American Journal of Primatology 38:303-314 (1996)

An Analysis of Birth Sex Ratio Bias in Captive

Prosimian Species
S. WATSON', J . WARD1, K. IZARD', A N D D. STAFFORD'
'Department of Psychology, The University of Memphis, Memphis, Tennessee;
'Department of Experimental Psychology, Duke University, Durham, North Carolina

The extent to which sex ratio bias is a common reproductive characteristic

of prosimians has not been well established. The present study analyzed
reproduction in 13 breeding groups of captive prosimians for evidence of
birth sex ratio bias. A substantial male bias was demonstrated in nongre-
garious, but not gregarious, breeding groups. Analyses of birth sex ratios
of individual mothers suggested that the observed bias did not result from
the tendency of a few mothers to overproduce males, but rather from a
small but reliable excess of male births in general. An examination of
infant mortality revealed that male Otolemur garnettii and Microcebus
murinus infants were more vulnerable to preweaning mortality, whereas
female Eulemur fulvus albifrons infants were more vulnerable. An anal-
ysis of birth order by sex found that mothers of one group (0.garnettii)
tended to produce males initially and females later. Additionally, a dis-
tinct pattern of birth seasonality was noted among Malagasy prosimians
that was absent in the African prosimians. Greater length of period of
sexual receptivity for nongregarious females as compared to gregarious
females is proposed as a possible mechanism of male birth sex ratio bias.
0 1996 Wiley-Liss, Inc.

Key words: birth sex ratio bias, prosimians, birth order

INTRODUCTION
Sex ratio bias exists for any species when the sex ratio of the male and female
progeny differs systematically from 1:l.Fisher [ 19301 predicted that when parents
invest equally in male and female offspring, an unbiased sex ratio will be main-
tained. However, for some species it may be the case that reproductive fitness is
enhanced by a tendency to produce greater numbers of either male or female
offspring [e.g., Trivers & Willard, 1973; Clark, 19781. Previous research suggests

Received for publication July 24, 1994; revision accepted August 29, 1995
Address reprint requests to Jeannette P. Ward, Ph.D., The University of Memphis, Department of
Psychology, Memphis, TN 38152.
Sheree Watson's present address is Bowman Gray School of Medicine, Wake Forest University, CMCRC-
Friedberg Campus, Medical Center Boulevard, Winston-Salem, NC 27157-1040.
Donna Stafford's present address is Department of Biological Structure, University of Washington,
Seattle, WA.

0 1996 Wiley-Liss, Inc.

304 I Watson et al.
that this adaptive strategy may occur in a number of mammalian species, includ-
ing primates [for review, see Clutton-Brock & Iason, 1986; Hrdy, 19881.
The attention of the authors was drawn to the issue of sex ratio bias in pros-
imians by the observation of a disproportionately large number of male births in
the breeding population of small-eared bushbabies (Otolemur garnettii) at the Uni-
versity of Memphis (UM). There is evidence that some prosimian species produce
a disproportionate number of male offspring in both wild and captive populations
[e.g., Clark, 1978, 0. crassicaudatus; Foerg, 1982, Cheirogaleus medius; Perret,
1990, Microcebus murinus; Izard & Simons, 1986, Galago moholi, 0. garnettii, 0.
crassicaudatus]. However, the extent to which a male sex bias is a general char-
acteristic of prosimian species has not been well established. Therefore, a statis-
tical analysis was conducted of the birth sex ratio of the UM colony as well as the
prosimian breeding groups resident a t the Duke University Primate Center
(DUPC).
In addition to analyzing birth records for evidence of sex ratio bias, other
analyses were conducted on these records in order to gain some insight into pos-
sible factors that might illuminate the functions and/or mechanisms of biased sex
ratios. The theory of differential mortality [Clutton-Brock & Iason, 1986; Leigh,
19701 suggests that a surplus production of males might be desirable if males are
especially vulnerable to preweaning mortality. Thus, infant deaths were examined
for evidence of a sex bias during the period of parental investment. Also, if male
infants were more fragile than female infants, male births might be expected to be
timed such that optimal environmental support is available for the development of
young [Clutton-Brock & Iason, 19861. For this reason, seasonality of births was
examined for evidence of preferential production of males during particular sea-
sons.
In the natural habitat, sex ratio biases may be related to the differential
emigration of young. Local resource competition (LRC) theory predicts that birth
sex ratios should deviate from unity if one sex is more likely to emigrate [Clark,
19781. The emigrating offspring do not compete with the mother for resources,
whereas the nonemigrating offspring may expropriate maternal resources and
thereby jeopardize the mother’s subsequent reproductive success. It is apparent
that, under these circumstances, the mother’s reproductive fitness would be en-
hanced by production of greater numbers of the emigrating sex. Among prosimian
species, males are typically the emigrating sex. This strategy would be maximally
adaptive if the emigrating sex were produced early in the mother’s reproductive
career, thereby reducing the duration of the deleterious effects of the competing
sex [Clutton-Brock & Iason, 19861. An analysis of birth order for sex bias was
conducted to determine whether this maximally adaptive strategy could be ob-
served in captive prosimians.
METHODS
Subjects
Archival records of prosimian births at two established breeding colonies, the
UM and the DUPC, were examined for evidence of sex ratio biases. The birth
records spanned the period between 1960 and 1993, collectively. Of the 27 breeding
groups for which records were available, only the 14 for which 30 or more births
occurred were chosen for analysis. The breeding groups and evaluation periods
were as follows: Cheirogaleus medius, 1969-1988; Eulemur fuluus albifrons, 1970-
1989; E . f . collaris, 1964-1989; E . f . fuluus, 1962-1987; E . f . rufus, 1961-1989; E.
macaco, 1969-1989; Galago moholi, 1961-1990; Lemur catta, 1967-1989; Mirza
coquerli, 1983-1987; Microcebus murinus, 1971-1988; Otolemur crassicaudatus,
 Birth Sex Ratio Bias in Prosimians J 305
1961-1990; 0. gurnettii, DUPC, 1960-1990, UM, 1985-1993; and Vurecia uurie-
gutu, 1966-1989. Only one breeding group from the UM colony, 0. gurnettii, met
this requirement. For most measures, the 0. gurnettii of the UM and DUPC col-
onies were analyzed as separate populations.
Analyses
Sex ratio. Birth sex ratios of individual mothers were analyzed for bias in
order to evaluate statistical independence prior to group analyses. Only mothers
with five or more infants (binomial P = 0.55 = 0.03) for whom sex had been
determined were included in this analysis. Previous research has suggested that
prosimian twin births are dizygotic [Pasztor & Van Horn, 19761. Evaluation of the
twin births in the present study supported this assumption in that of 394 twin
births, 46.5% were male-female twin pairs whereas 32.5% were male twins and
21.1% were female twins. Therefore, multiple births were statistically treated as
independent events. To permit evaluation at several taxonomic levels, group anal-
yses were conducted by evaluating z scores in terms of the normal curve distribu-
tion for the entire group of prosimians assessed, and for gregarious and nongre-
garious prosimians separately, as well as a t the levels of genus, species, and
breeding group. The analysis of gregarious species separately from nongregarious
species was required by the observation of systematic differences in the birth sex
ratio data. Categorization of prosimian species as gregarious or nongregarious was
made in accordance with the work of Charles-Dominique [1978], Bearder [1987],
and Nash [19931. Analyses were conducted by breeding group because each breed-
ing group in this study has been maintained as a separate and distinct population.
There were some infants of each breeding group for whom sex was not recorded (a
group average of 9% of all recorded births) and who were necessarily excluded from
this analysis.
Differential mortality. Percentages of male and female infants that died
within 1month of birth were assessed via tests of significance of proportions, using
the formula z = PI - P,/square root of p(1 - p)/N, + p(1 - p)/N,, where p equals
NIP, + N$,/N, + N , [Bruning & Kintz, 19681. One month was defined as a
calendar month. For example, an infant born on January 1was considered 1month
old on February 1.
Seasonality. Male and female births per month were depicted graphically
and inspected for evidence of differential seasonality by sex. Percentages were
calculated for males and females separately and displayed as monthly percentages
of total male or female births per breeding group.
Birth order. The relationship between mother’s parity and sex of offspring
was assessed via a 2 x 3 x2 contingency table to determine whether infants occu-
pying a given birth position differed from the expected value of 50% male offspring.
In order that parity could be properly assigned, only colony-born mothers were
used in this analysis. Because there was a preponderance of first- and second-born
infants relative to later-born infants, births were categorized for analysis as first-
born, second-born, or third- and later-born infants which composed 31.0%, 22.8%,
and 46.2%, respectively, of the 1,335 births included in this analysis. Statistical
analysis of this data was conducted only by species.
RESULTS
Sex Ratios of Individual Mothers
The mothers in this study produced an average of 4.47 infants. By breeding
groups, the average births per mother ranged from 2.88 to 6.59 infants. Table I
summarizes the results of the analysis of birth sex ratio of the individual mothers.
306 I Watson et al.
TABLE I. Individual Mothers With Birth Sex Ratio Biases Greater Than Chance (L = >
1.96,P < 0.05) Shown With Total Number of Mothers Per Breeding Group and Number
of Mothers Per Breeding Group That Had Five or More Births
Mothers Birth
Total with >5 Individual sex
Breeding group mothers births mothers ratios (M:F) z score
C . medius 29 12
E. f. albifrons 19 11
E . f. collaris 12 3
E . f. fulvus 36 14 Ino 7:1 2.13
E . f. rufus 36 10
E. macaco 17 8
G . moholi 57 16
L. catta 27 12
M. coquereli 14 5
M . murinus 33 15
0. crassicaudatus 70 23 Iago 12:2 2.67
0.garnettii-DUPC 65 9 White-Tail 8:1 2.30
0. gurnettii-UM 18 7 Hepburn 9:1 2.53
V. uariemta 37 16 Selene 0:5 2.23

This table shows the total number of mothers per group as well as the number of
mothers per group that gave birth to five or more infants during the study period.
Although five breeding groups contained an individual mother who produced a
disproportionate number of same sex infants, no group had more than one female
who displayed a significant sex ratio bias. Therefore, there was insufficient evi-
dence to conclude that inequity in sex ratios reflected the tendency of particular
females to overproduce offspring of either sex.
Examination of the total data set suggested that the observed biases resulted
from a small but persistent birth sex ratio bias in those breeding groups that did
produce a preponderance of one sex. To evaluate this hypothesis with respect to
each breeding group, the number of mothers that produced more male than female
offspring was compared to the number of mothers that produced more female than
male offspring. Analysis of the 14 breeding groups found only three that had more
mothers with a birth sex ratio bias: 0. garnettii-DUPC, x2 = 5.09, P < 0.025; 0.
garnettii-UM, x2 = 4.64, P < 0.05; M. murinus, x2 = 5.58, P < 0.025. In all three
of these groups there were more mothers that produced a simple majority of male
offspring.

Sex Ratio Analysis by Group

An initial analysis of the ratio of male (n = 1,126) to female (n = 977) births
by z score revealed that there was a male bias for this entire prosimian group (z =
3.25, P < 0.01). Results of analyses by genus, species, and breeding group are
shown in Tables I1 and 111. At the level of genus, sex ratio biases were found only for
Microcebus and Otolemur. The three breeding populations found to have sex ratio
biases were M. murinus, 0. garnettii-MSU, and 0.garnettii-DUPC, and births of
males were predominant for these. All significant sex ratio biases occurred in
nongregarious species. Furthermore, it was noted that all but one (M. coquereli) of
the other nongregarious species had a majority of male births. Therefore, the
possibility of a male bias in nongregarious species as a group was examined by
summing the total number of male and female births of these and calculating a z
 Birth Sex Ratio Bias in Prosimians / 307
TABLE 11. Sexes of Infants Born to Nongregarious Prosimians and
Deaths Occurring Within 30 Days Postpartum
Group Born 2 Died (%) z
Total nongregarious
Male 638 3.93*** 132 (20.7%) 0.71
Female 505 96 (19.0%)
Cheirogaleus (C. nedius)
Male 104 1.31 18 (17.3%) -1.57
Female 86 23 (26.7%)
Galago (G. rnoholi)
Male 104 1.54 35 (33.7%) 1.01
Female 83 22 (26.5%)
Mirza ( M . coquereli)
Male 30 -0.62 5 (16.7%) -0.62
Female 35 8 (22.9%)
Microcebus ( M . murinus)
Male 92 2.07* 18 (19.6%) 2.11*
Female 66 5 (7.6%)
Otolemur (sp.)
Male 308 2.87** 86 (27.9%) 1.54
Female 235 52 (22.1%)
0. crassicaudatus
Male 150 1.57 49 (32.7%) 0.80
Female 124 35 (28.2%)
0.garnettii
Male 158 3.13** 37 (23.4%) 1.8
Female 111 17 (15.3%)
DUPC subgroup
Male 107 1.97* 28 (26.2%) 2.07*
Female 80 11(13.8%)
UM subgroup
Male 51 2.21* 9 (17.6%) -0.20
Female 31 6 (19.4%)
*P < 0.05.
**P < 0.01.
***P < 0.001.

score: total male births exceeded total female births (z = 3.93, P < 0.001). A
similar analysis did not yield significant results for gregarious prosimian species
(z = 0.52, NS). An independent samples t test verified that the z scores of the
nongregarious breeding populations (mean z = 1.44) were reliably different from
the z scores of the gregarious groups (mean z = 0.283), t(12) = 2.64, P = 0.02.
Thus, it may be asserted with confidence that the male birth sex ratio bias found
in the total group of prosimians resulted from the contribution of the nongregar-
ious breeding groups.

Differential Mortality
As shown in Tables I1 and 111, few breeding groups had differential mortality
by sex. The percent attrition was calculated separately for male and female infants
as compared to the numbers of male and female infants born. Greater proportions
of male than female infants died during the first month of life in two breeding
groups, M . murinus ( z = 2.11, P < 0.05) and 0. garnettitDUPC (z = 2.07, P <
308 I Watson et al.
TABLE 111. Sexes of Infants Born to Gregarious Prosimians and Deaths
Occurring Within 30 Days Postpartum
Group Born 2 Died (%I 2

Total gregarious
Male 488 0.52 77 (15.8%) -1.38
Female 472 90 (19.1%)
Eulemur (sp.)
Male 303 0.72 60 (19.8%) -0.51
Female 279 60 (21.5%)
Eulemur fuluus (sp.)
Male 252 0.72 50 (19.8%) -0.14
Female 236 48 (20.3%)
E . f. albifrons
Male 66 1.19 7 (10.6%) -2.01*
Female 53 13 (24.5%)
E. f. collaris
Male 25 0.29 8 (32.0%) 1.21
Female 23 4 (17.4%)
E. f. fuluus
Male 78 -0.39 11 (14.1%) -0.10
Female 83 13 (15.7%)
E. f. rufus
Male 83 0.47 24 (28.9%) 0.80
Female 77 18 (23.4%)
Eulemur macaco
Male 51 0.82 10 (19.6%) -1.17
Female 43 12 (27.9%)
Lemur (L. catta)
Male 68 0.24 8 (11.8%) -0.05
Female 66 8 (12.1%)
Varecia (V. uariegatu)
Male 117 -0.64 21 (17.9%) -0.94
Female 127 29 (22.8%)
*P < 0.05.

0.05). Conversely, there was evidence for greater female mortality during the first
month of life in E.f. albifrons ( z = -2.01, P < 0.05).

Seasonality
Sample sizes of male and female births per breeding group distributed over a
12 month period did not permit inferential statistical comparisons. The proportion
of births by month was calculated separately for males and females as a percentage
of total births for each sex. Graphic depiction of monthly births showed evidence of
distinct birth seasons in L. catta, E. f. albifrons, E. f. collaris, E. f. fuluus, E . macaco,
E. f. rufus, V. uariegata, C . medius, and M. murinus. In contrast, G . moholi, M.
coquereli, 0. crassicaudatus, 0. garnettii-DUPC, and 0.garnettii-UM reproduced
throughout the year and displayed no distinctive peak in births. Graphic depiction
failed to display evidence that births of either sex were favored in any given month
or season (Figs. 1,2). Rather, male and female births appeared to occur with ap-
proximately equivalent probability in any given month. Species with a preponder-
ance of male births had a small but stable excess of males across the calendar year.
 Birth Sex Ratio Bias in Prosimians / 309

40

'"I
33

!O

10

JM Fan Msr Apt May JWI JU aug SSP OCt NOV D k

4D
f

33

20

10

J n FSD I M r &X my JUI JW Wg Qap OCI NW D&

Percentage of Male and Female Births

Male Female
Fig. 1. Percentage of male (black) and female (hatched) births by month for gregarious species. a: E. f. albi-
frons; b: E . f. collaris; c: E . f. fuluus; d E . f. rufus; e: L. catta; E E. macaco; g:.'1 uariegata. Births by months
were calculated as a percentage of total same sex births per breeding group.

Birth Order
Deviations from the expected frequencies of male and female offspring pro-
duced in first, second, and later births occurred in three species, E. f . rufu..s (x2 =
310 I Watson et al.

40 *G
a (b i
I

10

JW FeD Mer &r May Jun JuI W Q Sep OCL NOv D&

_I

30-

10

JW FeD Me, npr Mey Jun JUI L\uD SBp OCI NW Dffi

33

20

10

0
Jm FsD MI apr M y Jun JUI /uq Sap Ocl N~hr DBC

10

JW FED Mar Apr May JuO JUI Aug Sep OCI NW DeE

Percentage of Male and Female Births

Male Female
Fig. 2. Percentage of male (black) and female (hatched) births by month for nongregarious species. a: C.
medius; b M. murinus; e: M. coquereli; d G. moholi; e: 0.crassicaudatus; fi 0.garnettii-DUPC; g: 0.garnettii-
MSU. Births by months were calculated as a percentage of total same sex births per breeding group.

9.419, P = 0.0091, M. coquereli (x2 = 6.57,P = 0.0371, and 0.gurnettii (x2 = 6.51,
P = 0.0385). The deviation from expected frequencies in E . f : rufus appeared to
result from a disproportionately large number of males as second births (17 males
vs. three females) as did the deviation from expected values in M. coquereli (nine
males vs. two females). The deviation from expected values in 0. gurnettii, how-
ever, reflected a preponderance of males as first births (30 males vs. 12 females).
 Birth Sex Ratio Bias in Prosimians / 311
DISCUSSION

For breeding groups that did have biased birth sex ratios, an examination of
individual mother’s ratios showed that the bias resulted from a small but consis-
tent excess of male offspring across the mother populations, rather than from male
overproduction by a few aberrant females. This finding is compatible with the
views of Clarke [1978],whose LRC theory predicts a group-characteristic birth sex
ratio bias. The results of the present study are also somewhat compatible with
”rivers and Willard’s [19731 theory which hypothesizes that individual mothers’
birth sex ratio biases are influenced by maternal nutritional status. For these
captive breeding groups, food was readily available throughout the year and nu-
tritional status has been shown to influence the reproductive parameter of twin-
ning in L. cuttu [Pereira, 19931. However, if diet was a factor in this study of birth
sex ratio bias, its influence was limited to the nongregarious breeding groups.
Although nongregarious species contributed only half of the data points in this
sample, what initially appeared to be a male bias in birth sex ratios for the entire
sample was, in fact, a function of the preponderance of male births for nongregar-
ious prosimians only. All except one of these groups had a greater number of male
than female infants born, a systematic bias not found among the gregarious
groups. This male bias was especially pronounced within the genus Otolemur and
the species M. murinus. The male sex ratio bias in these captive groups of Otole-
mur parallels that reported in field studies of 0. garnettii “ash, 19831 and 0.
crassicaudatus [Clarke, 19781. The congruence in sex ratio bias between captive
and wild bushbabies suggests that the phenomenon may be only slightly depen-
dent upon environmental cues. However, Perret’s [1990] study of M . murinus in-
dicated that environmental cues may influence birth sex ratio bias in captive
prosimians. She found that females were more likely to have sons when housed in
proximity to other females and suggested possible mediation of this effect by ol-
factory and/or auditory cues. Because the nongregarious prosimians resident a t
DUPC and UM were housed indoors, such cues would have been concentrated and
may have contributed to the birth sex ratio bias effect.
Clarke [1978] proposed that male-biased birth sex ratios are adaptive for wild
populations from which male offspring emigrate and female offspring remain on or
near the natal territory and compete for resources. Adult females of nongregarious
prosimian species occupy a small foraging territory, solitary except for the inter-
mittent visitations of adult males and periods of resource sharing with subadult
young. Social systems which feature emigration of pubescent males are character-
istic of many of the nongregarious prosimian species [Bearder, 19871. In captive
situations, both male and female infants are typically removed from the mother at
some time between weaning and pubescence, but emigration, as such, does not
occur. Even if proximal cues resulting from male emigration were important to
birth sex ratio biases in natural habitats, it is not likely that they could be a
significant factor in captive colonies. Thus, we propose that the male birth sex ratio
bias in nongregarious prosimians is mediated by an endogenous mechanism.
A characteristic of nongregarious prosimian females that suggests itself as a
possible mechanism in the control of birth sex ratio bias is found in the periods of
female sexual receptivity, which are quite long. Periods of receptivity for nongre-
garious prosimian females are measured in terms of days, whereas those for gre-
garious females are measured in terms of hours [Hrdy & Whitten, 19861. Several
researchers have proposed that the time of fertilization within the estrous period
influences the gender of the resultant offspring [James, 1971,1976,1980; Harlap,
1979; Guerrero, 1974; Pratt et al., 19871. Specifically, it has been suggested that
312 I Watson et al.
human female concepti result from inseminations closer to the ovulatory event and
male concepti result from inseminations further removed in time from ovulation.
If a similar mechanism operates in prosimians, a nongregarious species would
have a greater window of opportunity for producing a male. It is likely that the
long period of receptivity for nongregarious female prosimians evolved to accomo-
date a social system in which the male was not continuously present [Bearder,
1987; Charles-Dominique, 19771; however, this does not preclude such a n adapta-
tion from constituting a mechanism of birth sex ratio bias.
An examination of birth order by sex revealed that primiparous 0. garnettii
mothers tended to produce a preponderance of male offspring. If reproductive strat-
egies in wild 0.garnettii populations are influenced by competition between moth-
ers and daughters for resources, it would be adaptive for mothers to produce males
early and females later [Clutton-Brock & Iason, 19861. This would reduce the total
amount of time that the mother and her daughters would compete for resources,
thereby maximizing the reproductive fitness of both mother and daughter. The
present analysis included only mothers who were born in captivity and had not
experienced resource competition. The fact that these well provisioned mothers
produced a preponderance of male offspring as early births further supports the
proposal of a n endogenous reproductive mechanism which is independent of re-
source availability.
This study found only limited evidence for differential mortality by sex during
the first month of life. The excess of early male deaths for 0. gurnettzi-DUPC and
M. murinus could have been predicted by Fisher’s [19301 proposal that differential
infant mortality by sex for a given breeding group should be associated with higher
birth rates for the less viable sex. However, factors contributing to the selective
preweaning mortality of male infants are as yet undetermined. It should be noted
that differential male mortality occurred in the DUPC colony of 0. garnettii but
was absent in the UM colony. This discrepancy may be attributed to the fact that
in the DUPC colony mothers and their infants were, at one time, housed contin-
uously with the sires [Izard & Simons, 19861. This practice was never employed in
the UM colony. 0. gurnettii adults have been reported to kill neonates when the
mother was not sufficiently protective [Welker & Welker, 19891. Allowing sires to
remain with mothers and neonates could result in the selective deaths of male
infants if primiparous mothers were less protective than experienced mothers.
No seasonal differences were noted for births of male and female offspring for
any breeding group. The observation of birth seasonality in the gregarious breed-
ing groups is consistent with previous reports for both wild and captive populations
[Rasmussen, 1985; Richard, 19871. However, the absence of seasonality in most of
the nongregarious breeding groups differs from field observations [Bearder, 1987;
Nash, 19831. The Malagasy lemurs exhibited more pronounced birth seasonality
than the African bushbabies. The gregarious Malagasy species were housed in
outdoor enclosures and were thus exposed to the normal spectrum of environmen-
tal cues. In contrast, the nongregarious species, all nocturnal in habit, were cage-
housed indoors and, for portions of the evaluation periods, some were maintained
on 12:12 light/dark cycles. Such maintenence has been shown to disrupt birth
seasonality in 0. crassicaudatus [Izard et al., in press; Eaglen & Simons, 19801.
The only two nongregarious species to maintain birth seasonality under the indoor
conditions were the Malagasy lemurs, M. murinus and C . medius. These two spe-
cies undergo seasonal torpor [Petter-Rousseaux, 19801 which may have resulted in
periodic absence of reproductive activity. Without specific information about cycles
of illumination for the African species housed indoors, a detailed analysis of birth
seasonality as a function of lighting conditions was not possible. However, the data
 Birth Sex Ratio Bias in Prosimians I 313
do suggest that birth seasonality is not a robust phenomenon in bushbabies or
galagos. Whether this constitutes a reliable distinction between African and Mal-
agasy prosimians is a question that requires further study.
Major advantages inherent to this archival study of captive prosimian popu-
lations were the very large number of recorded births collected over a n extended
period of time and the certainty of reliable sex identification. In general, our
results were congruent with those from wild-living populations. Because captive
circumstances facilitate the manipulation of variables that may influence birth
sex ratio bias, experimental study of this phenomenon in captive populations
should be pursued to evaluate adaptive theories of birth sex ratio bias and to
explore the physiological mechanisms by which this bias is mediated.

CONCLUSIONS
1. Male-biased birth sex ratios were evident for nongregarious, but not gre-
garious, captive prosimians.
2. The observed male birth sex ratio bias reflected the tendency of most moth-
ers to produce a small but reliable preponderance of male offspring.
3. Individuals of one species, 0.garnettii, displayed a tendency to produce male
offspring as first births.
4.The distinct pattern of birth seasonality observed in captive Malagasy spe-
cies was absent in captive African species.
ACKNOWLEDGMENTS
This research was supported in part by National Science Foundation Grant
BNS-8707754 to Jeannette P. Ward. Support was also received from a Centers of
Excellence grant awarded to the Department of Psychology, Memphis State Uni-
versity, by the State of Tennessee. We are grateful to the Duke University Primate
Center, Durham, North Carolina, for access to their records. This is DUPC Re-
search Report 573.
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