroup. Finally, all adult females (with the partial exception of Plum) who had been members of High Tail's Group before the merger, were in the same (right) sleeping sub group. As we shall see below, a mother's sleeping subgroup tended to include the individuals who were her daytime neighbors and interactants.
Daytime Neighbors
During the daylight hours, the baboons spend most of their time foraging on the ground, where they move more easily than in the trees, sometimes gather in clusters, and are not restricted by particular comfortable locations, as they seem to be in the trees. Infants can more readily move about and do so at a younger age. In the daytime I studied neighbor relationships in finer detail. Note that when I examine the identities of mothers' neighbors and interactants, I usually focus only on the infant's first three months of life rather than older ages. This is done for two reasons: because the mother's and infant's worlds most completely overlap during these months and because, as can be seen below, these are the months of greatest social involvement. These are also the only months during which baboon infants possess their distinctive black natal coat. Coat change begins during month three and is mostly complete by the end of month six.
Two aspects of neighbor relations may be of considerable importance to mothers and infants: the density of neighbors and the identity of these neighbors. From W. D. Hamilton's work on adaptive group geometry (1971 ) and W. C. Allee's early studies of the advantages of sociality (1931), one would predict that being surrounded by more conspecifics for more of the time would be advantageous to an individual in that it would provide additional protection against predators. This would be even more true if, as is sometimes the case in baboons, this circle includes at least one adult male: the large canine teeth of adult male baboons are formidable weapons. Alternatively, other arguments emphasize the potential disadvantages of being close to conspecifics, in particular the increased likelihood of disease transmission and of feeding competition. I shall consider the relevance of each of these suggestions in the context of the neighbor patterns found in this study.
The point-sample data allow estimation of the percentage of time that animals spend in proximity to each other. I recorded the identities of all individuals who were within 2 meters of the focal female, a distance chosen because it seemed to represent an individual's "personal space": there were few animals that an individual would tolerate that close to it for long. Neighbor data for greater distances were also gathered but will not be discussed further here.
Only rarely did baboons spend 10 percent or more of their time within 2 meters of new mother. In almost every such case the individuals near a mother were either family members, one or two adult males, or another mother with a same-aged infant. Five percent of the time spent in proximity, defined as within 2 meters, was about the lowest level that I could reasonably expect to measure when partitioning each female's point-sample data by months. For each female and each month of infant life, all individuals who spent at least 5 per cent of the time nearby were considered.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 27
The sharp increase in neighbor density during month one of infant life (i.e., between birth and one month of age) over that seen during pregnancy, some increase again in month two, and a consistent decrease after month two, dropping to late pregnancy levels by about month seven to nine. Note that the average number of neighbors per sample increases by approximately the amount that we would expect from the increase in the amount of time that there is at least one neighbor within 2 meters. That is, new mothers are "alone" about half as often as when they were pregnant; however, when there is at least one animal nearby, the number of neighbors is approximately two during pregnancy and at each infant age. We do not yet know the extent of protection against predation that is afforded by this pattern of neighbor spacing, but some degree of protection seems reasonable. This type of increase--neighbors increased time with at least one neighbor but not high densities of neighbors--probably does not result in appreciable increases in feeding competition.
The opportunities for disease transmission may well be greater than during pregnancy because they probably depend not only on the time spent with others nearby, but on the number of different animals to whom a mother and infant are exposed. During the last month of pregnancy, females averaged three or four different individuals who were their frequent neighbors. Although after parturition the set of frequent neighbors still represented a small fraction of the total group membership of about 45 animals, for all of the females the number of different neighbors was greater during the first month of infant life than during the previous month, for an average of eight different individuals who spent more than 5 percent of the time nearby. Nine individuals spent at least 5 percent of the time nearby during month two, six or seven in month three, five in month four. Thus, many more different individuals were spending at least 5 percent of the time near new mothers than had done so near pregnant females, probably increasing the likelihood of disease transmission.
Both feeding competition and likelihood of disease transmission may also depend on variability in the tendency of particular individuals to engage in harmful or helpful activities when they are nearby. Some individuals may provide more assistance or threat than others, both immediately and as potential long-term associates. Who were the individuals who were frequent neighbors?
The tendency of particular adult males to associate with particular mothers is clear. For example, both Brush and Mom with Moshi had two adult males who spent time nearby, but it was Ben and Max for Brush, Peter and Slim for Mom. After an infant was about two months of age, some associated adult males also spent time with the infant when it was separated from its. Male associates were members of the mother's sleeping subgroup and usually had previous associations with the mother, sometimes including mating when the infant was conceived. Their interactions will be examined in more detail in later sections.
Although four- to six-year-olds interacted with active, especially semi-independent, infants, they rarely spent much time near the mothers. During the year of this study, all members of this age class were males (Dogo, Nog, Toto, and Swat). Females of this age class are approaching menarche and their first year of cycling. Their interactions with mothers and infants might be quite different and are now under study in Amboseli.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 28
The two- to three-year-old juveniles varied considerably in the time they spent with new mothers, even with their own mothers when they had infants. Females Cete and Nazu were only a year and-a-half old when their mothers gave birth to Summer and Pedro; it was during those pregnancies that these young juveniles were probably weaned. They did not stay near their mothers during the first months after Summer and Pedro were born and only later interacted with their siblings. The same was true for male Ozzie and female Pooh when their mothers' next infants were born. Females Janet, Striper, and Fanny, by contrast, were two- to three-year-olds when their current siblings were born, and each of their mothers had experienced at least one other pregnancy since their births. They all showed considerable interest in their siblings, as did two-and-a-half-year-old female Dotty when her sister Spot gave birth.
With family data excluded, there are still clear overall differences among these juveniles. Janet, Nazu, Fanny, and Striper were common neighbors of mothers: each of these juveniles was included in one fourth to one-third of the data for mothers with infants in the first three months. In contrast, Dotty, Cete, and male Jake appeared in less than one-tenth of the data. Thus most juveniles who tended to be interested in infants to whom they were related were interested in most unrelated infants as well. Neither age nor dominance rank explains the differences among the juveniles. Follow-up studies of these juveniles and of the later cohorts are under way. The yearlings spent much less time near new mothers than did the two- to three-year-olds, and the time they did spend was usually spent in conjunction with their own mothers. Often a weanling would get on its mother's nipple while its mother groomed a new mother. Female Alice, offspring of the highest-ranking female, was an exception: she was highly interactive in general and spent time with new mothers, often sitting in their ventrums and pushing aside these mothers' infants. She spent much time near Judy, Vee, and her sister Spot. Male Ozzie also spent considerable time with Spot and Vee but not with his mother, Oval, when she had a young infant. Female Eno and male Fred often were near Gin. Female Pooh was virtually noninteractive and spent little or no time with any of the new mothers, including her own when Peach was born.
No general patterns emerged from an analysis of adult female neighbors of mothers except that Gin and Mom were particularly likely to be neighbors of other mothers during the first month of life of the other mothers' infants. Some pairs of females who gave birth within a week spent much time near each other (Spot and Judy, Gin and Slinky); others did not (Mom and Brush, Vee and Plum). But even for those pairs that spent time near each other, the maintenance of the spatial relationship was often quite one-sided, as is shown in a later section. Moreover, some individuals who spent time together interacted seldom, and some frequent interactants did not spend much time together.
It is also clear that mothers varied considerably in the number of frequent neighbors they had, particularly during the first two months. Gin, Vee, Spot, and Scar had the most neighbors; Brush, Handle, Plum, Judy, and Mom with Moshi (but not with Misty) the fewest. In general, the former group is composed of females who are younger, rank higher, and have female infants, but it is not clear what variables determine variability in neighbor densities for neonates and their mothers. Females whose infants were born at about the same time varied--Vee with many neighbors and Plum with few, Spot with many and Judy with few, Gin with more than Slinky, Scar with more than Preg, Mom (with Misty) with more than Brush. Looking at these matched pairs does not clarify the picture. The direction is consistent with rank for four of five of these pairs, with age for four of five, and with infant gender for three of four (Plum and Vee both had female infants). Mom would be considered old and high-ranking when she had each of her infants. She had more close neighbors with her female infant, Misty, than with her male infant, Moshi.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 29
Whatever the cause, some infants more than others grow up in frequent, close proximity to many individuals. This may speed their social maturation. It may also increase their exposure to disease and increase the amount of feeding competition that their mothers encounter, but there was no clear relationship between number of frequent neighbors and disease or mortality in this study.
In summary, mothers spent more time than others in close company with at least one other animal, and more individuals spent time near new mothers than near females without infants or mothers with older infants. Particular members of each age-sex class tended to spend time near mothers and infants. Some individuals, particularly adult males, spent considerable time with some mothers, and no time with others. Some mother-infant dyads had many individuals frequently in close proximity; others had few. Kin relatedness was not a major predictor of observed individual differences.
I have suggested some implications of the results for feeding competition, disease transmission, and predator protection. To further evaluate neighbors' potential effects, it is necessary to examine the nature of the interactions that occur among individuals. I shall turn now to some of the dynamics of relationships, first the tendency of individuals to seek each other out, that is, to approach each other, and then agonistic and grooming relationships.
Approaches
Social bonds are often measured by considering only active behaviors, such as grooming or sexual mounting for evaluating positive bonds, or aggressive encounters for evaluating repulsive relationships. Yet it seems reasonable that calm acceptance and shared proximity are also important. Such relationships, if they exist, contribute to the spatial patterns seen and described above, and provide the physical availability of animals for more active behaviors such as grooming, play, infant carrying, and perhaps even fighting; but it seems likely that they more often represent positive or helpful relationships than antagonistic ones. Therefore, I shall first consider close approaches, accompanied by no other social behavior and response, that is, those that result in the two animals just sitting or standing in proximity. Then in the next section I examine approaches that are part of a sequence of social interaction. Spatial displacements, or supplantations, where it is the place rather than the individual that is approached, are considered separately in the section on agonistic interactions.
SIMPLE, OR NONINTERACTIVE, APPROACHES
For practical reasons, approaches by one individual to another usually can only be identified, in the absence of a response by the second individual, when the approacher comes within a few meters of its object. I recorded as simple, or noninteractive, approaches all those approaches that carried the actor at least 1 meter (approximately a body length) and that brought the actor to within 2 meters of the animal being approached. These could be reliably scored without response by the object animal. As noted above, 2 meters seemed to represent an animal's personal space. Thus I wanted to determine which animals would be approached that closely with no further interaction, and which animals' approaches to such proximity would be tolerated by an animal who was approached. With which animals do such calm approaches occur? Are they more common or less after parturition?
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Baboon Mothers and Infants by Jeanne Altman excerpts part 30
For all 13 females the average rate of approaches to them was greater than the approaches by them. Each mother had a small set of individuals whom she approached, and this was to a considerable extent a subset of those from whom she tolerated approaches. In particular, most mothers had one adult male with whom they shared these approaches, for example, Mom-Moshi with Slim, and Brush with Ben.
There were usually more individuals who approached mothers than the mothers themselves approached. From the last month of pregnancy to the first month of infant life, both of these total rates (to and by the mother) rose slightly. The amount of the rise was small, but the change in direction was consistent: it increased for seven of nine females in the approaches by them and nine of nine for the approaches to them. In some sense, then, one might say that the time of birth was a time of drawing together of individuals who were relaxed with each other, and that this is one of several dynamics that caused the increase in neighbors described previously. For later months of infant life the rate of noninteractive approaches varied little from month to month, and it was inconsistent in direction of change from individual to individual and month to month. Unlike the rate of interactive approaches I shall consider next, the rate of noninteractive approaches does not reflect developmental changes in mother-infant contact or other aspects of infant maturation.
In examining differences among mothers it is necessary to consider that rates of noninteractive approaches both to and by mothers were probably underestimated, and equally so, in the first few months of this study. Thus to compare the rates for various females I did not include data before late September. Considering the remaining data, I found no pattern in the rate at which various mothers made noninteractive approaches to others. In the simple approaches made to mothers, the three highest-ranking (Vee, Spot, Mom with Moshi) had the highest rates (but Mom with Misty had the lowest). Among the juveniles only Dotty consistently made noninteractive approaches to several different mothers. The lack of a very clear-cut pattern may be due to the overall low rates of these behaviors.
Except perhaps during the first month of infant life, the rate of these calm approaches probably contributes to, but is not sufficient to account for, the appreciable increases that we found in the number of neighbors; but it is unlikely that the rate itself has an appreciable influence on mothers and infants. What is probably more important is the actor-recipient selectivity of these acts: mothers are brought closer to particular adult males and in some cases to other females with like aged infants. Those cases for which we have both likely paternity and behavioral data during the infant's first months indicate that the adult males involved are likely to be the infant's father. The adult female associates are not likely to be closely related through maternal lineages, but rather share common needs and experiences, and their infants provide each other with play mates. Actually, mothers of like-aged infants, or the infants themselves, may be more closely related than we suppose from a consideration of only maternal genealogies. Females who are closely associated tend to associate with the same one or two males. Thus their infants are likely to be paternal siblings. If, in addition, these mothers are of the same age cohort and formed their association in infancy, they, too, may be paternal siblings. At this stage of our knowledge, however, these suggestions are purely speculative.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 31
INTERACTIVE APPROACHES
A much more common class of approaches consists of approaches that were accompanied by other social behaviors or that were followed by some detectable reaction on the part of the animal being approached. These interactive approaches were followed by sequences of behavior ranging from relatively long energetic ones, such as fights or other dominance interactions, handling and pulling of associated infants, and grooming, to brief lipsmacking, body aversion, or a slight touch. Most social interactions, both beneficial and harmful ones, begin with an approach. Spatial displacements, which involve approach to another animal's location rather than to the animal itself, are considered separately in the agonistic section below.
In general, mothers initiated interactive approaches slightly more often than they did the simple, or noninteractive, approaches, and did so at a rate that increased very slightly with the birth of their infants and then changed little with the maturation of the infants. This is in striking contrast to the rate of interactive approaches directed toward mothers, which soared with the birth of their infants and then steadily decreased in the subsequent months. In their infants' first few months of life, mothers were all the recipients of interactive approaches much more often than they were the actors; later the differences were small and reversed in direction for some females and for some months. The six- to eightfold increase in approaches after the birth of an infant is much greater than would be expected from the fact that two individuals (mother plus infant) are now being approached rather than just one, as before.
As an infant matures, its mother may be approached less often because the infant spends less time in contact with her, enabling others to approach the infant at times other than those when it is with its mother. The relationship between infant contact and approach rate can be examined at the level of the immediate behavior by determining whether at any given infant age a mother receives more interactive approaches when her infant is actually in contact with her than the expected number of interactive approaches, calculated from the proportion of time the infant spends in contact that month multiplied by the total number of interactive approaches to the mother that month. Using CRESCAT to perform sequential pattern searches, I determined whether the infant was in contact or not when an interactive approach occurred by identifying each change in mother-infant spatial relationships and then labeling each occurrence of a social behavior, including interactive approaches, according to whether the mother and infant were in contact at the time the act was performed.
When their infants were in contact, mothers received more interactive approaches than expected in 59 of 72 mother-months (plus 13 ties). Results of (binomial) significance tests are a function of probability (contact time) and N (total number of approaches) for any month, which vary considerably in this case. Although the magnitude of the difference between observed and expected values was sufficient to reach the .05 level of significance for only about half (34) of these positive-deviation months (and two of those with negative deviation), the probability of 59 of 72 results being in the same direction by chance is extremely low (P <.001). Moreover, if data from all females for each month are pooled, mothers were approached significantly more often when their infants were in contact during all but months one and twelve.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 32
Interest in the mother-infant pair may also wane if older infants are less attractive to others than are younger ones. To examine this possibility I looked at the rate of approaches to a mother when her infant was in contact. For four females data were available for the last month of pregnancy as well as for all of the first six months of infant life (but not necessarily for later months). For each mother I calculated the rate at which she received interactive approaches during the last month of pregnancy, which I considered her "base rate," the rate at which she would receive such approaches if she did not have an infant. Then, if a female's "attractiveness" was independent of her status as a mother and the presence of her infant, we would expect that she would receive that same base rate of interactions while her infant was in contact (or, at most, twice the base rate if one considers that two individuals are being approached) and that this rate would be independent of the age of her infant. After the base rate for each mother was subtracted from her rate for each month thereafter. They demonstrate that a female received approximately six times as many interactive approaches after parturition when her infant was in contact, and that the rate of approaches was independent of infant age until the infant was four or five months old. However, the rate dropped dramatically in the sixth month. It is interesting to note that at about month six infants have lost most of their distinctive black natal coat. The present results are consistent with the suggestion that infants who have lost their neonatal coat may be less attractive to other group members. However, the decrease in approaches to mothers of older infants may be produced by other factors. For example, infants may remain attractive but others may selectively approach them when the infants are not in contact with their mothers. I shall also examine the contingencies between infant contact and maternal activities at certain infant ages, and consider the relationship of those contingencies to alternative explanations for the remaining age effects found in approach rates.
Who were the individuals who approached and interacted with mothers, or whom the mothers themselves approached? For interactions with adult females the pattern was somewhat different from that for interactions with males. A few females seemed to exhibit interest in many infants, others in none. This individual variability is similar to that reported by Breuggeman (1973) for the Cayo Santiago rhesus colony. Thus Spot, Mom, Gin, Slinky, and Oval each appeared as frequent interactants for one-fourth to one-third of the new mothers, themselves of course excluded. At the other extreme, Plum, Scar, Brush, and Lulu never did. The "high-interest" females could also be identified by the amount of time they spent as neighbors of other mothers. None of the high-interest females was closely related to mothers through maternal lineages, and there is no basis for suspecting that through paternal lineages they are more closely related to mothers than are the low-interest females.
New mothers themselves rarely approached others during the early weeks of infant life. One exception was the approach made by mothers to other females with very young infants, an event that was more likely to occur if these other females were lower in rank than the mother.
For each mother there were only one or two males that approached and interacted; often the same males who spent more time near her and with whom she shared noninteractive approaches.
Usually I could predict the existence and identity of such affiliations with males before the birth of an infant, commonly from its mother's mating, grooming, and neighbor associations. An affiliated male was likely to be the infant's father, that is, the only male, or one of only two or three males, that copulated with the mother during the days that she conceived the infant
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Baboon Mothers and Infants by Jeanne Altman excerpts part 33
The older juveniles and subadults, all male, rarely approached the mothers. Among the younger juveniles there was considerable variability. The two highest-ranking juveniles were the ones who most frequently approached and interacted with mothers. However, there was no consistent relationship between juvenile dominance rank and tendency to approach. High-ranking Striper, who appeared as a frequent neighbor of new mothers, was also the juvenile who frequently approached and interacted with mothers, including her own. High-ranking Dotty appeared more frequently as an interactant than she did as a, neighbor. Low-ranking Janet and Fanny were intermediate as interactants, and the others rarely approached and interacted.
Also striking is the variability among mother-infant dyads in the number of these frequent interactants. Vee had many (13) such interactants, but most of them were juveniles and older infants. For all the other mothers the variability in the number of interactants was primarily due to the number of adult female interactants, with Gin, Slinky, and Judy all having 6 or more and Spot, Mom, and Vee having fewer than 2 adult females who frequently approached and interacted with them. The highest-ranking mothers had the fewest such interactants: there was a slight tendency for the number of interactants to be inversely correlated with a mother's rank even though there was no tendency for the total rate of interactive approaches to mothers to be rank related. Neither variable was obviously related to infant gender or maternal age. Whatever the origin of the differences, some infants had early exposure to interactions with many group members, especially adult females; others had few. Moreover, the infants differed in the particular individuals to which they had the most early exposure, a selectivity that is not easily related to kinship, as discussed previously. The nature of the social interactions in which mothers and infants are involved may provide important clues to differences in early experience. I shall now examine several major types of social interactions, some apparently beneficial, others stressful, in more detail. These interactions sometimes followed immediately upon the approaches I have just discussed. At other times they occurred between two animals that already were in proximity to each other.
Grooming
Assumed to be beneficial, grooming is the most obvious and time-consuming form of primate social behavior. The amount of time that grooming was done to or received from others for the females in this study. The values were obtained by using CRESCAT to locate grooming bouts in the files and then having the computer subtract the time that grooming ended from the time that it started, cumulate these times for each dyad during a month, and divide the cumulated time by the appropriate total observation time for the month. Females spent little time either being groomed (1 to 2 minutes per 100) or grooming (3 or 4 minutes per 100) during the last month of pregnancy and were more often actors than recipients of grooming at that time (six of the females groomed more than they were groomed, three tied, and one, Plum, received more grooming than she performed). During the first month of infant life the average rate of grooming by mothers decreased to between 1 and 2 minutes per 100 and the amount of grooming time that mothers received increased greatly (to 7 minutes per 100), further increasing to a peak of over 8 minutes per 100 during the second 2 month of infant life. The amount of grooming mothers received dropped greatly during the third month, to 3 to 5 minutes per 100, and decreased very slightly each month thereafter to levels of about 2 to 3 minutes per 100 (sample sizes became small and variability great). After their infants were two months old mothers tended to spend about 2 minutes per 100 grooming others.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 34
All mothers except Spot groomed their own infants more than they groomed any other individual, and all were groomed by others more than they groomed others. As new mothers, only Brush and Handle did much grooming of other adult females. For other new mothers almost all their grooming was directed to their own neonates, their juvenile offspring, and one or two associated males. These males and juvenile offspring also groomed the mothers, as did a few other associates. The identity of these other groomers differed among mothers with the striking exceptions of adult female Slinky and juvenile female Janet, who did appreciable grooming of most of the new mothers. For Janet, this included providing most of Oval's grooming, which Oval reciprocated. Strangely, Oval's own juvenile daughter, Fanny, neither groomed nor was groomed much by her mother when Oreo was young, although they were frequent grooming partners at other times.
There were no consistent relationships between a new mother's dominance rank and the amount of grooming done by or to her. This contrasts with the finding of Seyfarth (1977) for a group that included eight adult female chacma baboons. The other baboon studies have not mentioned a correlation between maternal rank and the amount of grooming received. Except for that of Saayman (1971), they do report more grooming of females with young infants than of females in most other reproductive stages.
Grooming is usually assumed to be beneficial for removal of ectoparasite and for social bonding itself of unknown biological cost or others. benefit. Although grooming is quite time-consuming, it is done primarily to, not by, mothers and, moreover, is initiated by others: that is, mothers rarely approach and solicit (i.e., present for) grooming. Groomees are in general quite relaxed, although mothers are somewhat less relaxed than other groomees because those who groom mothers often stop grooming and reach for the infant, at which point a mother responds immediately by clutching her infant. We can only say at this point that grooming, unlike some other interactions, probably is not harmful and may be beneficial. Although it is time consuming, and from that standpoint would seem to stress mothers' time budgets additionally, time spent being groomed may obviate the need for some of the usual rest time.
Dominance Relationships and Agonistic Interactions
For almost every pair of females one or both members of the pair experienced at least two different reproductive conditions during the year. However, no pair of individuals reversed the direction of their dominance-subordinance relationship as a result of birth or maturation of their infants, that is, dominance relationships were independent of reproductive states. For possible changes in agonistic relationships due to the birth of an infant we must look beyond the simple "winner" or "loser" outcome of agonistic encounters.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 35
SPATIAL DISPLACEMENTS
Sometimes one individual moves closer to another individual (or' would have done so if the second animal did not leave), but it is the space or location of the second animal that is being approached, rather than the animal occupying that space. These spatial displacements, or supplantations, are usually distinguishable from what I have called approaches by the direction of gaze of the approaching animal and the endpoint of the approach: spatial displacements do not involve "tracking" or following the recipient. The animal whose site is approached responds with mild submissive gestures, for instance, a slight cower or several rapid glances, suddenly discontinues its current activity, and moves away. These spatial displacements were not considered approaches to an animal in the previous discussions. Supplantations were commonly but not necessarily followed by the actor's assuming the same activity (feeding, grooming) that had occupied the animal who was displaced and doing so at the same spot. Actors thereby take over resources such as partially dug grass corms. "Approach-avoidance" interactions are frequently discussed in the primate literature, as are spatial displacements, but definitions are rarely provided and there has not been a tradition of consistent terminology. l have used a somewhat more restrictive definition of spatial displacements than Post did; he would include as displacements some of the approach-avoidance interactions that I discussed previously in the sections on approaches. Rowell (1966a,1969) and Seyfarth (1976) made a different pooling of categories, combining and considering as "friendly" approaches all approaches to the place or to the individual that did not include any threats by the approacher.
Spatial displacements are a common type of agonistic encounter. Averaged over all females, the rate of spatial displacements both to and by mothers was slightly lower during the first month of infant life than during the last month of pregnancy, but otherwise the average remained at fairly constant levels independent of the existence or age of a female's infant. Six of nine females were displaced at lower rates during their infants' first month of life than during the last month of pregnancy. The three whose rates increased were three of the four females whose infants were born during the latter part of the dry season. Post (1978) found that displacement rates in general are higher during that season. Unlike berries, flowers, and most other baboon foods, the main dry-season food, grass corms, requires considerable time and energy to process, because the corms are dug from the ground. Thus one animal can benefit appreciably by supplanting another from a corm that the second has been digging. From month one to two of infant life the rate of spatial displacement increased for six of nine females, as it did for six of nine between months two and three of infant life. By month three, five of eight females were supplanted at higher rates than they had been during the last month of pregnancy. Thus infant birth may bring with it slight immunity against spatial displacements, but this immunity has disappeared by the infant's third month.
Again, Brush and Mom provide an interesting contrast. During the month before Moshi was born, high-ranking Mom's rate of spatial displacements was approximately the same as Brush's was during her last month of pregnancy. However, Brush was displaced at a slightly higher rate during her infant's first month of life, whereas Mom was displaced appreciably less after Moshi was born. Even if we take into account the fact that no observations are available for Moshi's third month, Brush was displaced about twice as often as was Mom during the first few postpartum months.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 36
During the last month of pregnancy high-ranking females as a group were supplanted at much lower rates than were the low-ranking ones, but there was no consistent tendency for rank order and rate of spatial displacements to be related. The decrease in rate of displacements during the next month resulted in less difference in rate of displacement between high- and low-ranking females during their infants' first month than there had been before their infants' birth. . This perhaps accounts for DeVore's conclusion that new mothers gained in dominance rank. Although DeVore's matrix of dominance interactions did not suggest a rank change (change in outcome of agonistic interactions) as a function of reproductive state, DeVore indicated that rank changed when a female's infant was born. Perhaps this interpretation was the result of his perceiving reduced rates of spatial displacements at this time. However, as noted above, in the present study even this possible immunity was brief and had entirely disappeared two months later. Also, in the present study low-ranking mothers were displaced more often than high-ranking ones. That is, there was a general trend for maternal rank to be inversely related to displacement rate pooled over the first three months of infant life, the time when overall interaction rates were by far the highest. Somewhat more clear-cut are the data for the five females for whom several sample days are available for each of these three months. Using y, a measure of association indicating how much more probable it is that a pair of individuals will be ranked in the same, rather than the opposite, order on two variables (in this case dominance rank and rate of being supplanted), I obtained values of .40 for all females but .80 for just those five females. That is, pairs of mothers were much more likely than not to have the same relative dominance rank and rank order on 18 rates of supplantation.
OVERT AGGRESSION
It was extremely rare (less than once per 100 minutes) for mothers either to display or to receive acts of aggression or overt threat. Of the 192 cells (mothers x months observed x--actor or receiver), in only 1 was the rate over two acts per 100 minutes; in only 1 was it over four. In general, these females were more likely to be the recipients of aggression than the performers of aggressive acts. This was true for the last month of pregnancy and throughout the first year of infant life, with no consistent variability as a function of infant age. There were no consistent patterns of rates of aggression with respect to dominance rank. The absence of detectable developmental and rank differences is probably related to the general rarity of the behaviors.
DISTRESS
If females receive a considerable amount of grooming and little overt aggression when they have young infants, is it reasonable to view their increased social involvement as basically calm or "positive"? Perhaps not, as suggested by the reports of mothers frequently avoiding "friendly" approaches. I analyzed the data on behaviors of "distress" or "fear" during social interactions. These social interactions do not include, but rather are in addition to the spatial displacements that have already been discussed. I counted one occurrence of a distress response whenever a female gave any one or more of the submissive behaviors, regardless of the intensity or duration of such behavioral acts; that is, if a mother responded to a partner with several submissive behaviors such as a grimace and a tail-up, I counted all the behaviors as one distress response unless an act by the partner intervened. Thus, for the time being, I have ignored intensity of distress.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 37
During the first month of infant life, mothers averaged almost 11 distress responses per 100 minutes; individual response rates were strongly related to maternal rank, with low-ranking mothers having higher rates of distress responses than did higher-ranking ones. The same result, indicating the importance of dominance rank, is seen in the pooled data for months one through three, y = .80; that is, the birth of an infant greatly exaggerated the effects of dominance rank and dominance affected the amount of distress experienced by new mothers and, in all likelihood, by their infants.
The rate of maternal expressions of distress decreased with in creasing infant age until by month eight levels were near those of late pregnancy. An alternative way of viewing the effect of the infant on stressful social interactions for its mother is to plot these interactions for each mother for each month against the percentage of time the mother's infant was in contact that month, as was done for interactive approaches, again producing a striking correlation but with much more scatter because dominance rank did not affect the rate of approaches as it did the rate of distress responses.
The fact that the rate of distress responses by mothers drops off at 1; about the same rate as does mother-infant contact suggests that mothers may be subjected to stressful situations just when their infants are in contact, or perhaps even that they could avoid stressful situations by not being in contact with their infants. To investigate this possibility I analyzed the rate of distress responses when the mother's infant was in contact, as I had done for the interactive approaches to mothers. If infant presence is the relevant situational variable, one would predict that mothers would express distress disproportionately when their infants were in contact. Moreover, if the infant's immediate presence alone was what resulted in the mother being in stressful situations, regardless of infant age, the rate of distress responses when the infant was in contact would be expected to remain constant as the infant matured. Alternatively, if an older infant is less attractive to others and/or if a mother is less sensitive or fearful when her infant is older, then the overall decline in distress would also be present when mother and infant were in contact.
I first analyzed the rate of distress responses, contingent on the infant being in contact at the time of the interaction, using conditional pattern searches in CRESCAT, as described previously. Mothers did perform behaviors of distress disproportionately when their infants were in contact, particularly when their infants were less than six months old. The data partitioned by individual mothers and by months rarely reached statistical significance, but the direction of the results is quite consistent; in 60 of 83 mother-months (plus 4 ties), infant contact was linked to maternal distress (P <.05, binomial test). Pooling the data for all females for each month results in significant values for months one through six and months eight and nine.
As I had done with interactive approaches, I then examined the rate of distress responses when a mother's infant was in contact for those mothers for whom data were available for the last month of pregnancy and also the first six months of infant life. These females averaged approximately four distress responses per 100 minutes during the last month of pregnancy. When their infants were in contact the following month, there was an average increase of more than nine such responses per 100 minutes. As expected from the several previous findings, the rate of maternal distress when the infant was in contact decreased only slightly with increased infant age for the first few months of infant life. That is, mothers are, in general, put in stressful situations because they are with their infants, and this decreases relatively little during their infants' first few months of life.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 38
Although the absolute rate of distress responses during infant contact changed little during the first six months, the percent decline was more than for interactive approaches to the mother and, with the exception of an increase in the fourth month of infant life, there was a steady decrease in the rate for each of the first six months. This slow steady decrease continued for those two mothers for whom I also had continuous data for the next several months (months six through nine). The gradual reduction in distress responses during contact that occurs may be due either to the mother's decreased "feeling" of vulnerability when the infant is older or to reduced attractiveness of the infant, which reduces other animals' attempts to interact. However, the rate of distress responses when the infant was in contact decreased with infant age slightly earlier and more steadily than did the rate of interactive approaches to the mother (a rate that variably increased or decreased very slightly until month five) and showed no sharp change at any infant age; this suggests that the mother's feeling of vulnerability or evaluation of the potential danger in the interactions may decrease earlier and respond to gradual infant maturation, as contrasted with the behavior of others, which changed dramatically after the major loss of the natal coat.
Mothers directed the distress responses and acts of embracing or restraining of their infants disproportionately more toward other adult females than toward individuals of other age-sex classes, and especially toward those females who were both higher in rank than the mother and among the females particularly "interested" in infants, especially Spot, Mom, Gin, and Slink. Thus only male Slim and, to a lesser extent, juvenile male Nog caused appreciable distress to Spot, and only Spot and to a lesser extent Slim and male Peter did so to Mom. In contrast, low-ranking Brush often was distressed by male Red and by females Spot, Vee, and Slinky; she was somewhat less distressed by Fem and Gin.
Infant Handling and Pulling
What are some of the distress-provoking situations to which mothers are exposed? Recall that in the analysis of aggressive behavior, approach was not considered a threat, nor have previous researchers considered it such. Nor has handling or pulling of an infant appeared in catalogues of threat or aggression, and thus I did not include them as such in the analysis of aggression above. Yet a mere approach often elicits submissive (including avoidance) behaviors, as several authors have noted (e.g., DeVore 1963, Rowell et al. 1968, Seyfarth 1976). In addition, other animals often handle an infant, sometimes even pulling it away from its mother with sufficient force to produce counter-pulling by the mother, screeching by the infant, and other signs of distress in both. Such pulling and handling occurred often during the infant's first month, was in addition to mere touching or muzzling, and was primarily done by other adult females to infants of lower-ranking females. In this study it seemed clear that mothers "perceived" the mere approach or presence of certain individuals, and certainly handling and pulling of the infant, as a threat or a source of distress. In the extreme cases, snatching resulted in infants being kidnaped. Usually this kidnaping was brief, but in one instance the kidnapper, Gin, kept the infant overnight. When female Handle finally was able to regain possession of her two-day-old son about 15 hours after kidnaping, Hans was weak and appeared dehydrated. Hrdy (1976) reviews instances of kidnaping in several species.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 39
Repeated handling and pulling of infants of low-ranking mothers seemed to have at least two consequences. It was probably a major factor affecting maternal. In addition, some mothers, when they were repeatedly approached and their infants repeatedly yanked, would approach and sit next to a particular adult male associate. Thus, as suggested in previous sections, relationships with adult males provided a major source of variability in infant experience. These relationships will be discussed below.
Associated Adult Males
Adult males, like females, showed varied degrees of interest in infants. However, for males more than for females, interest depended strongly on the identity of the mother. Similarly, each mother's tolerance of a particular male's approaches depended on the identity of the male. That is, there were specific adult male-mother associations, and these seemed to reflect specific relationships between male and female that were established before the infant's birth. In most cases, at least two males attempted to associate with the mother and infant soon after parturition. The higher-ranking male sometimes exhibited herding behavior, followed the mother, threatened other males away, groomed the mother, and, in fact, exhibited all elements of a sexual consortship except for actual mounting. This was most striking in Slim's pursuit of Mom when Moshi was born. He frequently herded Mom from Peter and occasionally even fought with Peter in these situations. Likewise, Ben prevented Max from remaining close to Brush when Bristle was born. Wounded and dropping in rank, Ben was unable to do the same after Sesame's birth to Slinky in February. As with sexual consortships, the role of the female in these interactions was not that of a passive observer. A mother did not follow a lower-ranking male while being herded by a higher-ranking one, nor did she ever threaten males who followed her; but she selectively avoided certain males when they approached, reciprocated the following or not, selectively followed and groomed certain males, and so on.
The result was that some relationships between mothers and adult males were more reciprocal and enduring than were others. The associated male, either through overt behavior or through his mere presence, often provided a buffer between the mother-infant dyad and other group members. When the male was near the mother-infant pair, others approached more hesitantly; they gave repeated "anxious" glances toward the adult male, and veered in their approach to the mother so that they approached her on the side opposite the male. They sat farther from the pair than usual, watching, and then suddenly trotted toward the mother and infant as soon as the male moved away. The male some times overtly threatened those who approached the pair, particularly if the interloper pulled at the infant. One result of such male influence was that the rate of infant pulling during an infant's first month was usually lower in the presence of the associated male.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 40
The presence of Slim did not usually reduce the rate of infant pulling. This was because he was the major infant puller for those females with whom he associated. An additional effect of male presence was that in the second month of life, when most infants spent moderate amounts of time away from but still near their mothers, infants were out of contact more when the male was within 2 meters than they were otherwise.
Adult male-infant relationships persisted beyond the neonatal period. Older infants rested against their male associates and obtained rides from them, ran to them in times of distress, and took greater liberties with these males than with others by feeding near them or by getting meat scraps from them. Elsewhere I have referred to the male who had an enduring and beneficial relationship with an infant as its "godfather". Particularly striking godfather relationships were those of Pooh with Chip and then later with Max, Fred with Max, Alice with Peter, Bristle with Ben, Hans with Even and Grendel with High Tail.
There is another side to the male-infant relationships: the males sometimes derived apparent benefits from their relationships with infants. In several species males under attack sometimes take infants to their ventrum, an act thought to serve as an "agonistic buffer", that is, to mitigate attack. However, in this study, only if the male and the infant had an existing positive relationship did the infant remain riding and possibly serve this function when the male was under attack. Otherwise the infant usually refused to cling. Then it either dropped off or the male embraced it continuously, running three-legged. The male's locomotion was thus hindered and the infant usually screamed in distress, bringing other baboons to its aid. Thus, in summary, the same male was likely to be an infant's possible father, its protector, and its exploiter.
Summary and Discussion of Social Relationships
ADULT MALES
The males that were associated with mothers usually were fully mature rather than subadult or newly adult males, and the associated males tended to come from the upper half of the dominance hierarchy. Top-ranking Slim and mid-ranking Peter were the males who were associated with the most mother-infant pairs.
These patterns of male associations can be compared in more detail with the sleeping grove subgroupings. Mothers were associated with the same males that were in their sleeping subgroup; more specifically, they were usually associated with the adult males who were most consistently in their subgroup. Only for Plum and Mom was sleeping grove member ship contrary to the data on mating associations and to ad libitum data on their associations before the birth of their infants. Their male and their female associations after the birth of their infants were consistent with their previous associations, but not with their general sleeping subgroup membership.
DeVore (1963) reported that all mothers associated with a group of dominant males who prevented other animals from access to young infants. Ransom and Ransom (1971), and Seyfarth (1978), in contrast, described a variety of individual pairwise relationships, a description more consistent with the data from this study.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 41
ADULT FEMALES
Can the associations of adult females with mothers be depicted in the same way as the male-mother associations? These female associations do not emerge with the same clarity as do the male associations. First, more females than males are associated with each mother, as would be expected from the availability of 50 percent more adult females than males (this takes into account deaths, maturations, and migrations). Second, these female associations appear to be more diffuse or of lower intensity than the male associations, most receiving total scores of 1 rather than the 2 to 5 of male associations. Omitting interactive approaches and omitting relationships with a score of only 1 made little difference in relationships with males. Doing so with the relationships between mothers and other females eliminates most of the relationships. Lower-ranking females who sought out a mother-infant pair sat near the mothers and appeared in the data as their neighbors, but did not often interact with them. In contrast, approaches by higher-ranking females were usually followed by interactions, both because the high-ranking females were more likely than were low-ranking ones to initiate interaction and because mothers usually reacted to these higher-ranking females; one of the mothers' primary reactions was avoidance. Thus there was some tendency for higher-ranking females, especially infant-pulling ones, to be frequent interactants, but not to spend much time as neighbors, and the converse tended to be true for lower-ranking females.
So the difference in dominance rank between a mother and an other adult female and the presence of other higher-ranking female and male associates, as well as a female's tendency to pull infants and whether that other female herself has a young infant, all affect the degree of association between the two, modifying their tendency to associate at other times and producing a range of associations more complex than the associations with the adult males. Yet, mothers' female associates, like their male associates, came disproportionately from their sleeping subgroups. This is especially true if we consider all mothers other than Plum and Mom. For all the other mothers there was a total of 25 close relationships with other females (by the criteria used for the male relationships), 17 of which were with females in the same subgroup. Three of the remaining 8 associations that crossed subgroup lines were between females with like-aged infants. In contrast, 8 out of 9 of Mom's and Plum's associations as new mothers or with new mothers were with females of the other sleeping subgroup. However, even this inconsistency is reduced when we consider that during October 1976, Peach's first month of life, Plum was always in the "right" subgroup in the sleeping trees, the same subgroup that contained the adults with whom she was associated both during that month and before then. Thus only Mom consistently slept with one subgroup but interacted with the males and females of the other.
We have seen that those individuals who primarily compose a new infant's world are a subset of those available in the group, a subset that is different for different infants, with some consistency over various activities and with some continuity with the mother's relationships before the infant's birth. Thus although parturition is an event that brings dramatic change in a mother's social life, certain relationships are maintained or intensified at that time.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 42
Summary and Discussion of Specific Forms of Interaction
The picture of the social milieu that emerges is a complex one. Some behaviors and relationships were essentially independent of the existence or age of a female's infant: the rate of overt aggression and of noninteractive approaches and the direction of dominance-subordinance relationships were all of this sort. However, several other dramatic and time-consuming behaviors became much more common soon after parturition and remained infant dependent: behaviors of fear or distress, receipt of interactive approaches, and receipt of grooming. These increased dramatically during month one of infant life and peaked during either month one or month two. The increase in these was due to other animals, not to the new mothers, who themselves maintained or decreased levels of initiation of behavior. More specifically, interactive approaches to mothers and maternal distress occurred primarily at times when their infants were in contact. Attractiveness of infants remained high until the infants were about five months of age; but apparently the reduction in mothers' feelings of vulnerability or stress was more uniformly gradual over their infants' first six months of life.
Although all females experienced these social changes after the birth of their infants, there was considerable individual variability in these experiences, either in their magnitude or in the identities of those who composed these mothers' social world. Some mother-infant pairs had many neighbors and others had relatively few. Some were approached and interacted with almost 30 times per 100 minutes, others about half as often. None of the major demographic or sociological variables--maternal rank, infant gender, or maternal age--proved to be clearly related to these differences, although there was a slight tendency for higher-ranking mothers as compared with lower ranking ones to receive noninteractive approaches from more other females and interactive ones from fewer other females. This was probably due to the hesitancy of others to interact with the high-ranking mothers and the lower level of stress or fear that approaches produced in these mothers. Seyfarth (1976) and Cheney (1978) found some tendency for higher-ranking females to be approached and groomed more than were low-ranking ones, a result that is contrary to that of the present study. I suspect that individual behavioral tendencies, that is, personality differences, sometimes override or interact with demographic or sociological effects. Unlike some high-ranking females such as Alto, the two highest-ranking new mothers in my study, Spot and Mom, were particularly aggressive individuals. Perhaps if they had not been aggressive, Spot and Mom would have been approached or groomed more.
Cheney (1978) further suggested that juvenile females seeking interaction with new mothers do so primarily with their own mothers and with female nonrelatives that are higher-ranking rather than with nonrelatives that are low-ranking. At least the latter situation does not seem to have been the case in the present study. It was not tested with Cheney and Seyfarth's own data, and additional information would be required in order to calculate the appropriate expected values based on availability in the group of higher and lower-ranking adult females with infants, in relation to each juvenile. The problem in making the test for that study is further complicated by the absence of genealogical data beyond Cheney and Seyfarth's 18-month study and therefore by their assumption of kinship relations based on behavioral association. At present the hypothesis cannot be disentangled from other interpretations.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 43
Finally, there were two major variables in my study that were clearly correlated with maternal dominance rank, the rate of expression of fear or distress and the rate of being supplanted. By maintaining a close relationship with an adult male, low-ranking mothers can partially avoid or buffer these interactions, but they remain potent ones for mothers and their infants. For the most part, maternity exaggerated the effects of dominance rank. Occurrences of these stressful interactions may be the major factor determining the establishment of offspring dominance rank according to maternal dominance rank that has been reported for macaques and that we have observed in this group. Long before opportunities arise for maternal support of infants observed in the infants own agonistic encounters, infants have had ample opportunity to learn their mothers' rank and the identity of those individuals to whom their mothers respond with fear. The infants soon seem to respond selectively to other individuals. Perhaps these early interactions account for Norikoshi's (1974) observation that by three months of age the behavior patterns of play used by each member of a pair of Japanese macaque infants reflected their future dominance relationship. Over the past seven years, in the two instances of mothers who dropped in rank (Oval, Judy) and the two in which mothers (Alto, Fluff) died when offspring were about one-and-a-half years old, the offspring assumed the ranks their mothers occupied during the period of the offspring's infancy.
In this context recall that spatial displacements, as well as most interactive approaches by other adult females, were made by individuals that were higher in rank than the mother. In addition, it was also primarily in response to higher-ranking females that mothers embraced or restrained their infants or directed their distress responses. That is, there were few instances of undecided or reversed dominance interactions, and even nonagonistic interactive approaches followed the same partner asymmetry as did agonistic interactions. If these interactions are important in the early establishment of dominance relationships, I would predict that those higher-ranking females who were most interactive with a mother and infant would be the first to establish dominance relationships with that infant. For female infants this would mean early establishment of permanent dominance relationships; for male infants it would mean that those females would remain dominant to the infants longest--all juvenile males eventually become dominant to all adult females. The order and dynamics of establishment of dominance relationships in this group are currently under study.
The increased and time-consuming social life that is imposed on mothers perhaps provides some predator protection, but it also entails increased exposure to disease and perhaps to feeding competition. Not only does it result in mothers' receiving much more grooming and thereby presumably greater rates of ectoparasite removal, but because they do less grooming of others than previously, the amount of grooming received for the amount of grooming provided is even greater than would appear at first.
However, even nonharmful interactions take time and even small amounts of time add up to a considerable total. Just 25 sequences per 100 minutes, the rate of approaches that were followed by interaction, add up to 150 sequences per ten-hour day. At even ten seconds each, these sequences alone add about 4 percent of imposed socializing time to an already tight time budget. Thus social interactions inflict at least a time cost. Some clearly have additional costs, others may be beneficial; but for most behaviors, for example, those that aid in identification of future playmates or mating partners, we can only be even more speculative about potential benefits.
In general, mothers' behavioral interactions have traditionally been classified primarily on the basis of presence or absence of threats ' or overt aggression by the actor. On this basis, various authors refer to mothers' avoidance of "friendly" approaches. Handling, grabbing, and pulling of the infant are usually considered "interest" and ''investigation" rather than "aggression" or "threat." As suggested by the analyses in the present study, if we wish to understand the experiences of motherhood and infancy and to predict the consequences of these experiences, it may be more fruitful to consider interactions from the standpoint of the mothers and their infants rather than from the standpoint of their interactants.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 44
Maternal Care in the Postnatal Period
THE FIRST TWO MONTHS of life--the first few weeks, in particular--constitute a period of intense dependency for the infant that produces additional stress for the mother. Although baboon infants cling to their mothers from the day of birth, most infants need some clinging :. assistance in the first few days, some for a week or more. Individual differences in maternal response in the first weeks of life may affect the likelihood of survival of this latter group and of other high risk infants.
Parturition
The need to keep up with the group during the day seems to be a particular strain for new mothers and may be the major advantage of nighttime births, which offer at least a few hours of rest after parturition. Most primate infants are born at night or during the very early hours of the morning. For savannah baboons this means that births usually occur when the group is in the sleeping trees. When we arrive on the morning of a birth, we often find an infant with its hair still wet and matted, and its mother with bloody hands, mouth, and perineum. In these cases the umbilical cord is usually still quite light in color, thick, and pliable. After a few hours mother and infant are dry and the cord is dark and stringy with little flexibility. Probably these infants were born soon before our arrival. In other cases the appearance of mothers and infants when we first see them in the morning suggests that birth occurred at least several hours earlier than that, perhaps late the previous evening or night.
Mom went into labor with Moshi the evening of 31 May 1976, reaching a stage of labor in which she alternately jerked her limbs and rested stretched out on her side on the ground after all other group members had ascended the sleeping trees. Finally, as it became quite dark, Mom too ascended the sleeping trees, and I left the group when the baboons could no longer be discerned. By 0500 the next morning Moshi and Mom were dry, Moshi probably having arrived the previous night.
Occasionally infants are born during the group's day journey, as were Juma and Peach during this study. Judy appeared with Juma in mid-morning after not having been seen by the observer for about an hour. By the end of that day and after considerable traveling by the group, both mother and infant appeared even more fatigued than most females on the day of parturition. Plum was more fortunate; the group traveled little after Peach's mid-afternoon birth and Plum was able to rest overnight before a long journey on the following day.
Descriptions of one or more primate parturitions are gradually accumulating, primarily from captive animals of various species (see, e.g., Abegglen and Abegglen 1976, Bowdent et al.1967, Goswell and Gartlan 1965, Gouzoules 1974, Hopf 1967, Love 1978, Nash 1974). These reports indicate considerable variability, even within species, in the extent to which mothers separate themselves from the rest of the group and in the extent to which other group members are attracted to a birth. We have observed comparable variability in the several births we have seen (in one case we observed the entire period of labor and the birth). The overall description in the literature of labor and birth corresponds fairly well to Mom's and Plum's labor.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 45
The Neonatal Period
All baboon mothers, even primiparous mothers, seemed automatically to clutch their infants from birth and to respond to their neonates, distress cries with such "embraces." During these first few days of life the most conspicuous difference that I observed between primiparous and experienced mothers was in their subsequent responses. Rates in focal samples were too low for statistical analysis; descriptions are based on focal plus ad libitum sampling. If an infant was already in contact and being held or cradled but the infant cried or gave rooting responses in an attempt to reach the nipple, an experienced mother hands usually lifted the infant higher on her ventrum by means of repeated embracing movements, as a result of which the infant usually reached the nipple, clamped on with its mouth, and ceased its distress cries. Not only did the infant obtain its essential nutrition this way, but it was provided with a fifth anchor-point when riding, an advantage that seemed appreciable in the first few days of life. In contrast, primiparous mothers seldom gave repositioning or sucking aid even when, in the most extreme situations, the infant was carried upside down and backward, and required constant embracing by its mother. Infant contact seemed to be one undifferentiated state to these mothers. One often sensed that they were unresponsive to, or puzzled by, their infants' continued distress, despite the fact that nearby group members responded to the cries by watching the pair, moving closer, and in creasing the repeated grunts that are given to mothers and infants. Most mothers learned quickly, though: increased responsiveness sometimes was observed by the end of the first day and certainly in the next few days.
Only in the case of the one totally incompetent mother, Vee, did the incompetence seem to have serious consequences. The fact that her infant, Vicki, was so poorly treated on her day of birth could not be compensated for by the mother's appreciable improvement by the second day and relative competence later on. The early deprivation seemed to be the most likely cause of Vicki's death in the fourth week of life. The recent research of Mason and his colleagues has emphasized the superiority of a responsive "attachment figure" to the immobile surrogate originally used in their laboratory rearing studies of rhesus monkeys. However, even the most unresponsive wild baboon mother is probably more appropriately responsive than the researchers' most sophisticated surrogate (a dog). It remains for future field and laboratory research to document more adequately the normal range of responsiveness and to determine the consequences of various levels of normal responsiveness both for survival and for such deficits in development as the cognitive deficiencies.
Mothers persist in the apparently automatic embracing of their infants even after infant death. They continue to carry the decomposing and increasingly dehydrated corpse, despite the fact that this usually means that they walk three-legged, setting the corpse down whenever they stop to feed and then retrieving it again, surely a tiring and difficult way to forage for several days. During 1969, one mother in an other group often carried the corpse of her infant in her mouth or walked on all fours, dragging it on the ground. After several days the head of the corpse was gone and the corpse was quite dehydrated. On the fifth day only some dry extremities remained. As the mother sat she began to gnaw on the hard dry piece in her hand, occasionally looked at it, seeming "puzzled," and eventually stopped gnawing it, clutched the scrap again and walked off. This is the only time I have seen such behavior. After about three days other mothers leave the corpse on the ground for gradually increasing periods of time while they forage at greater distances away, eventually either lose it or leave it, looking back at the corpse with repeated signs of conflict and am bivalence and sometimes giving alarm barks. The two corpses I have recovered after desertion were both badly decomposed.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 46
Maternal Style
By the end of the second week of life, all healthy infants climbed clumsily about in the mother's ventrum and made their first attempts to break contact. This was a period of close maternal attentiveness when ever mother and infant were not in contact and of rapid return to contact at the slightest disturbance of any sort. These first stages of separation were initiated primarily by the infant and were sometimes limited by the mother. Although most mothers were protective of their infants when intruders approached during the first months, mothers differed considerably in reaction to infant exploration during this period. The most restrictive mothers, Handle and Slinky, allowed virtually no break in contact for almost two months. The more "laissez-faire" mothers tolerated separation, although they themselves seldom moved away from their infants during the first month. These laissez-faire mothers watched or followed their infants less often, even when the infants were quite young.
By the end of the second month most mothers initiated some separation, and the infants oriented to and followed their mothers. During month three restrictiveness and overt maternal attention to the infant rapidly disappeared except during emergencies, whereas infant attention to and following of the mother became the. This was accomplished partially through one of the few examples of what I would call teaching in baboons. A mother began to take a few steps away from her infant, paused, and looked back at the infant. As soon as the infant began to move toward her, she again moved slowly away. At first this sequence was repeated every few steps, but soon a mother seemed to be able to initiate a long bout of following by just one such pause. Hinde and Simpson have described and well illustrated this behavior for caged rhesus macaques, labeling it "mother infant leaving game" . Basically, mothers could be characterized dichotomously in the first two months as being either in the range rejecting to laissez-faire or the range protective to restrictive in their behavior toward their infants. In comparing the first group (hereafter called "laissez-faire" mothers) with the second (hereafter called "restrictive" ones), we find that the laissez-faire mothers not only restrained their infants less and completely stopped doing so when their infants were younger--the main basis on which the initial classification was made--but that they also rarely if ever followed their infants. They seldom made contact with their infants, and at a younger infant age they increased the distance between themselves and their infants more often than they decreased it. At an earlier age they ignored their infants or directed punitive behaviors toward them when the infants attempted contact or suckling.
Seven of the 12 mothers for whom there were at least some observations during month one were classified as laissez-faire, 5 as restrictive. The average dominance rank at the time of parturition was 6.1 for the laissez-faire mothers, 11.2 for restrictive ones (recall that by convention the top-ranking animal is assigned the number 1, the next isnumber 2, and so on). Six of the 7 higher-ranking mothers were laissez-faire; 4 of the 5 lower-ranking ones were restrictive. Three of six female infants and two of five male infants had restrictive mothers. Mom, the only mother who was observed during month one with each of two successive infants, was laissez-faire with both female Misty and male Moshi. Thus maternal rank was a good predictor of maternal style; sex of infant was not.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 49
Infant Development and Mother-Infant Spatial Relationships
WE ARE NOW READY to turn to the infant itself, its development and the progress of its independence from its mother during the first year of life. Again, I shall consider individual differences as well as those features common to all infants. I shall examine the effects of ecological variables, maternal time budgets, and maternal style on mother-infant relations and independence. First, though, I shall consider the general pattern of infant maturation.
Physical Maturation
Baboon infants change appreciably in both appearance and behavior over the first year of life. At birth the infant has skin that appears bright pink and a natal coat of black hair along with disproportionately prominent ears, and the males have a particularly visible pink penis. This coloration contrasts strikingly with the adults' yellow-brown coat and grayish skin.
During the third month of life healthy infants all developed some gray skin pigmentation in their hands and feet and some gray mottling in their paracallosal skin, as well as some gold hair in their coat, particularly around the wrists and brow region. By month four Sesame and Pedro were noticeable for (1 ) absence of gray skin coloration, (2) loss of hair on tail, and (3) absence of gold hair. Their skin color change occurred later than did other infants, and not only was their coat color change delayed, but the hair was thin and white or beige when it did change. These abnormalities may be related to nutritional deficiencies, and it is notable that they became obvious at about the age that caged baboon infants whose mothers were fed a low-protein diet dropped below the range of weights for infants of mothers on normal laboratory diets.
By six months of age infants have reached a number of very visible milestones. Virtually all of their natal coat and most natal skin color have been replaced by skin and hair similar in appearance to that of adults. However, a few tinges of pink remain in the muzzle and ears almost until the first birthday, and the male's scrotum remains pink for at least another year after that.
At birth infants' locomotory abilities are quite undeveloped. During the first weeks objects that baboon infants put in the mouth are picked up directly with the mouth rather than with the hands. Their hind limbs are even more flexed than their forelimbs and their gait is extremely wobbly during the first few weeks. Locomotory attempts usually end with the infant falling over after only a few steps. When not clinging to their mothers, neonates sit hunched over or flop on the ground. It also seems to take a few weeks before the infant recognizes its mother and selectively orients and clings to her in preference to other animals. This lack of selectivity may make infant snatching especially dangerous in these early weeks: at this stage infants cling tenaciously to whoever carries them.
Initially, infants ride only ventrally. Soon, however, some attempt to climb up the mother's legs to her back. Fred, Alice, Ozzie, Eno, and Pooh all rode dorsally, although Pooh had considerable difficulty doing so until she was about eight months old. Fred, Ozzie, and Alice all did so with ease by three months of age, as did Eno. Summer, Grendel, Sesame, Juma, Safi, and Moshi all rode dorsally by two months of age, Oreo a little later. However, Pedro, Bristle, and Hans never did ride dorsally, nor did Misty by the time of her death at a little over two months. We have less systematically observed this variability in dorsal riding in other Amboseli baboon infants. It does not seem to be a useful age or developmental marker, as was indicated at first. Nor is any strong gender difference apparent.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 50
Physical and behavioral maturation proceed at different rates for various characteristics. Infants became quite competent in locomotion on the ground by the end of the first month of life, and during month two they mastered climbing over small logs and other obstacles. By this age, too, they were able to manipulate the plants and other objects they found on the ground, no longer getting these to their mouth just by mouthing them.
During the third and fourth months of life, infants began to spend increasing amounts of time in peer interactions, particularly play, though most did not stray far from their mothers to do so. At this age they began to feed on particularly accessible foods, could negotiate fairly well within trees, and could climb into and out of some umbrella trees but could not climb any of the fever trees, perhaps because of their smooth bark. They were still dependent on their mothers for virtually all food and transportation, but increasingly it was they who attended to their mothers, who maintained contact and proximity, rather than vice versa.
For the second six months of life, my data for behavior outside the mother-infant pair come primarily from ad libitum observations, checks of physical maturation, and the focal samples of mothers (an infant's own mother and others). A more complete picture of infants' interactions with others must await the more detailed studies of the older infants themselves. By six months of age, umbrella trees were negotiated with ease as were the lower and more branched of the fever trees. By eight months infants descended alone from even the tall, unbranched trunks of the fever trees used as sleeping trees.
During months 9 through 12 dominance relationships began to be evident among the infant dynds. These older infants also lost their general immunity from spatial displacement and other aggression by juveniles, and they could not climb on or sit immediately next to many adults with impunity. Older infants still slept with their mothers but usually had to descend the sleeping trees alone in the morning. By its first birthday every infant was quite independent and perhaps could have survived its mother's death (although three infants 17, 20, and 21 months old were the youngest known surviving orphans we have had). The mothers of approximately half the infants in Amboseli resumed estrus cycles when the infants were 8 to 12 months old, most others a few months afterward; mothers then became pregnant during the off spring's second year of life. However, those young juveniles whose mothers had not yet become pregnant retained occasional daytime nipple contact and continued to sleep at night in the mother's ventrum and probably on the nipple, the oldest observed individual being a two-year-old male, Jake. Despite its increasing independence, an infant's primary relationship throughout the first year is with its mother. I shall now examine in more detail the development of mother-infant spatial relationships through the first year and the changing contingencies between the behavior of mother and infant; then rejection and weaning will be considered.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 52
Infant's Use of Space
The young infant's world can be viewed as a circle with its mother at the center. We might, then, ask what percentage of its time does an infant of each age spend within circles of different radii? Because I do not have continuous records for distances greater than arm's reach, I used the point-sample records, estimating the probability of an infant being at various distances from its mother by the number of point samples at that distance divided by the total number of point samples. Monthly percentages were calculated for each infant and were then averaged over all infants. Related data through month eight are presented in such a way as to highlight individual differences. The expansion of the infant's physical world is abundantly clear from these data.
Over the first six months of life, contact time dropped steadily from 100 percent in the first two weeks to about 32 percent in the sixth month. After the first two weeks, time spent out of contact but within arm's reach remained at about 8 to 10 percent of total time (but it was a decreasing proportion of the noncontact time). Time spent at each of the greater distances showed successive increases. By the sixth month, approximately a third (32 percent) of the time was spent more than 5 meters away, another third (36 percent) out of contact but within 5 meters; whereas during month two, only 5 percent of the time was spent more than 5 meters away, another 20 percent out of contact but within 5 meters.
Although infants made increasing use of space more distant from their mothers, their use of this space was not at all independent of the position of their mothers. Employing a model of mother-infant distance based only on available area and taking 1 meter as a mother's arm's reach, one would predict that infants would spend approximately three times as much time in the zone between arm's reach and 2 meters as in the zone between contact and arm's reach, seven times as much between 2 and 5 meters as between arm's reach and 2 meters. Yet at no time during the first year did the actual distribution even approach the random one. Always, infants used space nearer their mothers appreciably more than expected.
I took as a criterion the distance from the mother that accounted for 90 percent of an infant's daytime. For example, during month two, a radius of 2 meters from the mother on the average accounted for 90 percent of an infant's daytime, but for Grendel the radius required was 5 meters, for Moshi and Misty (Mom's two successive infants) it was between 2 and 5 meters, 2 meters for both Juma and Safi, arm's reach for Pedro. These are all the infants of laissez-faire mothers. By contrast, all the infants of restrictive mothers were within arm's reach 90 percent of the time during month two.
By the end of month eight all infants spent at least 40 percent of their time more than 5 meters from their mothers. By month six for most infants, and by month eight for all, the distance from their mothers required to encompass 90 percent of their time was no longer a useful measure-- was over 20 meters, and I often could not locate the two at the same time.
During the second half of the first year of life, contact time declined from 30 percent at month six to 10 percent by about month ten. Younger members of this class rode during part of long day journeys. Otherwise, contact at this age occurred primarily during external alarms and during a long, dozing nursing bout, commonly in the early evening, at times when mothers were either resting or involved in grooming interactions with others.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 53
Spatial Relations: Dynamics
During the focal samples, I recorded the time and the identity of the individual who effected (performed) each change in the mother-infant spatial relations (classified as contact, arm's reach, greater than arm's reach) whenever a change occurred from one of these states to another and that same actor did not immediately (within 0.05 min) again change state; if the actor did immediately change state again, the 0.05 minute and actor identity criteria were again applied. For these spatial-state records four categories of actors were distinguished: focal mother, her infant, mother and/or infant but I could not tell which, and "other" (with the individual identified). Over 90 percent (usually well over 95 percent) of all transitions were effected by exactly one member of the mother-infant dyad. Transitions for which I could not distinguish between the mother and infant as actor accounted for less than 1 percent of the transitions for most months, with higher percentages occasionally occurring at times of group alarm or during months of highest rates of state changes. During the first two months of infant life, "others" effected 2 to 5 percent of the transitions for infants of laissez-faire mothers by carrying the infant. For restrictive mothers total rates of transition were too low during these early months to permit reliable estimation.
On average, 70 to 80 percent of all changes in spatial state that were made by one member of the mother-infant dyad were made by both the infant. Infants made over 85 percent of the breaks in contact during the first five months, 60 to 80 percent thereafter. They made 50 percent of the contacts during month one, over 90 percent after month three. Again, however, the pattern of individual differences reveals that Grendel, Juma, and Safi were making 90 percent of the contacts by month two: Bristle, Hans, and Sesame not until month four or later. Data for Misty are quite variable, probably because of her poor health. Moshi made 61 percent of the contacts in month two, 99 percent in month five; no data are available for him for months three and four.
Data on changes from one of the three spatial states to another were then reanalyzed by collapsing the categories into increases or decreases in the distance between mother and infant: both a change from contact to within arm's reach or to greater than arm's reach and a change from within arm's reach to greater than arm's reach were considered "increases"; changes in the opposite direction were considered "decreases." The percentage of decreases and the percentage of increases that are due to the infants are graphed for the first year of life. These provide estimates of the conditional probabilities that a decrease (and increase) will be effected by the infant, given that it was performed by only one member of the mother-infant dyad. These are approximately the %A and %L of Hinde and his coworkers, but these workers did not include transitions between contact and "within arm's reach" in their definitions of "approaches" (A) and "leaves" (L). In a field situation, with the animals usually moving along while foraging, I found it important to include these classes--it appeared that contact and arm's reach were quite different to the animals. However, for comparison with the other studies I also reanalyze these data by Hinde's "approaches" and "leaves" in a later section.
As noted earlier, the first stages of independence were characterized by infants primarily increasing the distance between the two and mothers primarily decreasing it; that is, the earliest stages of spatial independence were infant initiated. Only later did mothers reverse the dynamics of the spatial relationships, increasing the distance between themselves and their infants more than decreasing it. This stage was reached later for the restrictive mothers than for the laissez-faire ones. By this stage mothers had stopped following their infants; rarely if ever paused, looked at, or waited for them; and usually ignored their distress cries as they (the mothers) moved away.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 54
Clearly, not only the identity of the member of the pair making the transition but also the transition rate changed appreciably as the infant matured. In particular, the peak transition rates occurred soon after the time that infants switched their role and began decreasing distance to their mother more than increasing it. It is at this age that the observer has the impression that infants are frequently following or chasing after their mothers, sometimes even screeching and "throwing tantrums". I shall return to discussion of this "weaning" stage in the next chapter.
It is interesting to note that not only did the laissez-faire mothers leave their infants more than did the restrictive mothers, but their infants also left them more often, owing at least partly to the absence of restraint. Thus very different interactional patterns and rates of independence were established within dyads during the first month of infant life. Infants as well as mothers surely contributed to this interactional pattern, and most changes in spatial relationships were actually effected by the infant; but the data from this study suggest that differences between mothers were the major or initial determinant of differences between pairs.
Because infants differ in the age at which transition rates peak, the averaged data produce a flatter curve for monthly transition rates than do the data for individual infants. The pooled data thereby mask the appreciable developmental change in transition rates. This is illustrated by taking only the data for those six infants (three male, three female) for whom there are data from a sufficient number of continuous months to identify the peak month of transition rates. Peak transition rates occurred during month four for Safi, Sesame, and Grendel; month five for Hans; and month six for Summer and Bristle. Aligning the data for these infants by these peak months rather than by age of infant, we obtain a mean of 72 transitions per 100 minutes for the peak month. The time course of each infant's development probably corresponds more to this pattern than to the age-determined. I do not know whether it is a coincidence that the peak transition rates for the three female infants were all higher than any of those for the three males.
By month 7, infants were doing about 90 percent of the reductions in distance and 50 percent of the increases in distance between mother and infant. That is, when mother or infant moved closer to the other, nine times out of ten it was the infant who did so, whereas when they moved apart, the mother was as likely as the infant to do the moving. No clear change in these percentages emerged after this time, with decreases in distance due to the infant fluctuating slightly around 90 percent and increases in distance fluctuating somewhat more about 50 percent. These fluctuations are probably due to the small sample sizes for these later months: not only did I sample for fewer days per month and have fewer animals for months 10 to 12, but rates of transition dropped sharply after month 7, remaining below 30 transitions per 100 minutes throughout the rest of year one. Thus it is the rate of transitions, or changes, in mother-infant proximity rather than who effected these changes that best characterizes the dynamics of development within the second half of the infant's first year.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 55
Infant Contact as a Contingent Behavior
In the previous section I treated an infant's contact with its mother as a state that was independent of the mother's other activities. In fact, I found that the probability that an infant would be in contact was dependent on a mother's current activity, and this contingency itself changed appreciably during the infant's first year of life. I plotted the probability of contact in two different maternal activity states: grooming or resting, and walking. Young infants, up to about four months of age, were disproportionately out of contact when their mothers were resting or engaged in grooming interactions and disproportionately in contact when their mothers were walking. For infants over seven months of age, the relationship was just the opposite: infants were much more likely to be in contact when their mothers were resting or grooming that when their mothers were walking. Another way of viewing this reversed contingency is to examine the probabilities that a mother was engaged in various activities, given that her infant was in contact. For mothers of older infants the probability was over 90 percent that the mother was resting given that her infant was in contact at the time, this despite the fact that mothers spent only about 25 percent of their total daytime resting or in grooming interactions.
The data on contact, when averaged for all infants, mask the individual variability and damp the maturational effects, as was true for the change in rates of spatial transitions discussed in the previous section. Again, this occurs because the form of the graph is essentially the same for each infant, that is, each infant goes through the same stages, but the time course varies the reversal of contingencies occurs at different ages for different infants and, in particular, occurs later for those infants whose mothers were described above as being restrictive where the data are partitioned by maternal style. Having identified a contingency between a mother's immediate activity and the probability that her infant was in contact, we might then ask whether at some ages a mother's time budget for the day determines the amount of infant contact that day: that is, whether there are conditions under which a mother's activities are a limiting factor in determining infant contact.
Ecologists consider a resource to be limiting for some process if increased abundance of that resource results in growth or increase of the process. The fact that an organism utilizes are source or that organisms preferentially seek the resourceor clump around it does not mean that the organism would be more abundant if only more of the resource were available. In the case of maternal contact as a resource for an infant, the fact that at some age infants are disproportionately in contact when their mothers are resting does not tell us whether infants would be in contact more at times of the year or in locales (e.g., zoos) in which mothers spent more time resting, or conversely whether differences found between situations or habitats can be accounted for by differences in time spent in the various maternal activity states in the several situations. Even though an infant's contact with its mother is contingent on its mother's immediate activity, that activity may or may not be a factor limiting or determining the amount of contact an infant has with its mother during a day: each day there may always be enough resting time in total so that available resting time does not limit the total amount of mother-infant contact time for the day, even for older infants. When a very young infant has been out of contact with its mother, it resumes contact virtually every time its mother moves more than a few steps. At first this happens by the mother scooping the infant to her ventrum, later by the bottom infant itself joining its mother, as discussed previously.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 56
By the infant's third or fourth month, it uses its mother for transportation primarily if the group progression is quite rapid or the journey has been very long that day. Thus we would predict that a very young infant would be in contact more on days when its mother walked more and a somewhat older one would be more influenced by the length of the day journey. An infant more than about seven months old would be expected to spend disproportionately little time in contact on days when its mother walked a lot. For each sample day on each infant, I calculated from the continuous records the percentage of time that the infant spent in contact with its mother (as described previously). For most infant ages this value is very close to that estimated from the much more easily obtained point, or instantaneous, samples. From the point-sample data I estimated the percentage of time the mother spent in each activity that day; from the demographic data we know the age of the infant; and the length of the day journey was calculated from the day journey maps. Using SPSS Scattergram, I then examined the effect of day journey length and time budget on the amount of time that infants of each age (by months) were in contact with their mothers, using as the dependent variable the contact residual, that is, the deviation in observed contact time for that day from the contact I time predicted for that age (by day) from the linear regression.
When infants were in the first month of life, they were in contact so much of the time that existing variability in time budget or day journey did not affect the time spent in contact. Likewise, for infants over seven months of age, who were, overall, in contact less than 15 per cent of the time, there was only 1 of 15 regressions that was "significant" at .05 level about what would be expected by chance. However, during intermediate ages daily contact time was significantly related to maternal time budget. Infants were in contact more on days of long day journeys or when their mothers spent more time walking; they spent less time in contact on days when their mothers spent more time feeding or resting. Day journey length or time spent walking usually explained about 15 percent of the daily variability in contact residuals during these months. We can tentatively conclude that maternal time budget is to some extent a limiting factor determining the degree of independence at intermediate ages.
For young infants these results are as expected at ages when infants were disproportionately in contact while their mothers walked the time a mother spent walking during a day was a partial determinant of contact, and therefore noncontact, time for that day. At ages for which infants were neither disproportionately in or out of contact during walking, it was day journey length that became important. Surprisingly, the expected negative relationship between time spent walking and mother-infant contact did not occur for older infants, and it is not just that the correlation failed to reach significance--a fact that could be considerably influenced by sample sizes for those months. The results even fluctuated in direction for these older infants. A finer-grained analysis, involving more infants, will be needed to elucidate the factors affecting this period, but at this time it appears that mothers' activity budgets do not limit the contact time of older infants, but do limit that of younger ones.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 57
From a causal standpoint I would like to know whether mothers alter their behavior as a result of their infants being in contact or whether infants change to (or from) contact with their mothers as a result of the mothers' activity. Ideally, I would need to have continuous rather than point-sample data of the maternal activities as well as of infant contact so that I could examine the lag time to a transition in infant behavior given a transition in maternal activity and compare it to the overall lag time in such transitions. If it is infants who are adjusting their behavior to mothers, the contingent lag times would be shorter after a transition to a contact-stimulating activity than it is in general. It would be easy to do such an analysis by using CRESCAT, but I was not able to collect the continuous records for most maternal activities; and I therefore must be content for the time being to consider the suggestive evidence from ad libitum data and from rare events. These data suggest that the accommodation is primarily by the infants rather than by their mothers. An infant repeatedly hopping on and off its mother's back interferes with the mother's feeding and walking, especially if the infant tries to ride ventrally rather than dorsally. I often saw mothers of 4- to 5-month-Olds rebuff contact attempts made while the mothers foraged. Sometimes, after being rebuffed the infant, either immediately or after it "threw a tantrum," rode quietly on its mother and the mother no longer pushed it off. In similar instances, mothers of older (6- to 8-month-old) infants dislodged their infants while walking. The dislodged infant then followed its mother, sometimes throwing a tantrum, and the mother eventually sat and then immediately embraced the infant to contact and a suckling position. In these instances, the persistence of the infants did seem to shorten the time to a change in maternal activity, but the change had to occur before the infants were allowed to make contact. More dramatic cases occurred with 8- to 12 -month-Olds. Sometimes I was following a mother during a sample and could not locate her infant at all. The mother began a grooming interaction or resting and immediately the infant would dart toward her from an activity over 20 meters away, make contact, and begin suckling.
Summary and Discussion
A baboon infant begins its first year of life with a distinct natal coat, undeveloped motor abilities, and a total dependence on its mother. By the end of that year, its appearance is distinguishable from that of adults primarily by size, it can perform most adult behavioral patterns, and it probably could survive its mother's death. However, it still occasionally goes to its mother in times of alarm and for rare nursing bouts and still sleeps in her ventrum, perhaps suckling, in the trees at night.
The role of the natal coat is of interest at various stages of maturation. This conspicuous coat probably does not attract predators as long as the infant spends most of its time in its mother's ventrum. Once the infant rides dorsally or runs about independently the less conspicuous brown-yellow adult pelage is probably safer. Another speculation is possible regarding the change to adult pelage. It has often been suggested that the function of a conspicuous neonatal coat is to clearly mark the infants for more tolerance and protective treatment by conspecifics. Yet, if this coat is one of the major stimuli that attracts other group members, as it seems to be, and if the attraction places an additional burden on mothers, as I indicated, the change to adult pelage may be of value for mothers by reducing social pressures at a time when nutritional and ecological pressures are becoming extreme. Proposals for both the advantages (benefits) and disadvantages (costs) of infant pelage remain speculative.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 58
Contact time between mother and infant, which was 100 percent just after birth, declined at a rate of about 8-9 percent per month over this first year, with infants of restrictive mothers consistently spending more time in contact with their mothers than did their peers at the same age. Illness also resulted in greater contact between mother and infant. However, infant gender did not affect the amount of time an infant spent in contact with its mother. Although maturing infants increasingly spent time at greater distances form their mothers, at all ages they disproportionately used areas close to their mothers.
A few studies report data with unbiased estimates of time that maturing infants spend in contact with their mothers during the day. They include Struhsaker's for vervets in Amboseli, Berman's for the provisioned rhesus colony of Cayo Santiago, and Konner's study of !Kung in the Kalahari. The more slowly maturing baboons spend more time than the vervets or the rhesus, and the human children spend much greater time in contact than all the others, especially in the second six months. As Konner points out, the limited data for European and American children, both home- and institution-reared, indicate values of mother-infant contact that are much lower than any of these, usually less than 25 percent of the time in contact even for quite young infants.
At every infant age changes in spatial relations between baboon mother and infant were made by the infant more often than by its mother. During the first few months of life, however, infants increased the distance between the two and mothers decreased it. At this time infants were about four times as likely to leave their mothers as their mothers were to leave them, with considerable individual variability. Later it was mothers who became leavers, infants approachers, as has also been reported in laboratory investigations of a number of primate species (e.g., Hinde et al. 1964, Hinde and Spencer-Booth 1967, 1971, Jensen et al. 1967, Kaufman 1974, Rheingold and Eckerman d 1970, Rosenblum 1974, Rowell et al. 1968) and by Nash (1978) for a few older baboon infants at Gombe and by Berman (1978) for free ranging rhesus macaques of the Cayo Santiago colony. In the present study mothers and their older infants were equally likely to leave each other, but reunion was nine times as likely to be due to the infants as to their mothers.
To facilitate comparisons I have excluded from the "increases" and "decreases" those transitions that were between contact and arm's reach to obtain in the "approaches" and "leaves"of Hinde and his colleagues. The main difference is that during the first month of infant life most of an infant's increases in distance from its mother take it no farther than her arm's reach. When these are excluded, the infant's contribution to leaving drops from over 80 percent to just over :. 60 percent. Moreover, the young infant's contribution, in general, to both approaches and leaves is about 10 percent less than when transitions between contact and arm's reach were included. Results past the fourth month of life are indistinguishable from those noted previously, with infants making 90 percent of the leaves from month four on and about 55 percent of the approaches from month five or six.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 58
Infants become leavers more than approachers in the second or third month for the baboons in Amboseli, the provisioned free-ranging rhesus of Cayo Santiago, and the caged rhesus in the Madingley colony, a rather amazing amount of similarity in results between species and between settings. The rhesus in both settings reach a constant level of %A-%L at about 12-15 weeks of age, and about a month later for the Amboseli baboons. Also, the asymptotic level is approximately 35 percent in Amboseli, 40 percent on Cayo, and only 10-20 percent in Madingley. That is, infant contributions to spatial proximity stabilized later in wild baboons than in captive or free-ranging rhesus and at the stable level there was much less difference between an infant's contribution to leaving and its contribution to approaching in the captive rhesus than there was in either the free-ranging rhesus or the wild baboons. From the published data it is not possible to analyze further the source of this difference.
Of particular importance is the contingency found between infant contact and various maternal activities. I suggest that this contingency is at least partially imposed upon the infant; that is, the infant must adjust its behavior to that of its mother rather than vice versa (the main exception being the mother's adjustment of feeding postures described earlier). The young infant learns not just that it can be out of contact sometimes but that it can be out of contact at particular times that are determined by its mother's activity. This is probably one of the earliest forms of socialization for an infant baboon and may provide the social sensitivity on which later social integration depends. One of the first areas of infant trauma seems to involve the reversal in these contact contingencies. Older infants must take advantage of their mothers' rest time for any contact that they want rather than using that time for play and exploration. Thus at one level maternal rest time remains the ; same: it is the time to do the "unusual". However, the unusual is lack of contact for young infants and contact for older ones. Thus at the level of immediate behavior it is a reversal. At four to six months of age the contingency reversal seems to precipitate some of the first infant "weaning," or distress, behavior. Next I shall consider various aspect of independence and weaning in more detail.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 60
Weaning and Infant Independence
PRIMATE MOTHERS PROVIDE their young with a number of important kinds of care. Infant development and independence involve the infant's gradual attainment of self-sufficiency in each of these areas. In the scientific literature, as in common English usage, the term "weaning" is sometimes used to describe this overall process of in creasing independence. However, the term also retains its narrow and original reference to only nutritional self-sufficiency from mother's milk and to self-sufficiency that is imposed rather than voluntary. I think it is important, if a bit more linguistically awkward, to consider separately the various areas of care and to leave open to theoretical consideration and empirical study the question of the behavioral source and imposition of independence in each area.
Initially, baboon mothers provide all their infants' nutrition through nursing. They provide all transport through primarily passive carrying of the clinging infant. Protection from disease and ectoparasite is accomplished through grooming and perhaps through restriction of early exploration and of social contacts. Protection from predators is provided at first by carrying the infant away from danger as the mother herself detects predators, responds to detection by other group members, and either flees or joins in group action against predators.
An infant's increasing size and independence affect its requirements in each of these various areas. For the infant, its mother, and for other group members, the cost and the individual's ability to provide care for the infant will be a function of the type of care and the characteristics of the potential provider: there are alternative ways of meeting requirements, alternatives that involve different individuals and different behaviors. Below, I shall outline these parameters for baboons, providing what I hope is the first step toward a complete specification of costs, benefits, and options that will lead to similar research on this and other species.
Transportation
The available evidence in the literature suggests that the energetic cost of carrying an infant is essentially proportional to the additional weight involved. One might at first assume that it is advantageous to a mother to stop providing her infant's transportation as soon as the infant can keep up with the group on its own, and in fact, as noted in the previous chapter, infants spent considerable time out of contact even during travel at ages at which they were still primarily dependent on their mothers for food. However, a small animal in general expends more energy than a large one in covering a given distance in a given amount of time. Therefore, if a mother is the sole source of nutrition for her infant, it may cost her more if the infant becomes semi-independent and provides some of its own transport through locomotion than it would cost her to carry it. We cannot assume that apparent independence is advantageous to a mother and costly to her infant. The true situation may be the opposite of the apparent one.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 61
Carrying is an area of care in which other group members could readily substitute for mothers, a behavior that occurs to varying degrees in other primate species including humans. Provision of transport by other group members is relatively rare in savannah baboons, although, as noted elsewhere, in the most extreme case of maternal rejection during this study (Gin's rejection of Grendel), the infant spent considerable time riding on adult male High Tail, and after High Tail's death unsuccessfully sought rides from other group members until many months later he began to be carried by an infertile adult female, Lulu. Among those whom one would most expect to provide care are siblings, but they are I quite likely to be immature in this slowly maturing species. A six month-old infant would impose an appreciable additional energetic cost on a half-grown juvenile sibling three years old. Four- to five-year old siblings are probably large enough to comfortably provide some transportation for infants. The only members of this age class during 1975-76 were immigrant male Nog and males Toto and Dogo, neither of whom carried their siblings, Eno and Pedro. Almost all adult female relatives will be either pregnant or supporting their own infants. This leaves subadult and adult males as the most likely candidates for such care, and the adults do provide some. Perhaps the adults would provide more infant care in a more structured mating system than that found in these baboons, that is, one in which individuals are more (predictably) closely related to any particular infant in the group. It is interesting in this context that adult male hamadryas baboons, who maintain long-term polygynous bonds, establish their first harem relationships by providing much of the transportation for young juvenile females. Immature hamadryas probably require carrying at an older age because the groups travel longer distances in a day than do Amboseli baboons. It seems likely that for any primate species the amount of observed kin involvement in infant care at any particular time will depend on the age, sex, and kin structure of the social group at that time and on the other demands placed on potential helpers.
Nutrition
As indicated previously, caloric requirements to maintain weight are probably proportional to weight to the three-quarters power. However, a growing infant also requires additional calories for the production of new tissue. Additionally, to the extent that an infant is more active when it is not in contact with its mother, to that extent caloric requirements will be greater in order to maintain the increased activity level. Thus from the standpoint of a mother who is providing all her infant's nutritional requirements, it is preferable for the infant to remain totally dependent, in passive contact, rather than to be semi-independent, running about.
Would a mother gain as much advantage for her infant, at less cost to herself, if she allowed it to be active and out of contact and to obtain some of its food independently? Only if the assimilable calories the infant so obtained were greater than the infant's increased calorie expenditure in obtaining those calories and in running rather than being carried (see "Transportation," above). Although this is obvious, the practical implications may not be. As I discussed previously, many foods just are not accessible to harvest or to assimilation at some times , of the year or before infants are at certain stages of physical development. Thus an infant's physical development and the availability of appropriate weaning foods determine whether alternatives exist, alternatives that would be advantageous to the mother as well as to the infant. In the short run, it could sometimes be more costly to the mother, Just in energy considerations, for an infant to try to obtain some of its own food than for the mother to provide it all through lactation. It is possible, of course, that there are some essential nutrients that the infant requires past a certain age that are not obtainable from milk and that are available in weaning foods. These and similar constraints favoring either milk or other foods would have to be taken into account I in complete analysis of the economics of weaning.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 62
Two other alternatives are used by some other animals to provide food for young that are unable to provide it for themselves--harvesting and/or partially processing foods for the infants. These systems have the advantages that other group members can share with the mother in the provision of food for the young and that perhaps less energy is lost than if the food is converted to milk. Aside from humans, most vertebrates that use these feeding systems have two common features--first, the use of a den, nest, or other predictable location where the young are kept and to which the feeders return with food, and second, the exclusive use of animal rather than vegetable foods. Perhaps vegetable foods are not as suitable for animals to harvest for other, individuals unless the food can be cooked, as it is by humans. Certainly, for a baboon group, which forages over a long day journey and feeds throughout the day, such a feeding scheme would be more difficult. Perhaps these are some of the reasons that no such feeding system has developed in baboons, but the answer is not totally satisfying, and this seems to be an area that warrants further study. The only way that group members provide food for a baboon infant, other than its mother via lactation, is through tolerance of infants at a food source, the result of which is that infants obtain scraps of foods that they themselves could not obtain by their own effort. This more subtle, passive form of food sharing may be a very important one for a number of animals and is one that has received less attention than more dramatic forms just because of its subtlety. This is a common problem in behavioral research--dramatic behavior receives more attention.
Disease and Ectoparasites
It would at first seem that an infant's need for grooming would increase more slowly than its nutritional and transport requirements because surface area increases more slowly than does weight or volume. However, an infant that is semi-independent, that is close to the ground and is running and playing in shrubs and grasses, probably has a higher density of ectoparasites than one that rides on its mother. The time required to free a semi-independent infant of ectoparasites is therefore probably greater, proportional to the infant's weight, than the time required for a totally dependent infant. Again, we have the suggestion that there are certain advantages to a mother in having her infant clinging all the time rather than being semi-independent. The infant itself cannot readily do its own grooming: it is not physically able to perform complete grooming manipulations until at least eight months of age, and there are many parts of a baboon's own body that even an adult cannot reach to groom. Grooming appears to be a function that could easily be assumed by animals other than the mother. Others can and do groom infants, infants of laissez-faire mothers in ill particular, but most of a young infant's grooming is done by its mother, often at the same time that the infant is suckling. Perhaps this cements the bond that leads later to the considerable reciprocity of grooming relationships that is seen among juveniles and their mothers. That is, mothers may provide most of their infants' grooming in part to obtain even more grooming in return when their infants mature.
To the extent that exposure to disease is a function of environmental and social contacts, it is also a function of time spent away from the mother, but probably is otherwise essentially independent of infant age or size. Mothers cannot effectively control their infants' exposure when they are out of contact and can only partially do so when they are in contact. While they are in contact, mothers provide some regulation of social interaction, as described in previous chapters. However, I have not seen mothers limit, for example, the nonfood items that their infants mouth from the ground, especially in their first weeks. Note that it may be preferable for baboons to be exposed to certain diseases as infants rather than as adults, as with the so-called childhood diseases in humans. Again, it may prove difficult to compare properly even relative costs or benefits of behavioral alternatives.
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I would appreciate it, if you could send information about Baboon Mothers and infants together with pictures for my 8 year old son. A topic he is studying at present in School.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 63
Protection from Predation
With protection from predation as with most of the areas of care that I discuss, it is harder, not easier, for a mother to provide care if her infant is not in contact, no matter what the size of the infant. That is, the same benefit to the infant is more costly for the mother to provide if her infant is not in contact. A mother must first locate and make contact with her infant before carrying it in flight. Additionally, if the two are separated, it would seem that infants can more readily keep track of mothers are larger, generally slower-moving individuals whose activities and location are probably more predictable. Thus maintaining in formation about the other's location is probably done at little cost by the infant but at high cost by its mother. This theoretical speculation was supported by several kinds of observations. I already described the sudden appearance of older infants when their mothers began to rest. Additionally, in alarm situations, I commonly saw mothers of semi-in dependent infants stop, look about, appearing unsure of their infants' location, and then wait as their infants dashed out from a bush or tree (I had not been able to locate the infant) and approached. The mother sometimes met the infant as it came close and in every case she embraced the infant to contact, and then dashed away. As infants stray farther and farther from their mothers, even this cooperative form of , flight probably becomes inefficient for both. We do not know at what age an infant can run, for at least short distances, faster than its mother, or at least faster than its mother can run while carrying the infant. Also, by seven or eight months of age infants can climb even fairly tall fever trees. Ten- to twelve-month-olds were often seen to run alongside their mothers in flight when the group fled from a predator or local Maasai tribesmen. It was also at these times of emergency that particular associated adult males were seen to carry infants and young juveniles.
"Weaning": Maternal Punishment and Infant Distress
In the present study detectable aggression (usually biting, pushing, grabbing, or hitting) directed by a mother toward her infant was very rare, for most mothers and most infant ages, less than one act per 100 minutes. Only Gin and Spot aggressed against their infants during month one; Gin in month two; and Gin, Spot, and Preg during month three. Yet by month five all mothers had bitten, pushed, grabbed, or hit their infants. Maternal aggressive or punishing behavior was commonly followed by gestures or vocalizations of distress by the infant, particularly grimaces and the vocalizations "eee," "ikk," and "coo". In extreme cases, infant distress took the form of a tantrum, very much resembling those thrown by two-year-olds in our society. An infant baboon throws itself to the ground in a "fear paralysis", screeching ("eee") and cackling ("ikk"), while waving its tail about and watching its mother, who usually ignores the infant. The infant may periodically run after her as she for ages and the infant also sometimes interjects long "coo" vocalizations. This dramatic display sometimes lasts at high intensity for five to ten minutes, recurs sporadically throughout the day, and leaves infants thoroughly exhausted afterward. Although these extensive tantrums as well as their briefer or milder counterparts do sometimes follow aggression or punishment by the mother, they occur five to ten times as often as do such maternal acts. More commonly, a mother causes such infant distress merely by just getting up and walking away when her infant tries either to make contact or to suckle. Or she shifts position slightly, making the nipple inaccessible. Or, if the infant climbs on her back while she is walking, she suddenly sits down, thereby knocking the infant off. It is very hard to distinguish objectively these subtle acts from a mother's normal activities except from the subsequent infant behavior, a criterion that would become quite circular. However, this behavior by mothers does seem-to be quite common during about months four to six, and I think that using cine film or videotapes from that period played back at slow speed might help obtain a better understanding of the communication between the two.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 64
What seemed to stimulate punishing or aggressive acts by mothers toward their infants? During the first few months, a mother gently with drew her arm if her infant tried to hold her food or arm as she fed. Later she pushed the interfering infant away. Finally, mothers roughly shoved away older infants, who were threatened away if they even tried to feed within the mother's arm's reach. Occasionally (frequently for Grendel) infants were specifically rebuffed from nipple contact when they were four to six months of age, but most of these rejections seemed mild and situational rather than general--mothers often allowed long nursing sessions shortly after a brief rebuff. As pointed out earlier, much so-called weaning behavior actually serves to condition the infant to the "proper" time for contact and nursing. Consistent with this are limited data on the time infants spent on the nipple, which indicate that "nipple time" dropped rapidly over the first four months without rejection, decreased no more rapidly in the next few months. Also, infants and young juveniles continued to spend some daytime on the nipple until part way through their mother's next pregnancy, usually in the offspring's second year, and perhaps consider able time on the nipple at night--yearlings still slept clutched in the mother's ventrum at that age. No data are available on the amount of milk obtained by older infants, but it seems likely that at least some milk is produced as long as some sucking occurs and the mother does not become pregnant. Recall, however, that I could only occasionally identify either sucking or swallowing. Usually, I could only determine that the nipple was held in the mouth in the usual nursing position. In this study, occasional overt nipple withdrawal and punishment occurred even at resting times and sometimes after the infant had been on the nipple for a considerable time. At other times infant distress seemed to be due not to punishment but to absence of milk; the infant repeatedly switched nipples and often pulled one or both nipples by hand with no interference by the mother. This pulling may stimulate,
Maternal punishment also occurred when five- to eight-month-old infants repeatedly hopped on and off mothers, and mothers sometimes shook infants off when they were riding dorsally. At the extremes were Gin, who hardly tolerated any riding even when Grendel was only four months old, and Scar, who did not resist Summer's very frequent hopping on and off Scar's back whenever Scar took a step or, alternatively, paused for a moment. Only during months seven and eight did Scar begin to lower her hips or shake them slightly until Summer jumped off.
A situation that often stimulated loud, persistent infant distress occurred during descent from the sleeping trees in the morning. During month seven most mothers descended the sleeping trees without their infants, leaving them behind. At first infants protested violently and mothers sometimes returned to sit halfway up the tree, waiting until the infant descended that far, and then carried it the rest of the way. Occasionally an infant fell in its attempts to descend: At other times infants obtained rides on their adult male associate. Mothers often sat behind the group, facing and watching the sleeping tree until the screaming infant descended, at which time the infant usually ran to its mother, rode, and nursed. This forced independence period lasted only a month or less; eight- or nine-month-olds almost invariably descended alone in the morning without signs of distress.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 65
Summary of Weaning and Gradual Independence
In the previous sections I have tried to indicate the variables affecting infants' requirements for care in several areas in which all care is initially provided entirely by their mothers. I have also tried to indicate possible alternatives to maternal care that are available at various ages and the effects of growth and of spatial independence on both the of infant's requirements and the mother's ability to satisfy those requirements. Our knowledge of the time course of each of these is clearly imperfect. My main hope is that outlining both the information necessary to understand the development of parent-offspring relationships and our current state of ignorance will stimulate the gathering of the necessary field and experimental data.
Despite the gaps in current knowledge, we can draw several tentative conclusions at this stage. Most of an infant's requirements for care increase proportionately with the infant's increase in weight. Infant addition, both the requirements and the cost to the mother of meeting the requirements also increase as a function of the time an infant spends actively out of contact with its mother. Other conspecifics contribute, but apparently little, to providing the necessary care. If we ignore increased requirements due to semi-independence and consider only the effects of weight gain, even at slow rates of infant growth, that is, 5 grams per day, provision of just an infant's nutritional requirements and transportation would appreciably tax a mother's time budget by the time her infant was six months old. Maternal punishment and rejection during the infants' fourth to sixth month results at first in dramatic tantrums, but infants soon seem to learn to restructure their schedules so that they obtain care without interfering with their mothers' major maintenance activities.
One interesting speculation is that low-ranking mothers, by being restrictive and as a consequence spending more time in contact with their infants, provide less costly care for their infants during the months of total nutritional dependence. If so, they might, thereby, counteract the effects of being supplanted from food resources more frequently than high-ranking females are, which would otherwise necessitate their spending more time feeding. This would account for my inability to detect any relationship between a mother's dominance rank and the time she spent feeding.
Evolutionary biologists propose that these energetic, attentional, and time budget costs and protection from sources of mortality are translated into costs of and benefits to survival and future reproduction. On this basis, and assuming a measure of heritability of the traits involved, they have considered the potential for the evolution of such traits and have in addition proposed evolution through natural selection as a potential explanation for the development of observed social behaviors. Although the basic ideas were formally proposed at least as early as 1930 by Fisher, it is due primarily to expositions by Alexander (1974), Trivers (1972, 1974), West Eberhard (1975), and Wilson (1975), based on the work of Hamilton (1964), that these ideas increasingly have been applied as explanatory devices to the behavior of primates, including humans. In the next section I shall review the major features of both the more heuristic and the formal population genetic models, particularly as they apply to parent-offspring and sibling interactions, and the empirical evidence from primates that bears on these models.
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. Baboon Mothers and Infants by Jeanne Altman excerpts part 66
Evolutionary Models of Parental Investment and : Parent-Offspring Conflict
Robert Trivers, in a particularly stimulating and influential pair of papers (1972, 1974) developed a line of arguments based on pedigrees to produce predictions regarding the nature and time course of parental behaviors as a function of costs to the parents' future reproductive success and of benefits to survival of the current infant. Trivers' basic argument was that because parents are in general equally related to all their offspring but any particular (current) offspring is more closely related to itself than to its siblings, natural selection will favor offspring who attempt to obtain more parental investment (behavior that increases the survival of the current offspring while reducing a parent's ability to invest in future offspring) than their parents are selected to provide. Trivers further suggested that this genetic "conflict of interest" is resolved by behavioral conflict between parent and offspring. He pointed to the existence of behavioral conflict between parents and offspring, particularly in humans and baboons, as support for the idea that genetic conflict of interest has produced parent-offspring behavioral conflict. Clearly, there are several different questions that are relevant to this sequence of arguments.
Could a gene spread that caused an offspring to obtain more parental investment than its parent was selected to give? If so, under what conditions? Do the existing conditions for any particular species allow for the evolution of such offspring behavior in that species?
Is behavioral conflict the inevitable outcome of genetic conflict of interest? Are there reasonable behavioral options that provide more favorable cost-benefit ratios and/or require more probable genetic changes than does behavioral conflict?
Are there explanatory mechanisms for parent-offspring behavioral conflict that provide viable alternatives to the evolution through natural selection that is due to genetic conflict of interest? I shall consider each of these in turn, particularly as they apply to primates.
Several authors (e.g., Charlesworth 1978, Charnov 1977, Levitt 1975, Wade 1978) have recently constructed population genetics models of altruism, particularly sibling altruism. Each has assumed some aspects of large, randomly mating populations with large sibships. These assumptions apparently are approximately satisfied by some insect populations.
In 1978 three papers (McNair and Parker 1978, Parker and McNair 1978, Stamps et al. 1978) appeared in which population genetic models of parent-offspring conflict were presented. These followed on a note published the previous year, the first at tempt, to my knowledge, to model formally Trivers' suggestion that parent-offspring conflict will evolve and Alexander's rebuttal that this is not possible. Alexander argued that such genes will be selected against and that selection for parental manipulation of offspring will evolve. As with the related genetic models of sibling altruism, each of these recent population genetic models of parent-offspring conflict includes slightly different sets of assumptions and modes of analysis, which make it difficult to interpret results that differ. However, each model does demonstrate some conditions under which parent-offspring genetic conflict of interest could occur and that behavioral conflict might evolve through natural selection.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 66
However, a possible problem arises when we attempt to apply either the existing sibling or the parent-offspring models to primate evolution. Primate sibships, and those of a number of other large mammals, differ considerably from those assumed in most models. First, sibships in these species are very small, probably less than ten for most species. Second, the individuals constituting a sibship are not members of a single litter or even two or three litters, but rather are born singly, spaced over fairly long time spans. This results in lower levels of overlap between siblings at any life stage such as the period of infantile dependency, and less overlap throughout their lifetimes, producing an even smaller "effective sibship size" (in the same sense as "effective population size" is used in population genetics). Further, to a considerable extent maternal siblings will be half-siblings in many species. Paternal sibships also may be discrete, potentially interacting units, for example, age cohorts, under some mating and social, but they will characteristically be different from maternal sibships, offering opportunities for sibling interactions different from those offered by maternal sibships.
Sibship size and the number of offspring produced by any mating combination seem to affect the assumptions of or appear in the formal models in several places. It is not obvious to me how robust the models are under the conditions that are produced by most primate systems of mating, reproduction, and rates of maturation. I have not seen a treatment of this in the literature, although Charnov (1977, corrected in Johnson 1979) and Stamps et al. (1978) consider several special cases in which certain aspects of small sibships are accommodated in particular formulations. I hope that these points will be addressed and clarified so that appropriate applications of the models to these systems can be pursued.
An additional question arises when we consider estimation of the parameters of the models. Crucial both to the earlier formulations and to their more formal population genetic counterparts are measures of costs of an act, usually to the actor (e.g., parent), and benefits of that act, usually to the recipient (e.g., infant), both costs and benefits to be measured in units of survival and reproductive success or reproductive value. What are the costs and benefits of maternal care?
If a current infant garnered more care than its mother "was selected to provide," there are two major ways in which a primate infant might reduce its mother's future reproductive success: either through increasing the probability of her death or through delaying the conception of her subsequent offspring.
Greater mortality risk might arise from several sources. If a female increases the attention she devotes to locating food and to keeping track of her infant, I suggest that she has less attention available for predator detection and is more dependent on the alarms of the other group members. Increased infant care might also render a mother more susceptible to predation if she were less able to keep up with the group and therefore stayed at the rear, as we have commonly observed, and were therefore more susceptible to predation, particularly so if she could not flee from a predator as fast because she had to retrieve the infant (if it was out of contact) and if she ran more slowly due to the greater weight, or just due to poor physical condition.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 66
However, a possible problem arises when we attempt to apply either the existing sibling or the parent-offspring models to primate evolution. Primate sibships, and those of a number of other large mammals, differ considerably from those assumed in most models. First, sibships in these species are very small, probably less than ten for most species. Second, the individuals constituting a sibship are not members of a single litter or even two or three litters, but rather are born singly, spaced over fairly long time spans. This results in lower levels of overlap between siblings at any life stage such as the period of infantile dependency, and less overlap throughout their lifetimes, producing an even smaller "effective sibship size" (in the same sense as "effective population size" is used in population genetics). Further, to a considerable extent maternal siblings will be half-siblings in many species. Paternal sibships also may be discrete, potentially interacting units, for example, age cohorts, under some mating and social, but they will characteristically be different from maternal sibships, offering opportunities for sibling interactions different from those offered by maternal sibships.
Sibship size and the number of offspring produced by any mating combination seem to affect the assumptions of or appear in the formal models in several places. It is not obvious to me how robust the models are under the conditions that are produced by most primate systems of mating, reproduction, and rates of maturation. I have not seen a treatment of this in the literature, although Charnov (1977, corrected in Johnson 1979) and Stamps et al. (1978) consider several special cases in which certain aspects of small sibships are accommodated in particular formulations. I hope that these points will be addressed and clarified so that appropriate applications of the models to these systems can be pursued.
An additional question arises when we consider estimation of the parameters of the models. Crucial both to the earlier formulations and to their more formal population genetic counterparts are measures of costs of an act, usually to the actor (e.g., parent), and benefits of that act, usually to the recipient (e.g., infant), both costs and benefits to be measured in units of survival and reproductive success or reproductive value. What are the costs and benefits of maternal care?
If a current infant garnered more care than its mother "was selected to provide," there are two major ways in which a primate infant might reduce its mother's future reproductive success: either through increasing the probability of her death or through delaying the conception of her subsequent offspring.
Greater mortality risk might arise from several sources. If a female increases the attention she devotes to locating food and to keeping track of her infant, I suggest that she has less attention available for predator detection and is more dependent on the alarms of the other group members. Increased infant care might also render a mother more susceptible to predation if she were less able to keep up with the group and therefore stayed at the rear, as we have commonly observed, and were therefore more susceptible to predation, particularly so if she could not flee from a predator as fast because she had to retrieve the infant (if it was out of contact) and if she ran more slowly due to the greater weight, or just due to poor physical condition.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 68
The data from Social Milieu further suggest that greater mother-infant contact leads to greater contact between the mother and other group members, which in turn, particularly for low-ranking females, leads to increased stress and greater energy demands, all of which would render a female more susceptible to diseases caused by bacteria and by viruses. It also is possible that mothers are in a negative energy balance, and may accumulate particular nutritional deficiencies as well, perhaps resulting directly in starvation or, more likely, nutritional diseases.
Delay of future reproduction may be accomplished through the direct hormonal effects of sucking or through nutritional and other stress, perhaps including weight loss. This is the one area in which evidence is most available for at least the broad outlines of apparent reproductive costs. Yet the information is still inadequate for determining the effects of differences in amounts of maternal care of infants at various developmental stages on the timing of subsequent reproduction in wild primates.
The question then arises, could mothers do more? The form of maternal investment most often considered is nutritional, through lactation, and the most obvious form of parent-offspring conflict is thought to be nutritional weaning. Initially, a mother provides all its infant's energetic needs, primarily through lactation. Is she being selfish not to continue to do so? Consider a baboon mother who ordinarily spends over 55 percent of her time feeding, approximately 23 percent walking, 20 percent resting or engaged in grooming interactions when her infant is about five months old. At this stage, her infant spends 30 percent of its time in contact with her. This is the age at which pronounced dramatic tantrums occur. It is six months before most mothers . resume cycling and nine to twelve months before most become pregnant again. It is most unlikely that the mother of even a six- to nine month-old could provide enough care for survival of the infant and also support the strains of a new pregnancy. Let us assume that a mother "decides" to nurse her infant more. In order to do so she will have to spend more time feeding. Yet the sum of the mother's resting, grooming, and walking time will be decreased if the time spent in the one remaining activity, feeding, is increased. Furthermore, walking time is primarily determined by the movements of the whole social group during its day route rather than being subject to appreciable individual variability. Thus it is the mother's resting and grooming that will suffer. To the extent that these activities usually are beneficial, there will probably be a loss to both mother and infant.
Deleterious changes in allocation of time, energy, or attention, if they have life history consequences at all, are likely to have immediate ones in terms of reduced chances of survival for the mother or her current infant. In the absence of evidence to the contrary, it seems more reasonable to assume that the effects on biological fitness of an act are usually a nonincreasing function of the time since the act occurred. Costs to the mother are therefore likely to be either immediate reduction in the chance of surviving or diminished ability to provide care for the current infant, for example, reduced feeding efficiency, reduced attentiveness to predators, or inability to keep up with the group. In this context it is important to remember that the current infant's survival is entirely contingent on its mother's survival: in all cases of maternal death in baboons, the infant (i.e. under a year of age) has also disappeared. In Trivers' terminology I am proposing that what he called "self-inflicted" costs to the infant are greater and play a more important role in parent-offspring relations than may have been assumed. In terms of the population genetics models, I am suggesting that the benefit of maternal care to the infant is not a linear function of cost to the mother but rather approximates a linear function of the log of cost and reaches an asymptote or upper bound.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 69
These immediately detrimental effects of any attempt by an infant to demand more of its mother, combined with an infant's increasing abilities to care for itself, abilities whose development is crucial to the infant's eventual independence, suggest that parent-offspring genetic conflict of interest may arise infrequently as a relevant variable in many real-life situations.
Trivers further predicted that maternal care should be a direct function of maternal age. This argument was based on reproductive value being a monotonic, nonincreasing function of age for adult females and on the assumption that the relative "worth" of each of a parent's offspring is determined only by the offspring's genetic relatedness to the parent. However, additional factors should be considered in evaluating the worth to a mother of her various offspring.
1. Survival of early offspring versus later ones represents a reduction in generation time and therefore, if increased maternal care improves infant survival, greater maternal care for early infants would be favored by natural selection unless the population is a decreasing one. Note, too, that a surviving six-month-old, having survived a life stage of high mortality, has a much higher reproductive value than a future unborn offspring.
2. The potential of early offspring for "help at the nest," sibling altruism, and more reciprocal altruism with the parent due to a greater overlap of lifetimes, would also lead to a prediction of greater maternal care for early versus late offspring.
3. Mortality rates are a nondecreasing function of age in mammalian adults. Thus a later offspring is more likely to be orphaned, so it is more important for its survival for it to become self sufficient at a younger age. To the extent that care beyond a certain level retards development of self-sufficiency, such care would be disadvantageous to the infant.
4. Because of decline in metabolism and other effects of aging, the immediate costs to an older female of providing a given amount of infant care are likely to be greater than those for a younger female, off setting any decreased long-term costs as a result of lower reproductive value.
These factors can be subsumed in the cost-benefit ratios of the various models, and in that sense they offer no problem to the models as they are generally established. However, they do affect the forms that the functions have been assumed to take. These factors also affect benefit to the current offspring or cost to the parent. For example, these effects are such that they produce predictions of more care for early offspring, that is, decreasing maternal care with increasing maternal age. Effects 1 and 2 are particularly compelling. Although effects 3 and 4 are more debatable than the other two, they are still plausible. In any case, we are left with the need to state explicitly a larger set of assumptions that have often been implicit, to measure the costs and benefits and, perhaps, to develop more appropriate genetic models before we shall be able to make evolutionary predictions relating, for example, parental care and parental investment to parental age. Will conflict of interest always lead to behavioral conflict? I have suggested that more often than supposed there will be no conflict of interest between a parent and its current offspring regarding the amount of care the parent should provide. It is still important to ask, what will happen when conditions of conflict of interest arise? Is behavioral conflict the inevitable resolution of genetic conflict of interest? The answer depends on the net cost to each participant in the conflict and on the net costs of available alternative behavioral strategies.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 70
Behavioral conflict itself probably incurs a cost. Baboon infants that throw long tantrums lose time from feeding and other activities during that time, and they appear to be thoroughly exhausted when they finally stop. Moreover, it has been suggested that loud vocalizing by baboons renders these individuals more susceptible to predators. Alternatively, an infant might obtain a greater net gain (benefit) by waiting to obtain more milk from its mother when she is resting than by fighting (often unsuccessfully) to be allowed on the nipple when she is feeding. Providing additional care at rest time will cost the mother less, and so she will be more willing to provide it. Such less costly care is lower investment in Trivers' terms than care provided when the mother is feeding, but it may provide the infant with as great, or greater, benefit than care it forced its mother to provide while she was feeding or walking, both because the infant would not suffer the cost of the behavioral conflict itself and because the infant would suffer fewer self-inflicted costs resulting from its effects on its mother's survival.
Essentially, I am suggesting that an infant often can gain a particular amount of benefit in several different ways, which will vary in their cost to its mother not in just one way at a fixed single cost, and that these alternatives, along with the costs of behavioral conflict, will determine the nature of the behavioral resolution of potential conflict situations. The restructuring of contact time, described in Infant Development and Mother-Infant Spatial Relationships, appears to be a behavioral resolution through change rather than conflict. Such resolutions may often be the optimal ones.
But having chosen nonconflict strategy in some situation, wouldn't an infant still try to obtain even more care? That is, wouldn't the older infant that learned to seek out nursing when its mother was being groomed or the human toddler who learned to wait to be held until its mother finished a chore and sat down to relax, still try to obtain somewhat more than its mother "wanted" to provide? Perhaps, but not necessarily. Again, costs and benefits of alternative strategies must be evaluated. The additional investment just obtained places the current investment level at a different, higher value on the cost-benefit curve than previously, and if the curve is horizontally asymptotic, discontinuous or has other features already indicated as likely in real-life situations, then the self-inflicted costs may be greater than the potential gain that was being disputed.
In sum, conflicts of interest do not invariably lead to conflicts at the behavioral level. Cooperation and compromise are likely alternative behavioral resolutions of conflicts of interest. The relative use of different behavioral options will be a function both of the values of the alternatives and of the animals' ability to make the relevant evaluations of strategy. In a socially living animal, cooperation and compromise in parent-offspring interactions may form a useful developmental base for group organization and cohesion among adults and serve as a model for resolution of potential conflict situations among adults. In contrast, resolution by conflict may be more generally appropriate to more solitary and territorial species.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 71
Chase (1980) has pointed out that most existing models and theories dealing with the evolution of behavior have dealt with existence statements--can a behavior evolve or not--rather than with apportioning time or making choices among various behavioral alternatives. Maynard-Smith's analysis of evolutionary stable strategies, based on a game theoretic approach, is one way of considering several viable alternatives, and Chase makes use of more general economic models to propose others. Recent interest in incorporation of time as a valuable and limited resource in models of human economic decisions may prove useful in biology. The future development of models of alternative options should provide another fruitful way of looking at the evolution of social behavior in general and parent-offspring relations in particular.
Behavioral conflict certainly may arise as the resolution to genetic conflict of interest. However, caution will be needed in predicting the situations in which it is to be expected, and exclusive focus on this form of resolution would certainly be misleading. Is similar caution needed in attributing observed conflict to genetic conflict of interest?
Behavioral conflict between parents and offspring of several primates, including humans, is an observed phenomenon that is some times quite dramatic and attention-attracting. Is it always a result of genetic conflict of interest? The classic tantrums of two-year-olds in our culture and the tantrums of five-month-old baboon infants provided the prime examples in Trivers' (1974) work. The theoretical discussions have focused on conflict in which the issue between parents and offspring seems to be the amount of care the parent will provide, with the infant seeking "more," the parent "less." However, in baboons as in a number of other species, it is the infant who initially seeks independence and the parent who restricts it; the result is often struggles. Rowell first described restrictive baboon mothering in her caged colony of baboons and speculated that it might be a product of captivity, but the present study has shown it to be a more important and general phenomenon, as it apparently is in macaques. Even conflict at older ages does not always fit easily into a parental-investment model, nor I can we reject a priori alternative explanations. In any species for which there is a long period of dependency and in which we acknowledge the large role of learning during the lifetime, it seems important to consider the potential value of learning in parent-offspring relations.
This flexibility allows for developmental fine-tuning based on initial and changing states of both mother and infant, which in turn are dependent on a variety of ecological, maturational, and social factors, as shown in previous chapters. Even if a mother's and infant's "interests" overlapped entirely, owing to different environmental histories, the two are, as a dyad, a new phenotypic combination just as they are new genetic combinations as diploid individuals. A mother will need to alter her care depending on the season of her infant's birth, the prevailing conditions when her infant is at various stages of development, characteristics of the infant such as health, and the particular demographic and social milieu. If these factors were constant over a female's lifetime and over a number of generations, then perhaps a more "fixed program" would have evolved as a more efficient system. If conditions were constant over a lifetime, then perhaps some sensitive period during juvenile years or the time a female spends caring for her first infant would completely determine a female's investment pattern thereafter. Variability within a lifetime, however, will require repeated adjustments. A mother is to its infant and an infant is to its mother one of many only partially predictable environmental variables, not a predetermined constant. Additionally, their fates are inextricably bound, probably more so than those of any other pair of individuals or environment--individual pair, yet it is unlikely that they could have perfect knowledge of each other either initially or during development. Feedback mechanisms are necessary within each and between the two to monitor their own and each other's often rapidly changing states, particularly those due to maturation. Thus in addition to the possibility that behavioral conflict may sometimes be a consequence of genetic conflict of interest, it also may have evolved without genetic conflict of interest as a result of the infant's immaturity at birth, its long period of dependency, and the changing conditions under which parenting and development take place over a lifetime.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 71
Chase (1980) has pointed out that most existing models and theories dealing with the evolution of behavior have dealt with existence statements--can a behavior evolve or not--rather than with apportioning time or making choices among various behavioral alternatives. Maynard-Smith's analysis of evolutionary stable strategies, based on a game theoretic approach, is one way of considering several viable alternatives, and Chase makes use of more general economic models to propose others. Recent interest in incorporation of time as a valuable and limited resource in models of human economic decisions may prove useful in biology. The future development of models of alternative options should provide another fruitful way of looking at the evolution of social behavior in general and parent-offspring relations in particular.
Behavioral conflict certainly may arise as the resolution to genetic conflict of interest. However, caution will be needed in predicting the situations in which it is to be expected, and exclusive focus on this form of resolution would certainly be misleading. Is similar caution needed in attributing observed conflict to genetic conflict of interest?
Behavioral conflict between parents and offspring of several primates, including humans, is an observed phenomenon that is some times quite dramatic and attention-attracting. Is it always a result of genetic conflict of interest? The classic tantrums of two-year-olds in our culture and the tantrums of five-month-old baboon infants provided the prime examples in Trivers' (1974) work. The theoretical discussions have focused on conflict in which the issue between parents and offspring seems to be the amount of care the parent will provide, with the infant seeking "more," the parent "less." However, in baboons as in a number of other species, it is the infant who initially seeks independence and the parent who restricts it; the result is often struggles. Rowell first described restrictive baboon mothering in her caged colony of baboons and speculated that it might be a product of captivity, but the present study has shown it to be a more important and general phenomenon, as it apparently is in macaques. Even conflict at older ages does not always fit easily into a parental-investment model, nor I can we reject a priori alternative explanations. In any species for which there is a long period of dependency and in which we acknowledge the large role of learning during the lifetime, it seems important to consider the potential value of learning in parent-offspring relations.
This flexibility allows for developmental fine-tuning based on initial and changing states of both mother and infant, which in turn are dependent on a variety of ecological, maturational, and social factors, as shown in previous chapters. Even if a mother's and infant's "interests" overlapped entirely, owing to different environmental histories, the two are, as a dyad, a new phenotypic combination just as they are new genetic combinations as diploid individuals. A mother will need to alter her care depending on the season of her infant's birth, the prevailing conditions when her infant is at various stages of development, characteristics of the infant such as health, and the particular demographic and social milieu. If these factors were constant over a female's lifetime and over a number of generations, then perhaps a more "fixed program" would have evolved as a more efficient system. If conditions were constant over a lifetime, then perhaps some sensitive period during juvenile years or the time a female spends caring for her first infant would completely determine a female's investment pattern thereafter. Variability within a lifetime, however, will require repeated adjustments. A mother is to its infant and an infant is to its mother one of many only partially predictable environmental variables, not a predetermined constant. Additionally, their fates are inextricably bound, probably more so than those of any other pair of individuals or environment--individual pair, yet it is unlikely that they could have perfect knowledge of each other either initially or during development. Feedback mechanisms are necessary within each and between the two to monitor their own and each other's often rapidly changing states, particularly those due to maturation. Thus in addition to the possibility that behavioral conflict may sometimes be a consequence of genetic conflict of interest, it also may have evolved without genetic conflict of interest as a result of the infant's immaturity at birth, its long period of dependency, and the changing conditions under which parenting and development take place over a lifetime.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 72
Maternal Care and the Infant's First Few Months of Life
A BABOON INFANT is born into its mother's world, and immediately that infant is affected by those things that affect its mother. The infant also intensifies the very factors that influence her, as it draws the attention of other group members and increases the time and attention she must devote to maintenance activities. The adult males with whom she previously associated usually are the ones with whom the infant associates. The mother's adult female and juvenile associates also tend to be the ones who associate with the mother-infant pair, but in addition there are others of these classes who are just interested in all infants and who interact with mothers with whom they have not had previous association. The attraction of others to the infant results in low-ranking mothers being put in fearful, stressful situations much more often than otherwise because they can no longer keep a distance from their higher-ranking peers. Because infants are often pulled and even kidnaped by others, these low-ranking mothers are much more restrictive of their infants' movements, and the result is that their infants remain dependent on the mothers several months longer than do their peers.
Can we identify the characteristics of "good" mothering and "bad"? If we use a biological criterion of survival to maturity and first examine infant survival, data from this small sample of mothers suggest that restrictive mothering is perhaps better mothering. However, we must be cautious not only because of the sample size but because of two other factors as well. First, it seems likely to me that restrictive mothering may result in higher survival during the early months because it probably reduces the chances of falls, disease, kidnaping, predation. However, I have shown that infants with restrictive mothers develop independence more slowly. Thus it also seems likely that offspring of restrictive mothers will be less able to survive their mother's death at an age that offspring of rejecting mothers could; for example, the former infants might be able to survive alone only after a year and a half, the latter by one year of age. The particular balance of mortality pressures between those producing early infant mortality and those causing maternal death might vary from year to year or because of environmental differences. Restrictiveness is a strategy that achieves a short-term gain at the cost of a long-term loss.
If this basic analysis is valid, one would predict that mothers will tend to become more laissez-faire as they age because their later offspring are more likely to be orphaned, and early independence would therefore be more important to these offspring. If it is true that each kind of mothering has different advantages, or has advantages at different infant and juvenile ages, then we must also consider that infants of high-ranking females are not as subject to the socially induced dangers of laissez-faire mothering as are infants of low-ranking mothers. Thus by this argument, high-ranking mothers can in fact "afford" to be more laissez-faire; that is, laissez-faire mothering might be good mothering for a high-ranking infant, bad for a low-ranking one, and vice versa for restrictive mothering. Many of the attempts to evaluate parental care in different Western social classes ignore the fact that not only the options that parents have but also the outcome situations to which infants must adapt may differ among these groups. In contrast, an thropological studies of child rearing often consider outcomes for the particular social organization as a major determinant of child-rearing practices. If infant survival and adult adjustment, survival, or reproduction are reasonable criteria for good parenting, then the particular parental behavior that constitutes good parenting may vary considerably even within a given human or monkey society and must be considered within its normative context.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 72
Maternal Care and the Infant's First Few Months of Life
A BABOON INFANT is born into its mother's world, and immediately that infant is affected by those things that affect its mother. The infant also intensifies the very factors that influence her, as it draws the attention of other group members and increases the time and attention she must devote to maintenance activities. The adult males with whom she previously associated usually are the ones with whom the infant associates. The mother's adult female and juvenile associates also tend to be the ones who associate with the mother-infant pair, but in addition there are others of these classes who are just interested in all infants and who interact with mothers with whom they have not had previous association. The attraction of others to the infant results in low-ranking mothers being put in fearful, stressful situations much more often than otherwise because they can no longer keep a distance from their higher-ranking peers. Because infants are often pulled and even kidnaped by others, these low-ranking mothers are much more restrictive of their infants' movements, and the result is that their infants remain dependent on the mothers several months longer than do their peers.
Can we identify the characteristics of "good" mothering and "bad"? If we use a biological criterion of survival to maturity and first examine infant survival, data from this small sample of mothers suggest that restrictive mothering is perhaps better mothering. However, we must be cautious not only because of the sample size but because of two other factors as well. First, it seems likely to me that restrictive mothering may result in higher survival during the early months because it probably reduces the chances of falls, disease, kidnaping, predation. However, I have shown that infants with restrictive mothers develop independence more slowly. Thus it also seems likely that offspring of restrictive mothers will be less able to survive their mother's death at an age that offspring of rejecting mothers could; for example, the former infants might be able to survive alone only after a year and a half, the latter by one year of age. The particular balance of mortality pressures between those producing early infant mortality and those causing maternal death might vary from year to year or because of environmental differences. Restrictiveness is a strategy that achieves a short-term gain at the cost of a long-term loss.
If this basic analysis is valid, one would predict that mothers will tend to become more laissez-faire as they age because their later offspring are more likely to be orphaned, and early independence would therefore be more important to these offspring. If it is true that each kind of mothering has different advantages, or has advantages at different infant and juvenile ages, then we must also consider that infants of high-ranking females are not as subject to the socially induced dangers of laissez-faire mothering as are infants of low-ranking mothers. Thus by this argument, high-ranking mothers can in fact "afford" to be more laissez-faire; that is, laissez-faire mothering might be good mothering for a high-ranking infant, bad for a low-ranking one, and vice versa for restrictive mothering. Many of the attempts to evaluate parental care in different Western social classes ignore the fact that not only the options that parents have but also the outcome situations to which infants must adapt may differ among these groups. In contrast, an thropological studies of child rearing often consider outcomes for the particular social organization as a major determinant of child-rearing practices. If infant survival and adult adjustment, survival, or reproduction are reasonable criteria for good parenting, then the particular parental behavior that constitutes good parenting may vary considerably even within a given human or monkey society and must be considered within its normative context.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 73
We have seen the limitations placed on mothers and infants by their social and physical world, but we have also seen hints of a wide range of possible modes of action within those constraints. Why, I wonder, do not low-ranking baboon females make more use of male associates? Why do some individuals seem to be able to stay away from trouble through use of space, while others seem to get caught in the middle of every fight? In particular, it appeared that among low-ranking females, older ones such as Este and Judy, as compared with Brush and Handle, were better at making use of male associates and of space. Why do some mothers such as Preg and Slinky invest for months, to their own peril, in obviously doomed infants while others such as Gin provide so little care and that so reluctantly?
What are the causes and consequences of some adult females, regardless of age, reproductive status, or dominance rank, being much more actively interested in or aggressive in their interest in new infants? And what of juveniles such as Janet, who stayed near and interacted frequently with mothers, while others, such as Dotty, did so appreciably less? We now know that Janet was still three years from menarche, Dotty only two, facts that perhaps make the differences in their behavior even more surprising. Will Janet be more competent with her first infant than Dotty will be with hers? In laboratory studies, only extremes of social deprivation during infancy have been shown to result in poor mothering during adulthood. Will restrictive mothers continue to be so with successive infants, and will their daughters be restrictive when they become mothers? We would predict considerable consistency from the persistence of dominance rank differences, but these would be mitigated during adulthood as mothers learned to make better use of resources such as male associates and as maternal aging put a premium for the infant on early independence.
The Period of Semi-Independence
When her infant is in its fourth or fifth month, a mother's social life is beginning to return to preparturition levels. Her infant spends much time out of contact, and some of those who wish to interact with it do so during those times. By this age the infant readily recognizes individuals and often avoids some, seeks out others. The mother's social life decreases rapidly just as she needs to feed much more to provide for this growing infant: as one set of pressures, the social ones, decreases, the ecological ones increase. Stress probably peaks for high-ranking mothers when their infants are about five months old. For low-ranking mothers, for whom social stress is great during the early months, pressures probably remain high for the whole first five or six months. Experiences in these first six months probably provide the main mechanism by which offspring assume their mother's dominance rank.
By the time infants are 6 months old, all have experienced some maternal punishment and have displayed their distress with tantrums. But in response they have restructured their time with their mothers. Also, they have play groups to join, they are agile, well coordinated, and fast. If the season is right they can obtain many foods for themselves, a fortunate thing, too, because they now weigh over 15 percent of their mothers' body weight instead of the 6 to 8 percent they did at birth. If their infants are healthy and weaning foods are abundant, mothers' lives can really begin to return to "normal" when their infants are in their second six months of life. Some mothers resume their menstrual cycles during this period and by the time their offspring are 18 months old, almost all mothers are again pregnant.
What of a mother's semi-independent offspring between 6 and 18 months of age? Increasingly, it is involved in play and agonistic interactions. Less often will its mother interfere when these interactions result in distress for the infant. It is at this age that older siblings and adult male associates may become increasingly important. I think that it is in the second year of life, when mortality rates are still high, that parent offspring conflict of interest is most likely to occur and that other group members potentially provide the greatest contribution to the youngster's survival.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 74
Demographic Influences
Our ignorance of the 6- to 18-month period constitutes a major gap in our understanding of baboon life histories. It should partially be filled by the current work of Stuart Altmann at Amboseli and of Nancy Nicolson at Gilgil. However, at this time it is worth considering which individuals potentially constitute the world of the older infant and young juvenile. Who these individuals are will vary, determined to a considerable extent by the demographic factors operating in a population. Most long-term primate field studies have been conducted with expanding populations, usually ones with low rates of mortality, particularly for infants; and in the case of provisioned groups, there may also be a higher rate of conception. What is of interest here is the group structure that results from various combinations of age-specific birth and death rates and migration patterns, because each produces a particular group composition and each might be comparable to particular conditions of various human groups. High birth rates and low rates of infant mortality, for example, result in large cohorts of immatures. In such groups we might expect to find the tendencies for play groups to segregate by age and sex, as has been observed in laboratory groups of other primates, and might be comparable to the choices human youngsters have in large communities. Alternatively, in a nonexpanding population with appreciable infant mortality such as that at Amboseli, infants will not have accessible a sufficient variety of playmates to select among them. For very young baboon infants, proximity of their mothers (due perhaps to subgroup membership or dominance rank, as in this study) and closeness of infants in age may be more important variables than infant gender. Later, gender can be increasingly important as cohorts consist of wider age ranges and as the youngsters' play becomes less dependent on their mothers' location. Both group size and composition will affect an infant's options. For human children, Draper (1976) and Blurton-Jones and Konner (1973) have discussed similar issues as determinants of play group composition, choice of games, and gender differences in play. These authors contrast play among children who grew up under demographic conditions similar to those at Amboseli with play of children who live in Western cities with large peer populations. Draper also discusses other factors of social organization and mode of production as they interact with demography: and affect children's associations. Likewise, Barker and Gump make some of the same comparisons for small towns and large urban or suburban environments.
We know from studies of both human infants and caged primates that some small gender differences can be detected at very early ages. Yet it appears that under some conditions such differences may be minor or undetectable in ordinary mother and infant behavior and under others become quite important and exaggerated. The former seems to have been the case in the present study, in which other factors swamped any gender differences that might have existed. Will these early experiences just delay emergence of some sex differences or will the modified early experiences in turn modify aspects of adult behavior?
Other ways in which demographic parameters will determine the social milieu and the options available to individuals include whether individuals can surround themselves primarily with kin or not. If mortality is low, mothers can often select kin as associates.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 75
Selective Case History Descriptions of All Mother-infant Dyads with Emphasis on Adult Male and Kin Associations. The descriptions are based partially on ad libitum notes. Infants are ordered by age starting with the oldest.
1. Female Pooh
Plum was very protective of her first infant, Pooh, who was small and exhibited locomotor disability when first observed to leave her mother. There is no information on any early associations. When I first observed Pooh at the age of eight months, she was thin, smaller than the six-month-olds, retarded in her skin color maturation, and had a short, scraggy coat and impaired locomotion. She was undergoing severe weaning with frequent distress vocalizations and occasional biting by her mother, who otherwise usually ignored Pooh. At the time Pooh was associated with male Chip, who sometimes carried Pooh, stayed behind the group with her, slept near or with her in the trees and fed near her. The day after Chip migrated to another group, when Pooh was 15 months old, Pooh appeared with what seemed to be a dislocated hip and could walk only very slowly and clumsily. Male Max immediately assumed Chip's role as "godfather," staying considerable distances behind the group with Pooh, especially during the first few days. Pooh's condition eventually improved to its earlier state. Her relationship with Max continued. One day when Pooh was 24 months old, male Even ran off with Pooh in his mouth, inflicting fatal wounds on her as he did so. Max chased Even until Even dropped Pooh. Max then stayed with Pooh, grooming her and sitting near her. Plum, with her new infant, Peach, ignored these interactions. When the group moved out from the sleeping trees without Pooh the next morning, Max stayed at the rear of the group, repeatedly climbed into trees and faced back toward the sleeping grove while giving barks (at least 26 times between 0815 and 1116) that are characteristic of lost animals. The next day Pooh appeared by a shrub near Favorite Grove. Max approached and sat near her, once embracing her. Max lagged behind as the group moved off (owing to Maasai and their cattle), and as Pooh weakly attempted to follow, moving only a few meters. Cattle separated both Max and Pooh from the rest of the group. Eventually Max rejoined the group after barking and looking back toward where Pooh was last seen. She was never seen again.
2. Male Ozzie
When Oval's healthy male infant, Ozzie, was born, her daughter, Fanny, was one-and-a-half years old. High Tail and Slim were Oval's consorts the week Ozzie was. No information is available on early adult male attachments, but at six months of age Ozzie was often associated with High Tail. Seven months later, High Tail and Slim were Oval's consorts during the period in which she became pregnant with her next infant, Oreo.
3. Female Alice
Alice, a healthy female infant, was born to the oldest and highest-ranking female, Alto. Although B.J., Crest, and Peter each mated with Alto during the week that Alice was conceived, Slim was Alto's consort during the days of most likely conception and was, therefore, probably Alice's father. Alto had two other known living offspring in the group, five-year-old Spot and one-and-a-half-year-old Dotty. At 6 months of age Alice was associated with adult male Peter. An association between Alto and Peter has existed at least since 1971; during the present study an association existed between Dotty and Peter but not between Spot and Peter. In fact, Peter had grooming relations with the other family members, was often near them, even in the sleeping trees, and he supported the younger sisters and Alto in agonist encounters with Spot. Alto died when Alice was 17 months old, at which age she was still sleeping with and occasionally suckling from her mother. During the next few months, Alice slept in contact with Peter or her sister, Dotty. Alice's association with Peter persisted until his disappearance during 1978.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 76
4. Male Fred
An early association was observed between male Max and Fem with her first infant, Fred. This strong, persistent relationship existed well into Fred's second year of life, with Fred often taking advantage of it, feeding immediately next to Max. In this way Fred frequently obtained scraps of mammalian prey when others of his age did not. Max was most probably Fred's father, but B.J. was also a consort during the last week of estrus.
5. Female Eno
Eno's next-surviving sibling was five-year-old male Toto, who was not often associated with Eno or with his mother, Este, during this year. However, Eno and Toto were associated during 1976-77. When Eno was first observed by me when she was two-and-a-half months old, only a slight association could be detected between her and male B.J., on whom she occasionally rode. No stronger relationship developed before B.J. disappeared from the group nine months later. Before B.J.'s disappearance he and High Tail were Este's most common consorts when she resumed cycling.
6. Female Summer
Both Stubby and Slim stayed near mid-ranking female Scar when her female infant Summer was born, 17 months after Scar's previous infant, Cete. However, Scar often avoided both males and neither she nor Summer developed an association with any other male. Perhaps because Summer and her mother remained in general rather isolated from other adults, Summer stayed in unusually close proximity to her mother as well as spending much time in actual contact. Cete associated with her younger sister only after Summer became independent 6 to 8 months later.
7. Male Pedro
Peter, Slim, and briefly Stubby were associated with mid-ranking older female Preg when her male infant Pedro was born. When Pedro was four months old, Stubby died after suffering a severe leg wound, probably inflicted by Ben. As with Slim's other such relationships that year, he did not carry the infant and soon lost interest in the pair. Uncharacteristically, Peter did so as well. Preg's two-year-old daughter, Nazu, was not associated with the pair; her four-year old son, Dogo, was with them only occasionally. By three months of age Pedro's locomotion was decidedly abnormal and his coat and skin color development were retarded. He deteriorated rapidly in the sixth month. At that time, Nazu began to associate with him and groom him. She continued to do so until his death at eight months of age. Slim was Preg's sole consort when she resumed cycling.
8. Female Misty
Slim and Peter were frequent neighbors in the period immediately after female infant Misty was born to Mom, an elderly, high-ranking female. B.J., Chip, her and Slim had been Mom's consorts during the week Misty was conceived. No relationship developed between Misty and either male before Misty's death at two-and-a-half months of age. Partly because Slim threatened Striper away, Misty's two-year-old sister, Striper, was only slightly associated with her and her mother during Misty's brief life. Striper was more closely associated with Mom when Mom's next infant, Moshi, was born. Slim was Mom's sole consort when she resumed cycling after Misty's death.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 77
9 Male Bristle
Brush's first infant was born with webbed fingers and died in the first two weeks of life. Thereafter, Brush became pregnant in one cycle. Ben and Max both attempted to intensify their existing relationships with Brush when male infant Bristle was born six months later. Ben displaced Max and thereafter established one of the strongest and most persistent male-infant relationships observed, interrupted only for several months during which Ben suffered a severe shoulder wound, probably inflicted by male Red.
10. Male Hans
Handle was, along with Slinky, one of the two most protective and restrictive mothers when her first infant, Hans, was born. When Hans was two days old he was kidnaped by adult female Gin and kept until he was dehydrated and unable to cling; finally, he was successfully retrieved by his mother the next morning. Like other primiparous females, Handle took more than a month to accept and reciprocate or make use of a male's attempts to remain nearby, the male in her case being Even. Once established, however, the relationship between Handle and Even was one of the most reciprocal, and often Handle, Hans, Even, and Even's consort female, Lulu, constituted a close subgroup unit. Red, who also attempted association with Hans and Handle in the first month, was displaced by Even and was repeatedly avoided by Handle, perhaps because Red had supported Gin when she avoided returning Hans after the kidnaping.
11. Male Grendel
Gin's association with young male Red persisted into male infant Grendel's first month. Gin was extremely rejecting and punitive with this first infant, and a relationship between Grendel and Red began to develop. At this time, Red was undergoing a period of rapid rise in rank and was frequently involved in agonistic interactions with other males, during which he probably inflicted Ben's serious wound. These interactions culminated in Red's leaving the group, at first for two months, then sporadically thereafter. No relationship with Grendel was reestablished. Rather, during month two Grendel established a relationship with male High Tail, who had no obvious association with Gin at the time. High Tail was the only potential father recorded for Grendel but we have incomplete consort data for the cycle in question. Grendel rode on High Tail often and for long periods, fed next to High Tail and sat in his shade on hot afternoons, ran to him in times of alarm or other distress, until High Tail was killed by a leopard when Grendel was four months old. Thereafter, Grendel spent several weeks attempting to ride on many other individuals, and also tried again to spend more time in contact with his mother, but his mother remained rejecting and punitive. Most group members will readily carry only black infants; they only allowed Grendel to ride on them briefly. Grendel survived this difficult period, soon adjusting to his greater independence.
12. Female Sesame
Slinky's first pregnancy resulted in stillbirth, her second in a severely defective infant, who neither clung nor suckled, and who died two days after birth. Her third infant, Sesame, seemed weak from birth, but survived. Sesame's locomotion was poor and her maturation retarded. When she was four months old she suffered a period of illness characterized by severe locomotor problems and perhaps partial paralysis. After Sesame's birth, Max and Ben both sought association with highly protective Slinky, Max rapidly displacing the recently wounded Ben, whom Slinky often avoided. In the first month, Even stayed nearby but was noninteractive. Slinky neither approached nor avoided him and he lost interest in the pair thereafter. Max remained moderately associated with the pair, but shifted much of his attention to Pooh when Chip emigrated. Either Max or Even was probably Sesame's father. Sesame disappeared and was presumed dead during the second year of life.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 78
13. Male Juma
A good example of the dynamics of the male associations is the case of Judy and her infant, Juma. We have only partial consort data for the cycle in which Judy became pregnant, but her frequent associate in general was male Stubby, who died early in the present study, shortly after Judy became pregnant. During late pregnancy Judy was mildly associated with male Peter. Male Slim persistently followed Judy when Juma was born and even more persistently followed infant-snatching Spot, who, with her own new infant, followed Judy. Spot sometimes groomed and stayed near Slim. Judy repeatedly moved away from both of them, but was often in their presence, primarily because they persistently followed her. Peter stayed somewhat farther away at these times. Peter and Judy did approach each other. In contrast to the way she reacted to Slim, Judy appeared calm near Peter, did not restrain her infant or avoid Peter, and did groom him. Three-year-old Janet was often nearby and engaged in grooming relations with her mother and younger brother. When Juma was two-and-a-half months old, Judy and Juma disappeared during the night following the first day of what appeared to be a virus epidemic in the group. Peter was sick that day. He was one of the last two baboons to leave the sleeping grove, and as he slowly joined the group he repeatedly stopped and looked back at the grove. He was the only one to do so. We could detect no reaction to Judy and Juma's disappearance either by Slim or by Janet.
14. Female Safi
During early 1974-75, Spot had been associated with male Stubby, who died soon after Spot became pregnant. On the cycle in which she became pregnant her only consort recorded on days of likely conception was Stubby, but data are not available for all such days. When Spot gave birth to her first infant, female Safi, Spot ranked second only to her mother (Alto) in the female dominance hierarchy. Only her yearling sister, Alice, and male Slim intruded upon Spot and her infant. Slim was virtually the only animal who pulled Safi. Alice frequently pushed into Spot's ventrum, as she sometimes did with the other mothers, sitting on or pushing the infant Safi actions that Spot tolerated until Alice became too persistent and was pushed away. Alice sometimes then enlisted support from her mother, Alto, from adult male Peter, or from her three-year-old sister, Dotty. At an early age Safi developed considerable independence. During the apparently viral epidemic in May 1976, Spot and her infant, Safi, then seven weeks old, were two of the sickest baboons in the group. During the first few days of their illness, characterized by minimal, swaying movement, long rests, and no feeding, juveniles and low-ranking females who had not ventured to do so before approached and handled the infant. There seemed to be a clear recognition of Spot's incapacity, yet Spot did not drop in dominance rank during her illness, that is, there were no reversals in agonistic outcomes. Spot recovered more quickly than Safi, who retained an appreciable limp for a month and never resumed dorsal riding.
15. Male Moshi
Moshi, a healthy male infant, took full advantage of Mom's nonrestrictive care and the relative lack of interference from others. He rapidly became one of the most independent infants. Moshi's sister, three-year-old Striper, spent considerable time with her mother and new brother. Slim, Mom's only consort during the cycle of conception, was almost surely Moshi's father. Through repeated threats Slim displaced Peter as an associate of the mother and infant. The subsequent association was mild, did not last long, and included Slim's usual infant pulling. When he was about eight to ten months old, Moshi's coat turned white, a sign of poor health that usually appears about the fourth month. Moshi fell from a tree and died during his second year of life.

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Baboon Mothers and Infants by Jeanne Altman excerpts part 79
16. Female Oreo When Oreo was born, one of the two males likely to have been her father, High Tail, was dead. The other, Slim, was early associated with the mother-infant pair). Little evidence of such an association remained when I first observed Oreo, during her third month. During the last trimester of Oval's pregnancy, all ties seemed to be at least tempo rarely severed between her and her previous offspring, Ozzie, a state which had not changed four months later. A close association did exist between Oval and her three-year-old daughter, Fanny, as did one between Fanny and Ozzie.
17. Female Vicki
Vee's first infant, Vicki, was not able to get on the nipple during her first day of life; her mother carried her upside down (ventrum down) and backward even dragging her and bumping her on the ground much of the first day. The infant seemed normal at first sight in the morning, but the extreme mishandling she received soon made it hard to tell the source of poor coloration and signs of weakness and dehydration observed later that first day. In the sleeping trees the next morning, the infant was on the nipple when first seen. Vee behaved much more competently but remained fairly unresponsive and quite unprotective of the infant. She was still incompetent compared even with the other primiparous females. Vicki's early deprivation is the most likely cause of her death at three weeks. She never regained adequate skin color or clinging ability and often looked somewhat emaciated. During these few weeks, Vee was associated with male Slim, most likely Vicki's father.
18. Female Peach
As the group foraged, Plum remained behind to give birth to her second infant, daughter Peach, after about one to one-and-a-half hours of labor. At the onset, she handled and then licked the vaginal fluids. Increasingly, she thrust her arm in a sudden jerky motion or grasped and strained against a large log, grimacing. She was restless and attentive throughout. Just as the infant's head began to appear, a Maasai tribesman walked by in the distance and Plum moved under a shrub. She reappeared with infant Peach after 20 minutes and very rapidly ate the afterbirth, making several gagging motions. She finished the placenta and bit the end of the umbilicus.
For almost two hours, Max remained about 100 yards from Plum, high on a fallen, dead tree, watching her and occasionally giving single, two-phased alarm barks. Peach's sister, Pooh, remained near Max. Max, Pooh, Plum, and Peach rejoined the group, staying near each other the rest of the afternoon. The next morning, Red kept Max away from Plum. Male Even then displaced and chased Red away, remaining in a close reciprocal relationship with Plum and Peach throughout the remainder of the study (three weeks). Plum, wary of mothers and quite protective of her infant, followed Even closely. Even was most likely to have been Peach's father. Red, Max, and High Tail (dead by Peach's birth) were the other consorts during Plum's last estrous period. At three months of age, Peach became sick and died of unknown causes.
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Baboon Mothers and Infants by Jeanne Altman excerpts part 80
Selective Case History Descriptions of All Adult Males.
FOURTEEN MALES WERE ADULTS in Alto's Group during some part of the 1975-76 study year, including Stiff, Red, Russ, and Stu, who matured to adulthood in the group during that year. Changes in the overall hierarchy occurred because of deaths and migrations. Below is an alphabetical listing of the males and a brief summary of each male's history. G. Hausfater, F. Saigilo, D. Stein, and Walters have contributed demographic data from mid-1976 through 1978.
1. Ben
Ben was a juvenile male when studies began in July 1971. During the summer of 1973 he was ninth- or tenth-ranking in the adult and subadult male hierarchy, twelfth-ranking in 1974, and during 1974-75 he rose to second rank. We suspect that he inflicted male Stubby's fatal wound in October of 1975. In February of 1976, he received a similar canine puncture wound probably inflicted by Red. He dropped in rank. His recovery was extremely slow, impeded by infection, but by fall of 1976 he was beginning to take part in social interactions again, he stopped limping in early 1977, and in . December 1978 I could detect no limp or other residual signs of the wound. During 1975-76 he was closely associated with Brush and Bristle, a relationship still detectable in 1976-77 and in 1978.
2. B.J.
B.J. was an adult male in Alto's Group in July 1971. In August he was seriously wounded, probably in a fight with the dominant male, Ivan, who immediately took over B.J.'s consort female. B.J. left the group, lived alone for a period, then joined another group temporarily. He eventually rejoined Alto's group and became the dominant male of the group for it one to two years. He was mid- to low-ranking thereafter. He disappeared from Alto's group in March 1976, and we have not been able to locate him since then.
3. Chip
Chip joined Alto's Group as a fully adult male in August of 1973. He was never very high-ranking, but remained in Alto's Group and during 1975 was associated with Plum and her offspring Pooh. He disappeared from the group in March 1976, was tentatively identified in Kit South Group soon thereafter, but was not seen there during 1976-1978.
4. Dutch
Dutch was a fully adult male of Alto's Group in July 1971. Although low ranking, he was frequently in consort with adult females. During 1974 he developed a skin ailment on his hands that greatly impaired his locomotion and resulted in his walking on his knuckles. During early September 1975 he developed a severe respiratory disease, appeared to be feverish, and was unable to follow the group. The group encountered him several times in their travels over the next week. He looked thinner and more ill each time. He was last seen by us on 6 September 1975. We assume he died shortly thereafter.
5. Even
Even was a juvenile in Alto's Group in July 1971. During 1974-75 he rose rapidly in rank and he frequently migrated in and out of Alto's group, often chased for long distances from the group by Slim. He remained in Alto's Group as second-ranking male for most of the 1975-76 year, during which time he was closely associated with Handle and Hans and then with Plum and Peach. During the latter part of this year he frequently engaged in long chases of the third-ranking male, Red, as Slim had done with him. Red finally became dominant to Even during April 1977. Even was one of 11 group members (5 adult males, 3 females with infants) who disappeared early in November 1 978. Despite extensive searching and censuring of all other groups, they could not be located anywhere in the Amboseli area and were presumed dead.
Brad and Trouble
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Baboon Mothers and Infants by Jeanne Altman excerpts part 81
High Tail High Tail was the only adult male of the small group that fused with Alto's Group in fall 1972. At that time he became the dominant male of Alto's Group. He was mid-ranking during 1975 76 and was closely associated with Gin's son, Grendel, in that year. He became quite ill during the presumed viral epidemic of May 1976. Just as he was recovering, he was apparently attacked by a leopard during the early morning hours of 16 May: we found him, partially eaten, in a tree that morning with the leopard nearby and the whole baboon group in an agitated state. When they made a late descent and moved compactly and rapidly away from the sleeping grove, several juveniles looked up into the nearby tree where High Tail's body hung, behavior that led us to the corpse.
7. Max
Max was a young adult male in Alto's Group in July 1971. He was fifth ranking in July 1975 and below the newly maturing males in rank during 1975-76. He protected Pooh after Chip disappeared and rescued her from Even when the latter inflicted Pooh's fatal wounds in October 1976. Max stayed behind the group while Plum was in labor and while she gave birth to Peach. In February 1977 he left Alto's Group and joined Hook's Group. He was gone from Hook's Group by July of that year and has not been seen since.
8. Peter
Peter was a fully adult male in Alto's Group in July 1971. He was often associated with Alto at that time and with her and her offspring afterward. During 1976 he was closely associated with Judy when Juma was born. When higher-ranking males were in consort with certain females, Peter often counter chased the males, screeching, grimacing, and cackling, the outcome of which was that Peter was in consort with the female. Peter was often associated with mothers and infants, including during 1975-76, was particularly gentle with infants, and often tolerated them very close while he fed on vertebrate prey. Although quite sick during the viral attack of May 1976, he recovered and remained in the group until his disappearance in October 1978.
9. Red
Red was a juvenile in Alto's Group in July 1971. During 1975-76 he was closely associated with female Gin, supporting her when she kidnaped infant Hans, and preventing Handle from retrieving Hans. Perhaps this is why Handle resisted his attempts to associate with her while Hans was small. Red rose rapidly to third rank in the dominance hierarchy during that year and then migrated in and out of the group during 1976 and 1977 before becoming dominant to male Even in April 1977. Red disappeared in November 1978 and was presumed dead.
10. Russ
Russ was a juvenile in Alto's Group in July 1971. From physical appearance, and less so from behavior, we strongly suspected that he was Alto's offspring. Russ suddenly and rapidly rose in rank in 1976 and migrated from the group that April. When located several times shortly thereafter he was an isolate rather than being in an association with any other group. He was last seen in 1977.
11. Slim
Slim was first identified as an adult male in Stud's Group. He joined Alto's Croup sometime between September 1973 and May 1974, at which time he was the dominant male of Alto's Group, and he remained so until mid-1978. During 1975-76 he was the only adult male who frequently pulled infants and in general was quite rough with them. During that year he seemed to lose his interest in each infant after it was a couple of months old, and older infants did not seem to seek his protection or feed near him. Slim disappeared and was presumed dead in November 1978.
12. Stiff
Stiff was a juvenile in July 1971. At least since that time he had a limping walk. He was relatively noninteractive, and he had not risen in dominance rank before he left the group in February 1976. For about a month we saw him occasionally, always as an isolate and in the same part of the home range each time.
13.Stu
Stu was a juvenile in Alto's Group in July 1971. In December 1975 and January 1976 he began a rapid rise in dominance. He migrated to Stud's Group in 1976, moved to Hook's Group for several months during 1977, and then returned to Stud's Group, where he remained as first- or second-ranking male in December 1978.
14. Stubby
Stubby was a high-ranking male in Alto's Group in July 1971 and remained so until he was fatally wounded, probably by Ben, in October 1975. The puncture wound became badly infected and by mid-November he hardly moved. He disappeared the night of 12 November and was presumed dead.
Brad and Trouble
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Baboon Mothers and Infants by Jeanne Altman excerpts part 82
Behaviors Recorded in This Study Analyzed in the Text
AN (s) PRECEDING A BEHAVIOR indicates that it was included in the set of submissive or distress behaviors analyzed in this study: a preceding (a) is used for behaviors of aggression. Approach: directed walk toward, with orientation to individual rather than to spot; distinguishable by direction of looking and by end-point of approach and by any reorientation if object moves.
Approach-avoid: limbs (especially forelimbs) of animal are flexed, shoulders lowered, chin thrust out, animal looks at object (often infant in mother's ventrum) and, rocks forward and back--forward often while looking at infant, back while tensing and looking (or giving brief repeated "anxious" glances) at mother.
(s)Avert body: directed turn of body away from another, but not with the tensing and twisting of a "cower" or the rigidity of a "cower" or of "fear paralysis."
(s)Avert head: turn of head away from; often also with "avert stare"; see also "avert body."
(s)Avert stare, avoid looking at: usually a fixed looking elsewhere than at the individual who is directly in front of the actor's eyes.
(s)Avoid: directed walk away from; identified by its being preceded or accompanied by brief rapid glancing or a very mild cower.
(a)Bite, nip: obvious.
Brush past: walk past, touching, and usually with body swept against, the object.
(a)Chase: rapid run toward with repeated change of direction, tracking the object animal, and not accompanied by gestures of fear or submission.
Clamber, climb on: obvious; usually done by infants.
(s)Coo, cry: pursed-lip vocalization usually done by infants; hard to locate; usually done in a distress (e.g., weaning) rather than a fright context (I have heard an adult give this vocalization only once--female Handle, --when her two-day-old infant was kidnaped and she was unsuccessful in recovering it).
(s)Cower: "lateral flexion of the spine, often from a seated position"; head often lowered into shoulders; incipient "avoid" --often followed by"avoid'; or incipient "fear paralysis," often followed by that behavior.
Eyelid displays: variety of eyelid and brow movements among which I usually did not distinguish.
(s)Eee: screech vocalization; occurs in a fear context.
Embrace (infant): arm about other animal.
(s)Fear paralysis: an extreme behavior in which the animal remains rigid, close to the ground; it often follows and grades from "avert stare," "avert head," "avert body," "cower."
Flex arms: arms of animal flexed lowering front half of body and usually with rear in air while looking at another, usually smaller individual; often accompanied by lipsmacking and cocking of the head; seemed to be a pacificatory and inviting gesture to infants; approach-avoid is this behavior alternated with a tense drawing away and anxious glancing.
Follow: repeated change of tempo and/or direction of walk with respect to and temporally "in response to" one animal's movements more than to the group or subgroup as a whole.
Brad and Trouble
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Baboon Mothers and Infants by Jeanne Altman excerpts part 83
(a)Grab, hold down: obvious.
Groom: picking through the fur, usually of another animal; one of the most visible, durative, and common primate social behaviors.
Handle infant: moving or manipulating some part of infant rather than just touching it; but see also "pull (infant)" for that distinction.
(a)Hit, Swat: obvious.
Hold nipple, attempt suck: reaching toward the nipple with hand or with mouth but without contact or barely touching; distinguished from "rooting" by hesitancy of movement and by the ability to locate the nipple.
Ignore: obvious; "it was sometimes obvious that a monkey saw, heard, or felt a behavior pattern that was directed conspicuously toward it, yet 'ignored' the other monkey or monkeys involved".
(s)lkk: also called gecker or cackle, a vocalization often alternating with "coo"; given in immediate response to, say, a maternal rebuff, then followed by a coo; emitted in a context of low-level fear as well as moderate distress.
Lipsmacking: rhythmic lip and tongue movements, repeated rapidly.
(s)"look anxious"): repeated rapid glancing without eye to eye contact, usually with some head jerking, tensing.
Look at: obvious; but not "stare at" or "look anxious."
Lose grasp, lose hold, fall: obvious.
(a)Lunge at: incipient chase.
Mounting (sexual): Non cycling females do not receive mounts with intromission; only brief pelvis-grabbing, usually in response to a sexual presentation, occurred more than rarely in this study.
Muzzle-muzzle: placement of one animal's muzzle in contact with another's.
Muzzle anogenital area: obvious.
Muzzle other body part: obvious.
Pause: break in motion, a turn, a look back at another animal; not reliably distinguished from "wait" except a pause is shorter, not held until another joins the actor.
Present for grooming: animal presents its trunk, broadside; if seated, trunk presented often with arm up and head lifted and tilted.
Present (sexually) (tail up): animal presents its anogenital area to muzzle , of another animal, usually with one leg back, tail often raised.
Pull (infant): when the infant is in contact, and is clinging or is being embraced, this pulling is distinguished from "handling" by the fact that the infant's body is moved toward the actor and away from its present location with sufficient force that present contact would be broken if infant and/or "surface" (e.g., mother) did not exert counteracting force.
(a)Push, shove, lean on: usually a gross body movement or trunk movement rather than a grab, hold down, or hit; but occasionally done with a hand and then distinguishable from a hit by being slower with more contact time and without a grasp.
Restrain infant: actively (hold, embrace, etc.) prevent infant from leaving contact while infant is making movements that would take it out of contact if not restrained; thus this is a dyadically defined behavior.
Ride ventrally: cling ventrally to the trunk of a standing or walking individual.
Rooting: mouth and head movement, primarily lateral but with some lifting, hand searching "for" the nipple; only done by very young infants.
(a) Run away from: rapid avoid; see "avoid."
Run toward: rapid approach; see "approach."
Sit in (or climb in) ventrum of: obvious; usually done by infants.
Soft grunts: low, rhythmic grunts usually given while animal is approaching or seated looking at an infant.
(a)Stare at: prolonged, intense, fixed look at.
Suckle: obvious; in practice I could usually determine only if an infant had the nipple in its mouth and not whether it was actually sucking.
(a)Tail up: tail is held high, often arched, but not in a posterior presentation
Touch: obvious; but see "handle infant" and "pull (infant)" for distinction.
Wait: pause in walk until the object individual reached the actor, at which time the actor immediately resumed its walk; see also "pause"--not reliably distinguished from pause except by duration and joining by the actor.
Brad and Trouble
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KRISTY

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The mommy monkeys are very protective of their young. People under estamate the mommy when they have a baby they shouldn't though because the mommy will hurt and probably kill them and it wont be to pretty if it did happen to you if you were the one who undereastamated her
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monkeys are cute!
with love,
stephanie
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