Bentley-Condit 2003

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International Journal of Primatology [ijop] pp985-ijop-472826 October 15, 2003 21:36 Style le version Nov. 18th, 2002
International Journal of Primatology, Vol. 24, No. 5, October 2003 (
C
2003)
Sex Differences in Captive Olive Baboon Behavior
During the First Fourteen Days of Life
Vicki K. Bentley-Condit
1
Received June 3, 2002; revised January 21, 2003; accepted February 18, 2003
I discuss newborn baboon behavioral and proximity sex differences in a pop-
ulation of captive olive baboons ( Papio anubis) living in a social group of
>500 individuals. The data are based upon 20-min focal observations of
42 mother-newborn pairs (n = 27 , n = 15 ) for infant-days 17 and 36
pairs (n = 23 , n = 13 ) for infant-days 814 collected late-May through
late-November 2001. I examined the rst two weeks of infant life via behav-
ioral, proximity, and approach-leave/contact analyses in order to determine
whether behavioral sex differences exist during the rst few days of life. I ex-
amine and analyze these 2 weeks independently due to different sample sizes. I
used data fromthe total available sample population of 57 infants (n = 36 ,
n = 21 ) to discuss birth, survivorship, and infant weight. Statistically signi-
cant age and/or age-sex interactions exist for all of the behavior and proximity
measures during either infant-week 1 or 2. Moreover, there is a statistically
signicant difference in the birth sex ratio in the sample population but no
signicant difference in infant mortality by sex. There are also relative and
signicant differences in mothers treatment of their newborn males and fe-
males. There are alsosome general tendencies for female newborns onaverage
to suckle less and to explore more per focal observation than male infants do
as they age. Conversely, male newborns average slightly more time per focal
observation 1 m from the mother than do female infants. However, the ob-
served differences may be inuenced by maternal behavior in that mothers
have higher rates of contact with their female than their male infants.
KEY WORDS: infants; development; sex differences; Papio.
1
Department of Anthropology, Grinnell College, Grinnell, IA 50112; E-mail: bentleyc@
grinnell.edu.
1093
0164-0291/03/1000-1093/0
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2003 Plenum Publishing Corporation
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INTRODUCTION
Researchers have studied cercopithicine infant development and in-
dependence for ca. 4 decades. Acquiring independence from the mother
and the process by which an infant comes to explore her environment are
important aspects of this developmental process. These issues have been
particularly well explored for a variety of macaque and baboon species.
Beginning with DeVores (1963) work and continuing through the 1980s,
the exploration of infant development in Papio spp. has had many contribu-
tors, including, Rowell et al. (1968), Nash (1978), Altmann (1978, 1980), and
Nicolson (1982). While each of them contributed to our understanding of
infant development in Papio and placed it within the complex social milieu
and ecological setting of life in a baboon troop, most focused upon neither
the very rst days of infant life nor possible infant sex differences.
For other cercopithecine species, macaques in particular, some early re-
searchers did incorporate the early days of infant life, and there were reports
of some sex differences in infant behavior, particularly in terms of the type
of play and levels of aggressions (Harlow, 1962; Hinde et al., 1964). Further,
cercopithecine mothers can treat their male and female infants differently
(Eaton et al., 1985; Hinde, 1982). Among macaques, female infants have
received and reciprocated more positive physical contact (Mitchell, 1968),
male infants have received more punishment, rejections, and breaks in con-
tact (Eaton et al., 1985; Mitchell and Brandt, 1970), and mothers have played
more approach-leave games with their male infants (Simpson et al., 1986).
However, many of these sex differences did not become apparent before
infants were 6090 days of age, and more recent studies showed no early
behavioral sex differences (Schino et al., 1995; Tanaka, 1989).
While macaque infant development has continued to be addressed
through the last decade (Maestripieri, 1994), there has been little research
specically addressing baboon infant development since the 1980s. Instead,
infant baboon research has focused on other aspects of the mother-infant
pair, ranging from infant handling (Bentley-Condit et al., 2001; Chism, 2000;
Henzi and Barrett, 2002; Nicolson, 1987) to maternal care as contingent or
strategic behavior (Altmann and Samuels, 1992; Barrett et al., 1995; Bentley-
Condit, 1995; Lycett et al., 1998; Smuts, 1985). Wasser and Wassers (1995)
work is an exception to this trend.
What becomes apparent from a review of the literature on Papio is that
no study has had possible infant sex differences in the earliest days of infant
life as the central focus. Many researchers did not address sex differences at
all and several did not include the rst few weeks of infant life in their anal-
yses. When baboon infant sex differences have been addressed, the focus
has generally been upon older infants with no signicant differences found
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Sex Differences in Captive Olive Baboon Behavior 1095
(Young and Bramblett, 1977; Young and Hankins, 1979) or no differences
until 6090 days of age (Ransom and Rowell, 1972). Thus there is a gap in
the baboon infant development literature, both in terms of time span and
behavior, that I addressed via an examination of a captive population of
olive baboons (Papio anubis) in a large social group setting at the Southwest
Foundation for Biomedical Research (SFBR). Specically, I sought to deter-
mine whether there are male andfemale infant behavioral differences during
the rst 14 days of life and the roles played by both infants and mothers in
proximity maintenance during this period.
In addressing infant behavioral sex differences, it is also important to
consider factors that could inuence both infant and maternal behaviors.
With another SFBR population, Coelho et al. (1984) showed that male in-
fants tend to be signicantly heavier than female infants at birth. As a dif-
ference in size could affect or cause differences in behavior, I examined in-
fant weight data. Additionally, Wasser and Norton (1993) and Maestripieri
(2001c) suggestedthat cercopithecine mothers candiscriminate their infants
sex and recognize it very early in its life (Maestripieri, 2001b; Maestripieri
et al., 2000) and may adjust birth sex ratios towards, and invest more in, the
sex most likely to survive. As a skew in investment could affect either infant
or maternal behaviors, I also included infant birth, survival, and sex ratio in
the following discussion.
Based upon data available from studies of macaque early infant devel-
opment and of later baboon infant development, the working hypothesis
must necessarily be that there is no signicant behavioral sex difference.
However, contrary to published reports, observations ad libitum (Altmann,
1974) of newborns in the SFBR population during 1997 and 1998 led me
to believe that there were some subtle behavioral differences between very
young female and male infants. By infant-day 3, male infants appeared to be
much more engaged and active, e.g., reaching for objects and others, to the
extent that infants could be successfully sexed based upon these behaviors.
These observations ledtothe following predictions for the SFBRpopulation:
1. Female and male infants differ in the ways in which they apportion
their time among the behavioral categories recorded.
2. Female and male infants differ in the ways in which they apportion
their time among the proximity categories recorded.
My project is unique in that (a) it has very early possible sex differences
as the central focus, (b) it addresses the issue in a large, captive population
unique in its size and make-up that has never before been the focus of a
behavioral studywheretheresearcher has workedwithinthegroups habitat,
and (c) it presents the opportunity for a relatively large sample size over a
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relatively short period of time. Thus the project has the potential to further
our understanding of the extent to which female and male newborn baboons
behave differently and how mother-infant interactions may vary by infant
sex during the rst 2 weeks of infant life.
METHODS
Study Site, Subjects, Data Collection, and Statistics
I collected data at the Southwest Foundation for Biomedical Research,
(Southwest Regional Primate Center) in San Antonio, TX. The subjects are
an olive baboon breeding population, established in 1979 (Goodwin and
Coelho, 1982), housed in a large corral (6-acres; radius = 88 .4 m, cir-
cumference ca. 600 m). They receive a basic nutritional regime of LabDiet
!
supplemented by occasional fruits and vegetables and have access ad libitum
to water. All corral residents receive physical examinations and are checked
for TB biannually. Ill or injured baboons are removed from the corral and
taken to the hospital for treatment and recuperation. Otherwise, they are
left to themselves. I have collected data from inside the corral every sum-
mer since 1997 and for 6 mo in 2001. The baboons are habituated to my
presence.
I collected most of the data upon which this study is based during the
6-mo period of late-May through late-November 2001, during which all in-
fants born, i.e., live births, over a 17-week period (28MAY0124SEP01)
served as subjects (N = 48; n = 31 , n = 17 ). I included 2 additional
infants born in early October 2001 (n = 1 , n = 1 ) to bring the total to 50
mother-newborn pairs. Mother-newborn pairs remained in the study until
the infant reached 60 days of age or died, whichever occurred rst. I also
included 7 mother-newborn pairs for which I collected data in previous sum-
mers. Thus, the database consists of 57 mother-newborn pairs (n = 36 ,
n = 21 ) and 1710 20-min focal observations (570 observation hours). I col-
lected data once per day between ca. 0645 and 1500 h, 6 days per week
(Monday-Saturday). I scheduled observations of each mother-infant pair so
that all 2-hblocks of time, (i.e., 0700-0900, 0900-1100, etc.,) were represented.
During the infants rst 14 days I amassed 6 20-min focals (2 observation h)
per week for each mother-infant pair. Because it was not always possible
to collect data on a mother-infant pair for all 6 days of any given week be-
cause of inclement weather, mother-infant hospitalization, or inability to
locate the subject, there are different sample sizes for infant-weeks 1 and
2. Therefore, I describe and analyze the 2 weeks independently. During fo-
cal observations for each pair, I recorded a continuous record (Altmann,
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1974) of mother and infant behaviors, their proximity to one another and
which one was responsible for transitions in proximity, (i.e., approaches and
leaves, making and breaking contact), and their interactions with other troop
members.
To examine whether there are very early behavioral sex differences
between male and female infants, I recorded 3 primary behaviors, (i.e., what
the infant was doing), and 4 primary proximity measures, (i.e., where the
behavior occurred), during each focal observation of the mother-infant pair
over the infants rst 14 days of life. I describe the 7 proximity categories
and infant behaviors in Table I; they closely follow the denitions of Wasser
and Wasser (1995).
Table I. The seven infant behaviors and proximity measures
Behaviors
SUC SUCKLE. The infant has oral contact with the
mothers nipple. He/she may or may not be
involved in active nursing
RST REST. The infant is on the mother and inactive,
but not on the nipple. She/he may or may not
be sleeping
EXP EXPLORE. The infant is on or off the mother,
active and engaged in oral or manual
manipulation of its environment
Proximities
NIP NIPPLE. The infant is on the mothers nipple.
This is essentially the same as behavioral
category SUC
CNT CONTACT. The infant is in contact with, i.e.,
touching, the mother but not on the nipple.
Contact may be ventral or dorsal and active or
inactive
PX1 PROXIMITY 1 METER. The infant is 1 m
from the mother but not in contact
PX3 PROXIMITY 3 METERS. The infant is >1 m
but 3 m from the mother
Denitions of proximity indices
APR APPROACH. One individual moves toward the
other and changes from one proximity category
to another, e.g., from Px3 to Px1, from Px1 to
CNT. May or may not include making contact
LEV LEAVE. One individual moves away from the
other and changes from one proximity category
to another, e.g., from CNT to PX1, from PX1 to
PX3. May or may not include breaking contact
MKC MAKES CONTACT. One individual moves from
an off position, i.e., PX1 or PX3, to an on
position, i.e., CNT or NIP, relative to the other
BRC BREAKS CONTACT. One individual moves
from an on position, i.e., NIP or CNT, to an off
position, i.e., PX1 or PX3, relative to the other
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1098 Bentley-Condit
Some of the behavioral categories subsume more micro-level behav-
iors. For example, explore incorporates both oral and manual interaction
with the environment; the proximity measures incorporate approaches and
leaves, making and breaking contact, and by which subject. As there is obvi-
ous overlap between one of the behaviors and one of the proximitiesthe
suckle behavior must necessarily occur at the nipple proximityI analyze
anddiscuss onlysuckle. I calculatedapproaches andleaves andmakes/breaks
in contact from the proximity data for the proximity indices (Table I). The
use of only key behavioral and proximity categories is not an unusual prac-
tice when focusing upon such young infants (Wasser and Wasser, 1995),
and these categories accounted for 99.5% of infant behavior during the rst
2 weeks of life. In discussions and analyses of all behavioral and proximity
data, I utilize durations whereas in the discussions of the proximity indices
I employ frequencies.
I used several proximity indices to evaluate responsibility for the main-
tenance of contact and proximity. Hindes (1983) Approach/Leave Index
[(Infant Approaches/All Approaches x 100) (Infant Leaves/All Leaves x
100)] reects the differences between approaches and leaves for which an in-
dividual, or a class of individuals, is responsible. Negative scores indicate that
the focal individuals, i.e., the infants, left more often than they approached
while positive scores reect the opposite pattern. Approaches and leaves can
only be recorded when the infant or mother breaks contact and/or moves
from one of the previously dened proximity categories to another. Chang-
ing fromNIP to CNTis not a proximity transition because the mother-infant
pair does not actually break physical contact. I utilized 3 other proximity in-
dices in addition to Hindes (1983) index. The Contact Index (Simpson et al.,
1986) is very similar to the approach/leave index but only considers making
and breaking physical contact rather than all approaches and leaves [100
Makes by Infant (IM)/Makes by Mother (MM) + Makes by Infant (IM)
Breaks by Infant (IB)/Breaks by Mother (MB) +Breaks by Infant (IB)]. The
second is an additional proximity index that Brown (2001) suggested better
reects which partner is responsible for the majority of proximity changes
than the approach/leave index [(IM + IB/IM + MM + IB + MB) 100].
Thirdly, I included rates of mother contact [Mother Contacts/(1000s off )]
and infant contact [Infant Contacts/(1000s off )] (Simpson et al., 1986) be-
cause high scores indicate an intolerance of being out of physical contact
for an extended period of time. All of the measures combined help to de-
termine primary responsibility for proximity maintenance and whether this
responsibility varies for female and male infants.
Given the data, i.e., multiple observations on each mother-infant pair,
I used repeated-measures ANOVAs to compare the infant behaviors and
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proximities (Table I). ANOVA analyses allow the examination of the main
effects of infant sex, infant age, and their interaction for the various behav-
iors and proximity measures and are therefore appropriate to answer the
questions upon which the study is focused. To address any possible assump-
tion violations of a repeated-measures F, I used the Greenhouse-Geisser
correction throughout (Keppel, 1982). Because of the very small data values
represented in day-by-day comparisons, the presence of only one between-
subjects factor, and the potential bias of multiple t -tests requiring an in-
nitesimally small corrected (Bakeman and Gottman, 1986; Keppel, 1982;
Lehner, 1996), I did not use post hoc tests. Accordingly, the analyses are re-
stricted to the main effects of age and the age/sex interactions for each infant
week. I used Students t -tests for pairwise comparisons (where ANOVAa
are not used), z-tests in the comparisons of percentages, and chi-square tests
for the comparisons of birth and mortality sex ratios, mortality by week by
sex, and maternal style by maternal rank by infant sex. All tests are 2-tailed
and is 0.05. For all ANOVA statistical analyses, I utilized SPSS c 10.1.0
for Windows.
RESULTS
Infant Birth, Weight, and Survival
All infants born over a 17-week period are included as subjects.
Although infant births were not witnessed, times of birth are correct
within 1012 h. The nature of the study did not allow for the systema-
tic collection of weight data on newborns. However, weight data were col-
lected on all available infants over a 3-day period at the end
of October 2001, during a biannual round-up, for comparative
purposes.
The weight data include 58 infants (n = 26 , n = 32 ) of varying ages
(range = 1154 days). Forty-one of the 50 study subjects are included; 8 in-
fants had died and one was hospitalized at the time. The other 17 infants,
born late-September through October, were not part of the study sample but
provide useful data to show the distribution of infant weights by sex and age
(Figure 1). The mean masses for female and male infants are signicantly
different (x SE = 1.07 kg 0.058; x SE =1.51 kg 0.08; t = 4.21,
df = 49, p < .0001). However, because the male infant sample has an aver-
age age of 81 days and the female infants have an average age of 49 days,
the difference in average mass is not surprising. Although these data are
not amenable to any further statistical analyses, the plotted masses and the
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Fig. 1. INFANT MASS BY AGE A plot of each infants mass (N=58 infants; n = 26 females,
n = 32 males) at 1-154 days and the regressions for males and females.
regression lines show male infants in the sample to consistently weigh more
than the female infants.
Infant birth, survival, and sex ratio data for the study population are in
Table II. While there were signicantly more male infants born as part of
sample, there is no signicant difference in the mortality rates for male and
female infants and no signicant difference in mortality rates for this group
versus wild populations.
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Table II. Infant demographic data and comparative mortality data
All Sex ratio
2a
p
Study sample
Births 57 21 36 1.71:1 3.948 (df = 1) <0.05
Deaths
b
9 3 6 2:1 0.057 (df = 1) >0.5
Mortality 0.1579 0.1429 0.l667
Births Deaths
b
Mortality z
c
Comparative populations
Amboseli (Altmann, 1980) 18 2 0.1111 0.46 >0.6
Gilgil (Nicolson, 1982) 35 1 0.0286 1.847 >0.1
Tana River (Bentley-Condit, 1995) 11 0 0 1.42 >0.4
a
Males compared to females.
b
Infant deaths through infant-day 60.
c
Study to comparison population.
BEHAVIOR
Ages 17 Days (Table III)
I included 42 newborns (n = 15 , n = 27 ) in the analyses of EXP,
SUC, and RST (Table I). For exploration-time the infants show a signicant
main effect of age (F = 9.356, df = 3.437, p < .0005) and a signicant inter-
action of infant sex and age (F =3.027, df =3.437, p = .026) over the 7 days.
While one might expect a signicant main effect of age, the fact that the
changes occur during the rst 7 infant-days is noteworthy. Over the course
of infant-week 1, female infants devoted slightly more time to exploration as
reected in the average time per focal observation of exploring for females
( x SE=105.99 17.16 sec, x SE=93.14 10.18 sec). There is also a
signicant main effect of age on suckle-time (F=3.196, df =4.961, p = .009)
and resting-time (F = 3.045, df = 4.618, p = .014). A signicant interaction
of sex and suckle-time occurs across infant-week 1(F = 2.359, df = 4.961,
p = .042) while there is no signicant interaction of sex and resting-time
(F = .831, df = 4.618, p = .521).
I included 36 newborns (n = 13 , n = 23 ) in analyses of infant-
week 2. As with infant-week 1, there is a signicant main effect of age on
suckle-time (F = 2.89, df = 5.161, p = .015) and a signicant interaction of
sex and age (F = 2.453, df = 5.161, p = .034). Unlike the infant-days 17
data, there is no signicant main effect of age (F =.698, df =4.788, p = .620)
or a signicant interaction of age and sex on resting-time (F = 1.955, df =
4.788, p = .091). Males and females appear to have very similar resting-time
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1102 Bentley-Condit
patterns during infant-week 2. Like days 17, there is a signicant main ef-
fect of age on exploration-time (F = 3.647, df = 4.688, p = .005); however,
there is no signicant interaction of age and sex (F = 1.164, df = 4.688,
p = .329). In general, female infants average more time per focal obser-
vation exploring during week 2 than male infants do ( x SE = 284.21
27.86 sec, x SE =219.70 17.77 sec) and less time suckling ( x SE =
721.68 33 sec, x SE=791.37 23.5 sec). This equates to female infants
averaging an extra >5% of their focal time devoted to exploration and male
infants averaging an extra >5% devoted to suckling during infant-week 2.
PROXIMITY
Subjects in the analyses of proximity (Table I) are the same 42 new-
borns (n = 15 , n = 27 ) used in the behavioral analyses for infant-days
17. There is a signicant main effect of age on time spent at the proximity
category CNT (F =2.599, df =4.994, p = .027) and a signicant interaction
of age and sex (F =2.471, df =4.994, p = .034). However, there is no signi-
cant main effect of age on PX1 (F =.738, df =1.342, p = .432) nor is there a
signicant interaction of age and sex (F =.962, df =1.342, p = .357). While
neither sex spent an appreciable time at PX1 during infant-week 1, mother-
male infant pairs average slightly more time per focal observation at this
distance than mother-female infant pairs do ( x SE = 6.89 2.69 sec,
x SE = 2.21 0.78 sec). PX3 cannot be analyzed for infant-days 17
because infants were not >1 m from their mothers during then.
I used the same sample for this age group as in the behavioral analyses
(N = 36 newborns; n = 13 , n = 23 ). For CNT, there is a signicant main
effect of age (F = 2.833, df = 5.167, p = .016) and a signicant interaction
of age and sex (F = 2.799, df = 5.167, p = .017). The two other proxim-
ity categoriesPX1 and PX3also reveal differences. There is a signicant
main effect of age on PX1 (F =4.321, df =3.169, p = .006) and a signicant
interactionof age andsex (F=2.529, df =3.169, p = .050). Male infants aver-
ageslightlymoretimeper focal observationat PX1duringinfant-week2than
female infants do ( x SE = 29.18 4.77 sec, x SE = 17.4 4.68 sec).
At PX3, there is a signicant main effect of age (F = 3.482, df = 1.717,
p = .044) but no signicant interaction of age and sex (F =1.592, df =1.717,
p = .214). Infants spent essentially no time at PX3 from their mothers per
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Table III. Summary of data analyses
Behavior and Proximity
Ages 17 days Ages 814 days
(n = 15 , n = 27 ) (n = 13 , n = 23 )
F df p F df p
SUC
Age main effect 3.196 4.961 0.009 2.89 5.161 0.015
Age/sex interact 2.359 4.961 0.042 2.453 5.161 0.034
RST
Age main effect 3.045 4.618 0.014 0.698 4.788 0.620
Age/sex interact 0.831 4.618 0.521 1.955 4.788 0.091
EXP
Age main effect 9.356 3.437 0.0001 3.647 4.688 0.005
Age/sex interact 3.027 3.437 0.026 1.164 4.688 0.329
CNT
Age main effect 2.599 4.994 0.027 2.833 5.167 0.016
Age/sex interact 2.471 4.994 0.034 2.799 5.167 0.017
PX1
Age main effect 0.738 1.342 0.432 4.321 3.169 0.006
Age/sex interact 0.962 1.342 0.357 2.529 3.169 0.050
PX3
Age main effect N/A 3.482 1.717 0.044
Age/sex interact N/A 1.592 1.717 0.214
focal observation ( x SE = 1.49 0.58 sec, x SE = 1.76 1.24 sec)
during infant-week 2.
CONTACT AND PROXIMITY INDICES
In examining newborn behavioral and proximity sex differences, it is
also necessary to determine if any perceived differences can be attributed
to infant behavior or to maternal behavior. One way to approach this ques-
tion is to examine responsibility for proximity transitions and maintenance
and making and breaking contact. I calculated approach/leave, contact, and
proximity indices for all newborns, male infants only, and female infants only
for infant-weeks 1 and 2 (Table IV). The indices indicate that mother-female
infant relationships, as expressed by proximity, may differ frommother-male
infant relationships at a very young age.
DISCUSSION
The primary goal of the study was to determine if there were infant
baboon behavioral sex differences during the rst 14 days of life that could
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1104 Bentley-Condit
Table IV. Contact and proximity indices and rates
Ages 17 (n = 15 , n = 27 ) Ages 814 (n = 13 , n = 23 )
All ii All ii
Approach/leave
index (all
approaches &
leaves)
40 33.4 50 22.37 13.95 51.55
Contact index (all
makes/breaks in
contact)
40 33.4 50 24 15 55.9
Alternative
proximity index
(% changes in
contact due to
infant)
45.9% 48.89%
a
43.75% 64.6% 67.59%
c
54.8%
Mother contact rate
(mother contacts
per 1000s off)
14.33 11.52 30.22
b
11.38 10.13 14.92
Infant contact rate
(infant contacts
per 1000s off)
5.21 5.38 4.32 12.57 15.19 5.16
d
a
Males to females, z = .355, p > 0.1.
b
2
= 18.17, df = 1, p < 0.001.
c
Males to females, z = 1.984, p < 0.05.
d
2
= 5.34, df = 1, p < 0.05.
be characterized by a relatively simple set of measures. This appears to be
the case (Table III). Female and male infants in the population apportioned
their time differently between the behavioral and proximity measures, and
there are some patterns that emerge from the data and results, including 1)
behavioral and proximity patterns relevant to infant age, 2) behavioral and
proximity patterns relevant to infant sex, 3) patterns in birth ratio and infant
weight relevant to infant sex, and 4) patterns in the way mothers treat their
infants relevant to infant sex. Why these patterns occur is complicated as it
is likely that they are all inter-related.
First, there are behavioral and proximity patterns relevant to infant age.
For both females and males, there are positive relationships between infant
age and exploration-time and infant age and time at proximities contact
and 1-meter (Figure 2). Although the infants devote little time, proportion-
ately, to these measures, there are discernable increases from infant-week
1 to infant-week 2. There is a concomitant negative relationship between
infant age and suckle-time. Resting-time remains essentially constant over
the 2-week period. There are thus trade-offs occurring in time-budget al-
lotments to allow for the changes (Figure 3). The behavioral and proximity
changes with infant age reect a general pattern of increasing mobility and
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Fig. 2. FEMALE&MALEINFANTS INFANTAGE-BEHAVIOR/PROXIMITYRE-
LATIONSHIPS Male and female infant means and standard errors for each week and
the positive and negative relationships of behavior/proximity with infant age for suckle,
explore, and rest and proximities contact and 1-m. Proximity 3-m (PX3) is not plotted as
the time allotment is too small to show on the scale of the Y-axis.
capabilities with increasing age similar to patterns reported by Hinde et al.
(1964), Rowell et al. (1968), Ransomand Rowell (1972), Maestripieri (1994),
and Wasser and Wasser (1995). What is of interest here is not so much that
changes occur with age, but that discernable changes in behavior are occur-
ring at such a young age.
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1106 Bentley-Condit
Fig. 3. FEMALE & MALE INFANT BEHAVIORAL TIME BUDGETS WEEK 1 VS
WEEK 2 This stacked bar-chart presents a comparison of female and male infant time budgets
for Week 1 and Week 2 for the behaviors of suckle, rest, and explore.
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Second, there are behavioral and proximity patterns that are relevant to
infant sex. The analyses reveal signicant interactions of infant age and sex
for all of the behavioral and proximity measures except RSTand PX3 for ei-
ther infant-week1 or 2 (Table III). Female infants average more exploration-
time for bothinfant-weeks 1 and2 thanmales doandshowa steeper decrease
insuckle-time; these patterns are reectedinthe infant time budgets for each
week (Figure 3). With regard to the proximity measures, mother-male infant
pairs show a slight tendency to spend more time at PX1 than mother-female
infant pairs do. The data contradict other research showing higher activity
levels in both human and nonhuman primate male infants (Campbell and
Eaton, 1999; Eaton et al., 1985) and support those that have shown female in-
fants tobe more active (Michel et al., 1990). The data indicate that behavioral
sex differences exist at a very young age in this population.
Third, there are patterns in infant birth and weight, but not in survival,
by sex. All of the infant deaths occurred during the rst 14 days of infant
life, indicating that this period is a particularly risky time for the infants
in the population. Although more male infants than female infants died in
absolute terms, the survival and mortality rates for the 2 sexes are the same;
male and female infants do not differ signicantly in mortality for either
week 1 (
2
= .235, df = 1, p > .05) or week 2 (
2
= .127, df = 1, p > .05).
However, there is a difference in the live-birth sex ratio, with signicantly
fewer female infants born. This differs fromKoban et al.s (2001) report of no
sex bias in births for another population of olive baboons at the SFBR, but is
similar to other reports of male bias for female-philopatric species (Johnson,
1988). Further, there is an apparent tendency for male infants to outweigh
female infants over the rst 5 mo of life. The infant mass regressions for the
population appear similar to the mean birth weights reported by Glassman
et al. (1984) ( x = 0.94 kg, n = 45; x = 0.84 kg, n = 40) and are consistent
with ndings of statistically signicant differences in male and female birth
weights (Coelho et al., 1984; Glassman et al., 1984) at the SFBR. Although
the means reported by previous researchers equated to male infants being
ca. 10% larger than female infants, infant weight does not appear to have
had an effect on the reported sex differences as there is no consistent pattern
in both mobility and activity. Although male infants may be slightly more
mobile, (as reected by PX1,), female infants are more active, as reected
by EXP (Figs. 2 and 3).
Finally, there are patterns in the ways mothers act toward their new-
borns by infant sex. The proximity indices showed no signicant difference
betweenmale andfemale infants during infant-week1 inthe overall percent-
age of changes in contact due to the infants. Nor are the rates at which male
and female infants contact their mothers signicantly different. However,
mothers contact their female infants at a signicantly higher rate, compared
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1108 Bentley-Condit
to the (expected) rate for all infants, than they do their male infants dur-
ing this period. The higher rate reects that the mother-female infant dyad
is apart for a shorter period of time (ca. 33.1 sec) than the mother-male
infant dyad (ca. 86.8 sec) before the mother makes an effort to regain con-
tact. The more strongly negative scores for female infants on the Contact
Index and Approach/Leave Index, combined with the contact rates and Al-
ternative Proximity Index data, indicate that mothers are working harder
to maintain contact with their female infants. In fact, the earliest observed
mother-initiated rst break in contact with female infants did not occur until
infant-day 6, while with male infants, the earliest observed mother-initiated
break occurred on infant-day 3.
During infant-week 2, male infants are responsible for a signicantly
greater percentage of the changes in contact with their mothers than female
infants are. Concomitantly, male infants make contact with their mothers
at a higher rate than female infants do ( = once per 65.8 sec; = once
per 193.8 sec) and female infants have a signicantly lower contact rate,
compared to the (expected) rate for all infants, than males do. Mothers
continue to make contact with their female infants at a slightly higher rate,
and both the Contact Index and the Approach/Leave Index indicate that
mothers continue to take a stronger responsibility to maintain contact with
their female infants during week 2. For the mother-male pairs, the infant
must assume more responsibility for maintaining proximity, while for the
mother-female pairs it is the mother that does so. Thus, mothers appear
to behave differently toward their female infants demonstrating a higher
level of attentiveness, as expressed by approaches and making contact than
toward male infants. These results differ from other captive olive baboon
studies that suggested a lack of maternal behavioral bias during the rst few
weeks of life (Koban et al., 2001).
A sex-biased difference in mothers tendency to approach/contact their
infants may indicate higher investment in female infants in the SFBR pop-
ulation. However, the general cercopithecine pattern tends to be for a bi-
ased maternal investment to mirror a biased sex ratio (Maestripieri, 2001c;
Wasser and Norton, 1993) and, in the SFBR population, sex ratio and in-
vestment as reected by contact/approaches appear to be in opposite direc-
tions. Additionally, the data may reect variation in maternal style by infant
sex. Mothers approaching/contacting their male infants relatively less than
their female infants may indicate leniency towards male infants, even during
infant-week 1. If so, this differs from other research on baboon maternal
style (Altmann, 1980) that showed variation by maternal rank but not by in-
fant sex. In the SFBR population, style as reected by contacts/approaches
showed no statistical relationship between maternal rank, i.e., high, mid, or
low, and infant sex (
2
= 2.54, df = 2, p > .28).
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Mothers higher contact rates with their female infants might be related
toinfant vulnerability, particularlygiventheriskiness of therst 14days of in-
fant life. Primate mothers are more protective of their infants when they per-
ceive themto be at risk (Fairbanks, 1996), and protectiveness is positively as-
sociated with higher frequencies of handling and harassment (Maestripieri,
2001a). At least three of the 9 infant deaths in the sample occurred due to
fatal handling by other females. Given the presence of fatal infant-handling
and extremely shallow matrilines, (i.e., 1.66 adult matrilineal kin per adult
female Bentley-Condit, 2001) in the SFBR population, the close proximity
of other females could be potentially risky to their newborns. If mothers con-
tact/approach their female infants more often because they see themas more
vulnerable, a higher presence of other females 1 m of mother-female pairs
than of mother-male infant pairs could be an indicator of this vulnerability.
However, this does not appear to be the case for the SFBR population. For
the rst two weeks of life, there is no statistical difference between mother-
male pairs and mother-female pairs (t = 0.563, df = 48, p = .576) in terms
of time other females are 1 m. Thus, the higher mother contact rates and
more strongly negative proximity indices for mother-female infant pairs can-
not be explained by female infants receiving greater interest, as expressed
by proximity, from other adult females than of male infants.
In conclusion, we are left with the following outcomes in terms of the
original predictions.
1) Female and male infants are expected to differ in the ways in which
they apportion time among the behavioral categories.
This prediction is supported by the data. There are statistical differences
during infant-week 1 that reect age and infant sex interactions for SUCand
EXP and during infant-week 2 for SUC.
2) Female and male infants are expected to differ in the ways in which
they apportion their time among the proximity categories.
This prediction is also supported by the data. There are statistically
signicant differences during infant-weeks 1 and 2 that reect age and sex
interactions for CNT, and for infant-week 2 for PX1.
Other patterns emerge from the data that were not part of the original
predictions. First, there is a skew in the infant sex ratio at birth for the SFBR
sample population and a lack of correlation between infant sex and mothers
rank. Second, there are signicant effects of infant age on both behavioral
and proximity measures even within the rst 7 days of infant life. Finally,
the proximity indices and contact rates indicate mothers treat their female
infants differently from their male infants at a very young age. It is this
last point, in particular, that complicates the interpretation of the results on
infant behavioral sex differences as maternal inuence on infant behavior
and proximity cannot be excluded.
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1110 Bentley-Condit
Despite the above caveat, the male and female newborns in the SFBR
population behaved differently and did so almost immediately after birth.
While it remains to be determined the extent to which the patterns are
population specic or generalizable, my study reveals that there is still much
to be learned about baboon infant development, sex differences, and the
effect of the mother upon infant behavior and proximity.
ACKNOWLEDGMENTS
The research was funded by grants from the Committee on Faculty
Support at Grinnell College, Grinnell, IA. I am grateful to the Southwest
Foundation for Biomedical Research for their permission to conduct behav-
ioral research inside their breeding corral and for their continued support
of my research. I thank Linda Brent, Thomas Butler, Bill Cummins, Scott
Chambers, and other members of the Department of Laboratory Animal
Medicine animal care staff for all their patience and assistance. I also thank
Thomas Moore, Grinnell College mathematics department, for his contin-
ued assistance in all things statistical and two anonymous reviewers for their
constructive and benecial comments.
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