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Consistency in maternal behavior within families of free-ranging rhesus monkeys An extension of the concept of maternal style.

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American Journal of Primatology 22:159-169 (1990)
Consistency in Maternal Behavior Within Families of
Free-Ranging Rhesus Monkeys: An Extension of the
Concept of Maternal Style
CAROL M. BERMAN
Department of Anthropology, State University of New York at Buffalo, Buffalo,New York;
Caribbean Primate Research Center, Box 906, Punta Santiago, Puerto Rico
This paper assesses the extent to which free-ranging rhesus monkeys
(Macaca mulatta) on Cay0 Santiago show consistent mother-infant interaction with individual infants over time and consistent maternal styles
from one infant to another. Inter-pair differences in several measures of
mother-infant interaction related to proximity and contact were correlated positively and highly significantly between 6-week age periods.
Analysis of variance methods were used to show that, after the first 6
weeks, these measures of interaction were also more similar among pairs
with the same mother (i.e., families) than among pairs with different
mothers. Differences between families were also consistent between age
periods, and appeared to be due primarily to differences between mothers
rather than to differences between sets of infant siblings. The results suggest that the concept of maternal style can be used to describe variation
between both individual Cay0 Santiago mother-infant pairs and between
individual Cay0 Santiago mothers.
Key words: Macaca mulatta, maternal behavior, maternal style, Cay0
Santiago
INTRODUCTION
Inter-individual differences in mother-infant interaction among nonhuman
primates can often be described in terms of particular qualities of relationships
(e.g., restrictive, protective) [Hinde & Simpson, 19751. The description of such
qualities is generally based on the identification of differences in one or more
measures of mother-infant interaction between individual mother-infant pairs
which are consistent over time, and on consistent relationships between measures.
The measures are typically those concerned with the regulation of contact and
proximity between mothers and infants. Such consistency at the level of individual
mother-infant pairs appears to be common among cercopithecines, e.g., rhesus
[Hinde & Spencer-Booth, 1967; Berman, 1978, 1982; Simpson & Simpson, 19861,
yellow baboons [Altmann, 19801, vervets [Fairbanks & McGuire, 1987,19881, but
is not universal, e.g., bonnet macaques [Silk, in press].
Received for publication December 22, 1989; revision accepted May 25, 1990.
Address reprint requests to Dr. Carol M. Berman, Department of Anthropology, State University of New
York, Buffalo, NY 14261.
0
1990 Wiley-Liss, Inc.
160 I Berman
The term maternal style has also been used to describe such consistency [Altmann, 19801. This refinement is justified when the primary responsibility for differences in interaction can be traced to differences between mothers rather than
infants, as it has been for a range of species [e.g., Hinde & Spencer-Booth, 1967;
Rosenblum & Youngstein, 1974; Altmann, 1980; Simpson & Simpson, 1986; Fairbanks & McGuire, 1987; see, however, Chalmers 119721. Because discussions of
maternal style are usually based on analyses of individual mother-infant relationships within a population over a limited time period, it is not clear whether
particular maternal styles are descriptive only of individual mother-infant relationships or of more enduring qualities of particular mothers. Some researchers
have casually noted tendencies for mothers to display similar maternal styles with
successive infants [Altmann, 1980; Goodall, 1986; see, however, Dolhinow, 1980,
19821, but so far there is little quantitative data available to assess the extent to
which this sort of within-family consistency occurs. Berman [ 19901 recently used
analysis of variance methods to show that free-ranging rhesus monkeys (Macaca
mulatta) who have reared several infants both differ consistently from one another
in the rates at which they reject infants, and display similar rejection rates with
each of their infants. Fairbanks [19891 had similar results for percentage of time
captive vervet mothers and infants spent in mother-infant contact with one another. However, the extent to which this is true for a variety of measures of
mother-infant interaction remains unclear. This paper examines the extent to
which the concept of maternal style can be used to describe variation in motherinfant relationships both among individual pairs and among mothers who have
reared a number of infants. This is done by (1) reassessing the extent to which
individual rhesus mother-infant pairs on Cay0 Santiago show consistent patterns
of contact and proximity interaction across infant age periods, using a larger sample than previously [Berman, 1978, 19821; (2) extending analyses of within-family
consistency to several measures of mother-infant interaction; and (3) assessing the
extent to which differences between families are consistent across age periods; and
(4) are due primarily to differences between mothers or between sets of infant
siblings. Detailed descriptions of particular maternal styles displayed will be presented elsewhere [Berman et al., in preparation].
METHODS
The study took place on Cay0 Santiago, Puerto Rico, between January 1974
and December 1986. Cay0 Santiago is a 15.2 ha island off the east coast of Puerto
Rico that has supported a closed, free-ranging, provisioned, predator-free population of rhesus monkeys since 1938. The population has been censused daily since
1956, and hence is well habituated and made up of known individuals with known
histories and maternal kinship relationships. See Rawlins & Kessler [19831 and
Sade et al. [19851 for detailed descriptions of the history, terrain, population, and
management.
The study group, group I, was formed naturally by fissioning in 1961 and was
organized in a manner typical of multimale, multifemale macaque groups [e.g.,
Lee, 19831. It consisted of about 64 individuals in 1974 and grew to a peak of about
320 in 1984, before fissioning into five smaller groups in 1985 and 1986. There
were three matrilineages in group I, with members spanning as many as four
generations. The sample included a total of 38 mothers and 124 infants born into
all three lineages (9 in 1974,12 in 1975,15 in 1979,21 in 1983,22 in 1984, 21 in
1985, and 24 in 1986). The 88 infants born between 1983 and 1986 were the
offspring of females observed previously as mothers and/or infants. Each mother
had from 1to 7 infants in the sample (XtSD=3.3&1.3).Male infants represented
Consistency in Maternal Behavior / 161
TABLE I. Measures of Mathe-Infant Interaction
Time off: the percentage of time mothers and infants were neither in ventroventral contact or
i n nipple contact was estimated from point time samples.
Time on uentrum: the percentage of time mother-infant pairs spent in ventroventral contact
was estimated from point time samples.
Time on the nipple: the percentage of time infants spent on the mother’s nipple was estimated
from point time samples.
Time >5 m: the percentage of time off that was spent >5 m from the mother was estimated
from point time samples.
Relative rejections: the proportion of all attempted nipple contacts, including those initiated by
the mother, that were prevented by the mother.
Maternally initiated contact: the proportion of all nipple contacts which were initiated by the
mother.
Proximity index: a measure of the infant’s relative role in maintaining proximity to the mother;
it is the percentage of approaches made by the infanC to the mother over a 60-cm limit minus
the percentage of departures made by the infant from the mother over a 60-cm limit (Hinde &
Atkinson 1970).
621124 or 50.0% of the sample. First-born infants represented about 25% of the
sample.
Observations began as soon as possible after birth and after the annual trapping period during January and February. In most cases they continued until the
infant was at least 30 weeks of age. Exceptions included the infants born in 1979
(who were observed only during weeks 1-12), 4 infants that died before 30 weeks,
and 5 infants that were born exceptionally late in the season. Focal-animal sampling methods [Altmann, 19741were used to record all social interactions of infants
with their mothers, along with the identities of the initiator and the recipient of
the interactions and the times they occurred. At two-minute intervals, point (instantaneous) time samples were taken identifying individuals in contact with the
focal animal, within 60 cm and between 60 cm and 5 m. All observations were
made between 700 hr and 1,200 hr local time, hours which included all major group
activities: feeding a t the food bins, foraging, traveling, socializing, and resting.
However, time spent near the feeder bins was not included in the observations. The
sequence of infants observed was determined by a randomized order arranged at
the beginning of each week.
Each mother-infant pair was observed for 3 hr during each 2-week age period,
generally in 3 half-hour sessions per week. This amount of time is comparable to
that typically used in other provisioned colonies where observation conditions are
excellent and where interaction rates tend to be high. These data were suffkient to
show stable individual differences across age periods [Berman, 1978, 19821 (see
aIso Table I11 below).
Table I lists seven measures of mother-infant interaction chosen for analysis,
along with their definitions. Four measures are basic measures of percentages of
time mothers and infants spent in contact and in proximity to one another. Three
additional measures deal with the dynamics of contact and proximity maintenance. These measures are particularly usefkl in determining the relative responsibility of mothers and infants for differences between relationships [Hinde, 19691.
In addition, they provide a basis for direct comparisons between studies done in
captivity [e.g., Hinde and Spencer-Booth, 19671and in the wild [e.g., Berman et al.,
in preparation]. Measures concerned with the dynamics of nipple contact are emphasized because differential patterns of nipple contact are thought to have par-
162 I Berman
TABLE 11. Numbers of Infants in the Sample for Each Mother During Each 6-Week
Age Period
Age Period (weeks)
Mother’s name
526
604
WE
530
673
FH
348
501
3R
233
446
262
454
497
234
44 1
317
498
676
470
443
AA
cs
528
585
A43
584
674
A47
587
A41
678
633
A27
A26
A3 1
A25
C62
Total mothers: 38
Total infants: 124
1-6
2
2
1
3
1
2
3
2
1
1
2
3
1
2
1
3
1
2
1
2
2
1
3
1
1
1
26
44
7-12
1
1
2
3
1
-
4
3
2
4
4
2
4
1
6
3
4
2
1
3
1
2
2
2
2
4
2
1
3
2
1
3
2
1
2
2
1
36
84
13-18
2
1
3
3
2
3
3
2
5
3
2
4
1
6
2
2
1
2
3
1
2
1
3
3
3
3
2
3
3
2
3
2
1
2
3
1
36
88
19-24
25-30
3
3
1
1
4
3
2
1
3
3
2
6
3
3
5
1
5
2
2
3
4
2
1
3
4
2
6
3
3
4
5
2
2
1
3
3
1
2
2
3
3
3
3
2
4
3
2
5
3
1
3
3
1
37
100
1
3
3
1
2
3
3
3
3
3
3
4
3
3
5
2
2
3
3
35
101
Total hours
per mother
81
36
126
135
63
27
144
126
90
216
135
99
162
45
225
9
99
90
45
90
135
45
90
81
99
99
135
117
72
126
99
81
171
90
45
90
108
36
-
titularly important consequences for infant development and for subsequent maternal reproductive performance [e.g., Altmann, 1980; Nicolson, 1982, 1987; Lee,
1987; Stewart, 1988; Gomendio, 1989; Johnson et al., in preparation].
The data were analyzed within five 6-week infant age periods: (1-6 weeks,
7-12 weeks, 13-18 weeks, 19-24 weeks, and 25-30 weeks). Only mother-infant
pairs whose data were complete for each 2-week period within a 6-week age period
were included in the sample for that 6-week age period. Hence all mother-infant
pairs included within an analysis for a given 6-week age period were observed for
Consistency in Maternal Behavior I 163
TABLE 111. Consistency of Inter-pair Differences in Measures of Mother-Infant
Interaction Over Time: Pearson Correlation Coefficients
~
~
~
~~~~~
Between
nonadjacent
age periods
Between adjacent age periods
Time off
N(m-i pairs)b
Time on ventrum
Nb
Time on nipple
Nb
Time >5 m
Nb
Relative rejections
Nb
Maternally initiated
contact
Nb
Proximity index
Nb
1-6 wk
vs .
7-12wk
7-12 wk
vs.
13-18 wk
13-18 wk
vs.
19-24 wk
19-24 wk
vs.
25-30 wk
-0.06
39
0.59***
39
-0.10
39
0.33**
72
0.61***
72
0.32**
71
0.60***
72
0.76***
61
0.53***
0.41***
88
0.66***
88
0.41***
88
0.61***
88
0.69***
86
0.39***
0.36***
95
0.68***
95
0.35***
95
0.63***
95
0.57***
95
0.29***
60
0.54***
72
81
0.61***
88
90
0.72***
95
-
0.60***
28
0.65***
28
-
X
w
Proportion
positive and
significant"
0.26
0.08
216
0.64
0.46
616
0.19
0.04
116
0.61 0.48
313
0.66
0.50
616
0.47
0.29
516
0.62
0.38
313
aProportion of coefficients that are positive and significant a t least at a P<0.05 level
bNumber of mother-infant pairs.
*P<0.05.
**P<O.Ol.
***P<O.OOl.
the same amount of time (9 hr) during that age period. Table I1 lists the number
of infants included in the sample for each mother during each 6-week age period,
and the total amount of time each mother was observed. Since undisturbed observations could not begin until after the annual trapping period [Berman, 19891, a
number of infants born before the end of trapping could not be observed in their
early weeks. For this reason sample sizes increased with each 6-week age period.
Arcsine transformation were used with all proportional measures, such that (T
= arcsin
where p was the proportional measure [Sokal & Rohlf, 19691. With
such transformations (and in most cases without) the data met the assumptions for
parametric correlation analysis and analysis of variance. All statistical tests were
two-tailed.
6
RESULTS
Consistency over Age Periods for Individual Dyads
Pearson correlation tests were used to assess whether differences in interaction between individual mother-infant pairs were consistent across 6-week age
periods. With only two exceptions, correlations for each measure of mother-infant
interaction across adjacent age periods were positive and highly significant (Table
111). For each measure except Time on the Nipple and Time Off, most correlations
across nonadjacent age periods were also positive and significant, although they
tended to be less strong.
Consistency over Age Periods for Families
Pearson correlation tests were also used to assess whether differences in interaction between families were consistent across 6-week age periods. Mean mea-
164 I Berman
TABLE IV. Consistency of Inter-familyDifferences in Measures of Mother-Infant
Interaction over Time: Pearson Correlation Coefficients
Between
nonadjacent
age periods
Between adjacent age periods
1-6 wk 7-12 wk 13-18 wk 19-24 wk
vs.
vs.
vs.
vs.
7-12 wk 13-18 wk 19-24 wk 25-30 wk
-0.09
Time off
N (mothers)
24
Time on ventrum
0.47**
N (mothers)
24
-0.07
Time on nipple
N (mothers)
24
Time >5 m
N (mothers)
Relative rejections
0.61***
N (mothers)
22
Maternally initiated contact 0.58**
N (mothers)
22
Proximity index
N (mothers)
0.11
36
0.56***
36
0.09
36
0.45**
36
0.71***
34
0.57***
34
0.44**
36
0.35*
36
0.61***
36
0.34*
36
0.79***
36
0.72***
35
0.48**
35
0.68***
36
0.45**
34
0.79***
34
0.42**
34
0.79***
34
0.70***
33
0.27(*)
33
0.77***
34
-X
Proportion
positive and
significant"
0.20 0.11
116
0.61 0.35
316
0.19 0.06
2/6
0.68 0.42
313
0.68 0.51
616
0.48 0.38
4/6
0.63 0.44
3/3
"Proportion of coeficients that are positive and significant at least a t a P<0.05 level.
'"P<O. 1.
*P<0.05.
**P<O.Ol.
***P<O.OOl.
sures of mother-infant interaction were calculated for each family (i.e., for each
mother with all her infants in the sample). Not surprisingly, the results were
similar to those for individual mother-infant pairs (Table IV). With five exceptions, correlations across adjacent age periods were positive and significant. With
the exceptions of Time on the Nipple, Time Off, and Time on the Ventrum, most
correlations across nonadjacent age periods were also positive and significant,
although less strong.
Consistency of Mothers with Each of Their Infants
Analysis of variance (ANOVA) methods were used to assess the extent to
which mothers with two or more infants in the sample displayed similar measures
of mother-infant interaction with each infant. A separate test was carried out for
each measure and for each 6-week age period. In each test, a measure of interaction
was entered as the dependent variable and mother's identity was entered as a
factor. Because some measures varied with yearly changes in the size of the social
group [Berman, 19881, group size was controlled by entering it as a covariate
before the effect of the factor was assessed. Hence, each analysis was able to
determine whether the mother's identity could account for a significant proportion
of the variation after any variation attributable to group size was removed.
With few exceptions after the first age period, mother's identity emerged as a
significant or near significant source of variation between mother-infant pairs
(Table V). In other words, after the first six weeks, scores for mother-infant pairs
with the same mothers (i.e., families) tended to be more similar than those for pairs
with different mothers. There was no tendency for infant siblings to be of the same
sex (F ratio for infant sex ratio by mother's identity = 0.77, df = 27, NS). Although
Consistency in Maternal Behavior / 165
TABLE V. Consistency of Measures of Mother-Infant Interaction within Families: F
Ratios for Analysis of Variance of Mother-Infant Measures by Mother's Identity,
Controlling for Group Size
Age period (wk)
1-6
Time off
Time on ventrum
Time on nipple
Time >5 m
Relative rejections
Maternally initiated contact
Proximity index
N (mothers)
N (mother-infant pairs)
1.68
0.51
1.48
0.76
0.68
14
31
7-12
1.80*"
1.83*"
1.80*"
2.12*a
1.86*'
3.38***b
4.02***a
26
74"
66b
13-18
19-24
25-30
1.64'"
1.44
1.58"'
1.52"'
2.05*
1.53'*'
2.37**
29
82
1.63*
2.20**
1.83*
0.82
2.82***
1.82*
2.63***
30
94
2.20**
1.31
2.28**
1.17
2.54***
2.49**
1.90*
31
98
'"P<O. 1.
*P<0.05.
*+P<O.Ol.
***P<O.OOl.
infants within families tended to have similar numbers of older immature siblings
(F ratio for number of immature siblings by mother's identity = 2.24, df=27,
P<O.OOl), this factor could not account entirely for within-family consistency.
When ANOVA tests for within-family consistency in measures of mother-infant
interaction were repeated controlling for numbers of older immature siblings,
mother's identity generally continued to emerge as a significant or near significant
source of variation (Table VI).
Responsibility for differences.
Hinde's [1969] methods were used to assess whether differences in measures of
mother-infant interaction between families were due primarily to differences between mothers or between sets of infant siblings. Applying the method involved
hypothesizing two simple kinds of differences between mothers or sets of infants:
(1)mothers may have differed in their propensities to seek contact and proximity
with infants, or (2) sets of infants may have differed in their propensities to seek
contact and proximity with their mothers. Predictions were subsequently made
about the directions with which the various measures of contact and proximity
would differ for each hypothesized type of difference in prcpensities. For example,
if differences between mothers were primarily responsible for differences between
families, one would predict that relatively high levels of contact between mothers
and their infants would be associated with relatively low levels of maternal rejections and with relatively high proportions of their contact initiated by the mother
(i.e., negative correlations between time in contact and relative rejections, and
positive correlations between time in contact and proportions of maternally initiated contact). Similarly one would predict that relatively low levels of time at a
distance would be associated with infants taking relatively small roles in maintaining proximity to their mothers (i.e. positive correlations between Time >5 m
and the Proximity Index). On the other hand, if differences between infants were
primarily responsible for differences between families, one would predict correlations in the opposite directions. The hypothesis that simple kinds of differences
were responsible for differences between families would be supported to the extent
166 / Berman
TABLE VI. Consistency of Measures of Mother-Infant Interaction within Families: F
Ratios for Analysis of Variance of Mother-Infant Measures by Mother's Identity,
Controlling for Group Size and Numbers of Older Immature Siblings
Ape Deriod (wk)
Time off
Time on ventrum
Time on nipple
Time >5 m
Relative rejections
Maternally initiated contact
Proximity index
N (mothers)
N (mother-infant pairs)
1-6
7-12
1.59
7.25*
1.43
0.48
1.32
14
31
1.98'*'"
1.74"'"
1.94'*'"
1.82"'"
3.58**b
2.93**b
5.06***"
26
74"
66b
13-18
19-24
25-30
1.89*
1.81(*'
1.80'"'
1.92*
2.17*
1.37
2.19*
29
82
2.56**
2.34**
2.91**
1.02
2.53**
1.21
2.10*
30
94
3.17***
1.69'*'
2.83**
0.87
2.82**
2.70**
1.68'"
31
98
'"P < 0.1.
*P 10.05.
**P <0.01.
***P <0.001.
that marked correlations could be found between measures in the predicted directions. To apply the method, mean measures of mother-infant interaction were
calculated for each family. Relationships between measures were assessed using
Pearson correlation tests. The results (Table VII) were generally in harmony with
the hypothesis that differences between mothers were primarily responsible for
differences between families, at least after the first 6 weeks.
DISCUSSION
Both families and individual rhesus mother-infant pairs on Cay0 Santiago
show significant degrees of consistency over successive infant age periods in several aspects of interaction related to proximity and contact. In addition, mothers
who raise several offspring show a significant degree of consistency with each of
their infants, at least after the first 6 weeks. This within-family consistency is not
dependent on the sex of the infant or on the presence of older siblings, factors
frequently cited as important sources of variation between mother-infant pairs
[e.g., Hooley & Simpson, 1983; Berman, 1984; Simpson & Simpson, 1986; Silk, in
press].
These findings, along with indications that in rhesus monkeys both inter-pair
differences [e.g., Hinde & Spencer-Booth, 19671 and inter-family differences in
measures appear to be due more immediately to differences between mothers than
to differences between infants, are consistent with the concept of maternal style,
and suggest that an extension of the concept to individual mothers may also be
justified, at least for Cay0 Santiago rhesus. The extent to which such consistency
applies to other environments or other species remains to be seen. Simpson &
Howe [19861 found significant differences for several measures of mother-infant
interaction between different groups of captive rhesus mothers, but not between
mothers within each group. Fairbanks [1989] found consistent differences in percentages of contact between 11 captive vervet mothers of 21 infants, and hypothesized that maternal consistency may be due to temperamental consistency, particularly with reference to confidence and timidity. Fairbanks suggested that this
consistency is rooted in both genetics and the mother's experience as an infant.
Consistency in Maternal Behavior / 167
TABLE VII. Responsibility for Differences in Mother-Infant Interaction Measures
Between Families: Pearson Correlation Coefficients Between Mean Scores for
Individual Mothers
Aee ueriods (wk)
Relative rejections with
Time on ventrum
Time on nipple
Time off
Maternally initiated contact with
Time on ventrum
Time on nipple
Time off
Proximity index with
Time >5 m
N (mothers)
1-6
7-12
13-18
19-24
25-30
-0.24
-0.10
0.15
-0.40**a
-0.32*"
0.33*"
-0.31*
-0.26'*'
0.23
-0.46**
-0.48***
0.47***
-0.41**
-0.28*
0.33*
0.66***
-0.20
0.08
0.15"
0.02"
-0.02"
0.47**
0.27'"
-0.30*
0.65***
0.46**
-0.42**
0.38*
0.09
-0.03
26
0.43**b
34"
0.43**
36
0.35*
36
0.18
34
36b
'"P <0.1.
*P <0.05.
**P 10.01.
***P <0.001
However, later experience and relatively stable social circumstances during motherhood (e.g., rank) should also be considered [see also Berman, 19901.
The findings that mothers are primarily responsible for differences between
families do not necessarily imply that within-family consistency is due entirely to
enduring maternal attributes. Consistent maternal behavior could also result from
enduring aspects of the family's social environment. Nor do the results imply that
differences between infants do not exist or do not influence mother interaction.
Indeed, differences between infants could theoretically contribute to within-family
consistency in maternal style. For example, captive rhesus infants show early and
consistent differences in their responses to stressful situations [see review, Suomi,
19871. Such differences could influence mother-infant interaction directly or indirectly by influencing maternal behavior. That such early differences may be in
part inherited [Suomi, 19871 could lead to similarities between mother-infant
pairs with the same mothers. So far, however, cross-fostering studies suggest that
such infant-based factors do not account for consistent maternal behavior [Suomi,
19871.
Although the consequences of various maternal styles are not well understood,
there is evidence that aspects of maternal style may influence both the infant's
psychosocial development and the mother's subsequent reproductive performance,
including the length of interbirth intervals [see reviews, Nicolson, 1982, 1987;
Gomendio, 1989; Johnson et al., in preparation). An implication of the consistency
of maternal style from infant to infant is that the long term consequences of
particular styles may accumulate over a lifetime and have profound effects on life
history variables. For example, t o the extent that maternal style influences the
length of interbirth intervals, particular styles could have large effects on the total
number of offspring a female produces in her lifetime. Similarly, to the extent that
maternal style is associated with infant mortality, morbidity [Altmann, 19801, or
the rate or quality of psychosocial development [e.g., Kaufman and Stynes, 1978;
Dolhinow, 19801, consistent styles could lead to more consistent risks or advantages for offspring of particular mothers.
168 / Berman
CONCLUSIONS
1. Individual rhesus mother-infant pairs on Cay0 Santiago, Puerto Rico, show
consistent differences in measures of mother-infant interaction over successive
infant age periods.
2. After the first 6 weeks, measures of interaction are more similar for
mother-infant pairs with the same mothers (i.e., families) than for pairs with
different mothers.
3. Differences in measures of mother-infant interaction between families are
also consistent over successive age periods and are due primarily to differences
between mothers rather than to differences between sets of infant siblings.
4. Maternal style is a term that can be used to describe both consistent differences in mother-infant relationships between individual pairs and between
families of free-ranging rhesus monkeys.
ACKNOWLEDGMENTS
I thank the Donald Sade, Richard Rawlins, Matt Kessler, and Dell Collins for
permission to carry out research on Cay0 Santiago; and L. Busaca, A. Homer, E.
Kapsalis, N. Printer, L. Ritchie, and L. Smith for assistance in data collection.
Cay0 Santiago was supported by NIH contracts DRR 71-2003 and RR-7-2115, and
by NIH grant RR-01293 to the University of Puerto Rico School of Medicine. I
received support from the Wenner-Gren Foundation for Anthropological Research,
the Explorer’s Club, Sigma Xi Society, NIMH predoctoral fellowship MH 05195,
NSF postdoctoral fellowship SPI-7815548, and NIMH grant MH38647.
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