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Effect of the enclosure on carriers' body weight loss in the cotton-top tamarin (Saguinus oedipus).

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American Journal of Primatology 66:279–284 (2005)
BRIEF REPORT
Effect of the Enclosure on Carriers’ Body Weight Loss
in the Cotton-Top Tamarin (Saguinus oedipus)
SUSANA SÁNCHEZ1,2n, FERNANDO PELÁEZ1,2, ANA MORCILLO1, and CARLOS
GIL-BÜRMANN1
1
Área de Psicobiologı´a, Facultad de Psicologı´a, Universidad Autónoma de Madrid,
Madrid, Spain
2
German Primate Center, Göttingen, Germany
Infant carrying in cotton-top tamarins is a costly activity that results in
weight loss by the fathers and helpers. However, to date, measures of
carrying costs have been obtained in small indoor enclosures. We studied
body weight changes in adult and subadult individuals from eight large
groups (n45) for 9 weeks after the birth of infants. Four groups were
housed in large indoor/outdoor enclosures (42 m2 3.3 m high), and four
were housed in small indoor enclosures (12 m2 2.4 m high). All of the
individuals were weighed regularly at least three times a week.
Reproductive males lost more weight in the big indoor/outdoor enclosures
(mean=6.51%) than in the small indoor ones, as did male adult helpers
(mean=5.59%) and female adult helpers (mean=4.4%). Still-growing
subadult individuals also lost weight in the big indoor/outdoor enclosures
(mean=3.17%), although the differences did not reach significance
(P=0.07). These results support the hypothesis that cotton-top tamarins
living in more natural settings experience higher weight loss than those
housed under less-natural conditions. Am. J. Primatol. 66:279–284,
2005.
r 2005 Wiley-Liss, Inc.
Key words: enclosure effect; body weight loss; carrying costs; Saguinus
oedipus
INTRODUCTION
In cotton-top tamarins (Saguinus oedipus), the fathers and adult and
subadult individuals of both sexes (‘‘helpers’’) cooperate with the carrying of
infants that represent 15% of the mother’s weight [Snowdon, 1996; Tardif et al.,
1993]. Using data from saddle-back tamarins in the wild [Goldizen, 1987], Tardif
Contract grant sponsor: MCyT-DGI; Contract grant numbers: PB-98-0094; BSO-2002-02611;
Contract grant sponsor: Universidad Autónoma de Madrid/German Primate Center; Contract grant
sponsor: Universidad Autónoma de Madrid; Contract grant number: FPU-2003.
n
Correspondence to: Susana Sánchez, Área de Psicobiologı́a, Facultad de Psicologı́a, Universidad
Autónoma de Madrid, 28049 Madrid, Spain. E-mail: susana.sanchez@uam.es
Received 26 April 2004; revised 18 October 2004; revision accepted 7 February 2005
DOI 10.1002/ajp.20144
Published online in Wiley InterScience (www.interscience.com).
r
2005 Wiley-Liss, Inc.
280 / Sánchez et al.
[1997] estimated that carrying two 30-day-old infants represents for an adult an
increase of 21% in their energy expenditure. On the other hand, foraging and
feeding time is reduced while carrying is performed [Price, 1992a; Tardif, 1994].
Sánchez et al. [1999] found that after the birth of infants in indoor captive groups
of cotton-top tamarins, fathers and male helpers lost weight, and this weight loss
was related to their participation in carrying. The results also showed that
tamarins’ feeding time and energy intake decreased while they were carrying
infants, although no relation was found between these two variables and weight
loss. In contrast, other studies showed that lactating mothers progressively
gained weight as they reduced down their contribution to carrying, mainly during
the time when a new pregnancy could take place (weeks 4–5) [Ziegler et al., 1990],
and as a consequence of increased food intake [Kirkwood & Underwood, 1984;
Price, 1992b; Sánchez et al., 1999]. Furthermore, Achenbach and Snowdon [2002]
found that after births occurred in cotton-top tamarin groups, the adult males
also lost weight. However, in this case the weight loss was not found to be related
to carrying time. When Achenbach and Snowdon [2002] analyzed weight loss in
adult males of different size groups, they observed that the loss was higher in
small groups because infant carrying was shared by fewer individuals.
Captive studies in which measures of carrying costs were obtained have been
conducted in groups living in small indoor enclosures (e.g., 2 m2 by 2 m high
enclosures: Anthropological Institute, University of Zürich (C. jacchus [Nievergelt & Martin, 1999]); 4 m2 by 2.2 m high enclosures at the Psychology
Department, Wisconsin University (S. oedipus [Achenbach & Snowdon, 2002];
and 12 m2 by 2.4 m high enclosures at the German Primate Center, Göttingen
(S. oedipus [Sánchez et al., 1999])). However, cotton-top tamarin groups in the
wild occupy territories of 10.5–12.4 ha, most of which can be covered in 1–2 days
[Savage, 1990]. We expect that in the wild, the effort of carrying infants will have
a more dramatic effect on carriers’ weight loss. We predicted that even in
captivity, but in a more naturalistic environment (i.e., large indoor/outdoor
enclosures), carriers would lose more weight than in small indoor enclosures.
MATERIALS AND METHODS
Adult and subadult individuals from eight large groups (n45) of cotton-top
tamarins were observed. During the study, mothers gave birth to 13 infants (five
twin litters and three singleton litters). Juvenile individuals were excluded from
the analysis because their contribution to carrying is very limited [Price, 1992c].
Four groups at the German Primate Center of Göttingen (DPZ), were housed
in relatively small indoor enclosures (12 m2 2.4 m high) and kept at a
temperature of 261C, humidity of 65%, and light length cycle of 12 hr. The other
four groups were part of the tamarin colony at the Universidad Autónoma de
Madrid (UAM) and were housed in indoor enclosures (3.5 m2 2.2 m high) at a
temperature of 261C. The animals had continuous free access to exceptionally
large outdoor areas (42 m2 3.3 m high) that were well furnished with branches
and other enrichment elements. To avoid direct solar radiation, 80% of the
outdoor areas were covered by shading nets. A system of automatic water
sprinklers was placed outdoors to create mist to maximize cooling and humidity.
The group sizes and compositions are listed in Table I.
The weather conditions at the UAM colony are given in Table II. The
breeding period of two groups (F and G) took place during spring, and the
breeding periods of the other two groups took place in autumn (E) and in late
autumn through early winter (H), respectively.
Enclosure Effect on Carrier Weight Loss / 281
TABLE I. GROUP COMPOSITION of the Observed Families by Classes
Classes
Adult
a
Subadult
Juvenile
Group
n
Enclosure
No.infants
#
~
#
~
#
~
A
B
C
D
E
F
Gb
H
13
10
8
6
8
8
8
10
DPZ
DPZ
DPZ
DPZ
UAM
UAM
UAM
UAM
2
1
2
2
1
2
2
1
4
3
1
1
2
4
3
4
4
3
1
1
3
3
3
4
1
1
2
1
2
–
2
–
2
1
2
1
–
1
–
–
–
1
2
–
1
–
–
1
2
1
–
2
–
–
–
1
a
The adult males and adult females also include fathers and mothers, respectively.
An adult female and a subadult male, who were included in this table, were expelled from the group before week
8 in the study. They were not included in the analyses for that reason. Adults, 424 months; subadults, 13–24
months; juveniles, 6–12 months [Price, 1991, 1992a].
b
TABLE II. Weather Conditions at the UAM Colonyn
Group
RP
Season
M-min.
M-max.
M-daily
M-[13]
H
M-Sunshine
E
B
N-B
B
N-B
B
N-B
B
N-B
A-A
W-W
S-S
W-W
S-S
W-S
A-W
A-A
5.5
1.6
7.4
3.6
11.6
4.9
4.5
6.4
14.9
15.7
17.9
18.6
24.2
16.9
11.1
18.1
10.35
7.9
12.7
10.7
18.1
10.6
7.6
11.9
14.5
13.8
16.3
16.7
22.3
15.1
10.1
16.8
58.5
48.8
54.3
45.0
47.6
50.8
68.9
49.7
10:20
10:16
13:35
11:08
14:28
12:19
9:27
10:42
F
G
H
n
RP, reproductive period; B, breeding (9 weeks); N-B, non-breeding (7 weeks).
Season (onset-end of reproductive period): S, spring; A, autumn; W, winter. Temperature: M, mean (1C); M-[13],
mean at 13’00 CET; H, humidity (%); M-sunshine, daily mean sunshine duration (hh:mm).
UAM and DPZ used the same animal diets and feeding schedules, both of
which were strictly controlled during the study to avoid body weight fluctuation
caused by a variation in food supply. When infants were born, the groups were
provided with proportional extra food. The animals had free, easy access to food
placed indoors. Details regarding the diet, housing conditions, and feeding
schedule at DPZ are provided elsewhere [Sánchez et al., 1999].
All of the groups were observed for 9 weeks following the birth of infants.
This breeding period includes the infant total-dependence phase during the first 4
or 5 weeks [Kirkwood & Stathatos, 1992], when carrying costs are the highest
[Sánchez et al., 1999], and the beginning of independence. In the 9th week,
infants spend 50% of their time moving independently (in captivity [Price, 1992b]
and in the wild [Savage et al., 1996]). All individuals were routinely weighed on a
scale (71 g) (Sartorius Universal Type 46100) at least 3 times a week. The
noninvasive procedure used was previously described by Sánchez et al. [1999]. To
calculate weekly body weight changes, we subtracted the weekly mean weight
from the prebirth weight (i.e., the weekly mean weight of the last week prior to
birth). For reproductive females, their weight on the day infants were born was
282 / Sánchez et al.
used as the prebirth weight. All groups were also weighed during nonbreeding
periods (7 weeks).
Nonparametric Mann-Whitney U-tests and Wilcoxon T-tests were used for
most of the comparisons. Since we had predicted that after the birth of infants the
tamarins in big indoor/outdoor enclosures would experience greater body weight
losses, the Mann-Whitney U-tests for these analyses were one-tailed. A
significance level of Po0.05 was considered for all analyses.
RESULTS
Before the births of infants, all of the reproductive males, adult helpers, and
subadults at UAM weighed less than those at DPZ (mean=551.76777.80 g vs.
mean=634.84773.63 g; U=135, n1=25, n2=25, Po0.001). To be sure that the
difference in conditions between UAM and DPZ did not have an influence on
weight changes in the absence of births, we compared body weight changes of
adult tamarins in DPZ and UAM during a nonbreeding period of 7 weeks. No
differences were found between the nonbreeding periods of the two colonies
(U=108, n1=14, n2=21, P=0.18).
On the other hand, although no differences were found in the weekly mean
temperatures between breeding and nonbreeding periods at the UAM (U=439.5,
n1=36, n2=28, P=0.38), individuals lost more weight during the breeding periods
(T=13, n=21, Po0.001). During the breeding periods, temperatures were higher
in the spring breeding periods than in the autumn-winter ones (U=46, n1=18,
n2=18, Po0.001), but we found no differences when we compared weight losses
(U=65, n1=12, n2=13, P=0.48).
As Fig. 1 illustrates, when we compared the means of body weight losses after
infant births in the two different locations, we found that reproductive males lost
more weight in the UAM enclosures (mean= 6.51%73.4%) than in the DPZ ones
±Std.Dev.
6
±Std.Err.Big indoor/outdoor enc.
±Std.Err.Small indoor enc.
% Mean body weight change
4
Mean
2
0
-2
*
**
**
-4
-6
-8
-10
Fathers
Ad.Males
Ad.Females
Subadults
Mothers
Fig. 1. Body weight changes in fathers, adult male and female helpers, subadult helpers, and
mothers in big indoor/outdoor enclosures and small indoor enclosures.
Enclosure Effect on Carrier Weight Loss / 283
(mean= 2.2%72.88%) (U=2, n2=4, n1=4 P=0.04), and the same applied to
adult male helpers (mean= 5.59%72.43% vs. mean= 0.88%70.36%) (U=1,
n2=5, n1=9, Po0.01) and adult female helpers (mean= 4.4%71.88% vs.
mean=0.48%71.19%) (U=0, n2=5, n1=8, Po0.01). In the case of subadults,
we found the same tendency as before (mean= 3.17%74.26% vs.
mean=1.15%75.12%); however, this result did not reach statistical significance
(U=11, n1=4, n2=11, P=0.07). We also found no differences in mothers’ weight
changes between the UAM and DPZ enclosures (mean= 0.14%73.68% vs.
mean=1.33%73.36%) (U=7, n1=4, n2=4, P=0.38).
DISCUSSION
The results of this study support the notion that infant carrying in cotton-top
tamarins is a costly activity, even in captive conditions, since both reproductive
males and helpers experience weight loss after the birth [Sánchez et al., 1999]. We
suggested that it would translate into higher body weight losses in the wild, and
predicted that even in captivity–but in a more naturalistic environment, with big
indoor/outdoor enclosures–carriers would lose more weight than carriers living in
small indoor enclosures. Our results indicate that after the births of infants,
fathers, adult male helpers, and adult female helpers experienced more weight
loss in the UAM enclosures than in the DPZ ones (Fig. 1). The highest body
weight losses were 16.85% for fathers, 20.23% for adult males, and 14.70% for
adult females at the UAM enclosures. The fact that individuals weighed less at
UAM than at DPZ before the births emphasizes the risk for higher relative weight
losses in the more natural setting.
Considering our results and those of Achenbach and Snowdon [2002], one
may conclude that even in captive environments, where food availability is
guaranteed and there is no need to travel for foraging, both the number of
individuals in the group and the characteristics of the enclosure are related to
weight loss. The larger territory provided by the big indoor/outdoor enclosures of
UAM may impose higher carrying efforts compared to the restricted conditions
under which captive studies are usually conducted. Since abiotic factors, such as
the length of day, temperature, and humidity, affect the time when callitrichids
begin or cease activity (C. jacchus [Suchi & Rothe, 1999]), one could expect that
these factors would also have some influence on the carrying efforts of tamarins in
indoor/outdoor enclosures. Nevertheless, abiotic factors (such as temperature) did
not appear to play a major role at the UAM colony. No differences were found in
body weight losses between the spring breeding periods and the autumn-winter
ones, despite the differences in temperature. In fact, outside temperatures at the
UAM colony are far to be extreme, and animals can choose to go outside or remain
indoors. In any case, all of these abiotic factors should have relatively little
influence as long as nutritional resources are available [Suchi & Rothe, 1999].
A slight but not significant difference was found in the weight of mothers in
the UAM and DPZ enclosures. Mothers tended to gain weight in small indoor
enclosures, thereby improving their physical condition (Fig. 1) [Sánchez et al.,
1999]. Mothers’ weight gain after birth has been related to low participation in
carrying, and an increased food intake [Kirkwood & Underwood, 1984; Nievergelt
& Martin, 1999; Price, 1992b; Sánchez et al., 1999], mainly when they can become
pregnant again in weeks 4–5 after birth [Sánchez et al., 1999]. However, the
lactating mothers in the large indoor/outdoor enclosures at UAM did not gain
body weight as they did at DPZ.
284 / Sánchez et al.
In conclusion, this study supports the idea that in a more naturalistic
environment, carriers lose more weight than they do under the restrictive
conditions of captivity, where most studies have been conducted. Although we
still do not know the meaning of body weight loss in terms of its value as a
selective force [Sánchez et al., 1999; Tardif, 1997], studies under semi-free
conditions, including small groups, should provide data emphasizing the
importance of cooperation for successfully coping with the high energy demands
of infant carrying in callitrichids.
ACKNOWLEDGMENTS
We thank the primate breeding and husbandry staff of the DPZ and UAM for
their technical assistance, and the anonymous referees for their very valuable
comments and advice on the early version of this manuscript.
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