Zoo Biology 16:301307 (1997)

Age Specific Fecundity, Litter Size, and

Sex Ratio in the Chacoan Peccary
(Catagonus wagneri )
Christopher J. Yahnke,1,2* Jakob Unger,3 Bridget Lohr,2 Dennis A. Meritt,4
and Werner Heuschele2
1

Department of Biological Sciences, Northern Illinois University, DeKalb, Illinois

Center for the Reproduction of Endangered Species, Zoological Society of San Diego,
San Diego, California
3
12-1419 Rothesay Street, Winnipeg, Manitoba, R2G-1V3, Canada
4
Department of Biological Sciences, De Paul University, Chicago, Illinois
2

Ten years of data on the reproductive biology of the Chacoan peccary (Catagonus
wagneri) were analyzed to determine average litter size, sex ratio, timing of births,
and individual and age-specific fecundity. Data were obtained from a captive
herd of chacoan peccaries located in the western Paraguayan Chaco. Births peak
in the austral spring months of September, October, and November, with fewer
litters born during the dry season months of June, July, and August. The average
litter size was 2.4 with a sex ratio of 56:44 (M:F). There was no significant
difference in litter size among individual females or among females of different
ages. Finally, there was some individual variation in the age at which sows produced their first litter, but no discernible variation among sows in the average
time between litters. Zoo Biol 16:301307, 1997. 1997 Wiley-Liss, Inc.
Key words: Tagu; Chaco; captive breeding; reproductive biology

INTRODUCTION

The Chacoan peccary (Catagonus wagneri) has the most restricted range of the
three living peccaries. It is endemic to the Gran Chaco of western Paraguay, northern
Argentina, and eastern Bolivia, although its occurrence in the later country is poorly
known. The primary habitat type occupied by the Chacoan peccary is the xerophytic
thorn forest or dry Algorrobo-Palo Santo-Quebracho woodlands [Short, 1975; Mayer
and Brandt, 1982]. This habitat often has dense and spiny undergrowth making observations of this animal in the field difficult. The etymology of its local name, Tagu

*Correspondence to: Christopher J. Yahnke, Department of Biological Sciences, Northern Illinois University, Dekalb, IL 60115.
Received for publication 28 October 1996; revision accepted 7 March 1997.

1997 Wiley-Liss, Inc.

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is testimony to the ability of this animal to make paths into seemingly impenetrable
areas, whereas the local names of the other two species are more descriptive of appearance and refer to the white lower jaw of the white-lipped peccary (Tanyka-ti,
Tayassu pecari) and relative small size of the collared peccary (Cure-i, Tayassu tajacu).
In part, it was the different indigenous names which led investigators to the Chacoan
peccary in 1972 [Wetzel et al., 1975]. Only in the Gran Chaco are all three peccaries
sympatric.
Relatively little is known about the Chacoan peccary. Discovered in 1972 by
the late Ralph Wetzel, it was formerly known from the Lower and Middle Pleistocene of Argentina [Wetzel, 1977]. Recent studies by Taber [1991] and Unger [see
Handen et al., 1994] have suggested that only 4000 - 5000 individuals remain in
Paraguay, with smaller populations in Argentina and Bolivia. The work by Taber et
al. [1993; 1994] and Neris [1993] are the only data on Chacoan peccary home range,
and Mayer and Brandt [1982] studied the Chacoan peccary during three consecutive
winters providing the first reports of winter diet, reproductive biology, and behavior.
These studies represent almost all that is known about the Chacoan peccary in the wild.
The Proyecto Tagu was established in 1986 in the central Paraguayan Chaco
as a means of learning more about the reproductive biology and general behavior
of the recently discovered, yet highly endangered Chacoan peccary. This captive
breeding project is funded by the Center for the Reproduction of Endangered
Species (Zoological Society of San Diego). Details on the establishment of the
colony can be found in Byrd et al. [1988], and Benirschke et al. [1989]. A number of publications have already been reported on this colony and include notes
on neonatal mortality [Benirschke et al., 1995], methods of immobilization [Allen,
1992], feeding and social behavior [Handen and Benirschke, 1991], and general
remarks on litter sizes [Brooks, 1992; Unger, 1993]. Here we present partial results from the first 10 years of data collected on reproductive biology at the
Proyecto Tagu.
MATERIALS AND METHODS

The Proyecto Tagu is located at Fortn Toledo (S 22 21 21, W 60 20 24)

35 km west of the Mennonite town of Filadelfia in the central Chaco of Paraguay.
Currently there are four enclosures ranging in size from 0.5 - 3.0 ha. Each enclosure
contains native vegetation including natural food items such as mesquite (Prosopis
nigra, P. alba), Acacia sp., and prickly pear cactus (Opuntia sp.), and larger shade
trees such as Guayacan (Caesalpinia paraguariensis), Coronillo (Shinopsis quebracho-colorado), and Quebracho blanco (Aspidosperma quebracho-blanco). Because
there is not adequate naturally occurring food in each pen, food is supplemented
daily with mandioca, squash, pumpkin, and watermelon (the later two items available seasonally), and a balanced dry domestic pig food. Water is available in each
pen throughout the year.
Data on maternity has been collected since the first captive litter was born in
November 1988. Maternity, defined as the sow most likely to have given birth to a
litter, was determined by observing associations between new litters and females.
Occasionally, maternity could not be determined. For example, multiple litters born
to a herd on the same day did not appear to form consistent associations with a
particular female. In cases where maternity or discrete litters could not be deter-

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303

mined, the data were excluded from the analysis. Sex of newborn Chacoan peccaries
was determined at the time of first physical examination, usually between 3 and 9
months of age. Also, sex of newborn peccaries found dead in pens was recorded as
part of the necropsy exam. Occasionally, newborn individuals disappeared from pens
before their sex could be determined. These individuals were probably removed by
avian scavengers, such as caracaras and vultures, soon after death. Each animal was
given a colored ear tag and an ear notch for individual identification, and reliable
dates of birth were available for all of the peccaries, enabling us to assign the sows
to age groups.
Kruskal-Wallis nonparametric statistical analysis was used to determine differences in litter size among age groups. Since multiple litters were observed for each
female, we also used this test to determine differences in litter size and interbirth
intervals among females. Although we recorded 14 sows giving birth to at least one
litter, only the seven most productive sows were used in the later analysis. Chisquare analysis was used to test for deviations from a 1:1 sex ratio in newborns and
in newborn mortality. Chi-square analysis was also used to test for a seasonal difference in the number of newborns born at the project.

RESULTS

Chacoan peccaries are born throughout the year at the Proyecto Tagu and
were observed in every month (Fig. 1). Births peaked in the spring months of
September, October, and November, with very few individuals born during the
driest months of June, July, and August (Fig. 1). The number of individuals born
during the wet season (defined as October-March, which have greater than average rainfall) was significantly greater than the number of individuals born during the dry season (X2=5.26, 0.01<P<0.025). Based on 80 captive litters, litter
size ranged from one to four and averaged 2.4 (SD=0.8). Of this total, triplets
were most common and accounted for 43.8% of the litters, followed by twins
40.0%, singles 10.2%, and quadruplets 3.8%.
Based on 164 individuals for which sex was determined, the sex ratio at birth
was 87 males to 77 females or 53:47 (M:F). This was not significantly different from
1:1 (X2=0.61, 0.25<P<0.50). The infant mortality sex ratio, defined as those animals
which died prior to three months of age, was 48 males to 38 females or 56:44 (M:F)
which also was not significantly different from 1:1 (X2=1.16, 0.25<P<0.50). The
cumulative first year mortality rate of juveniles born at the Proyecto Tagu was 50.3%
at three months, 53.3% at six months, 54.3% at nine months, and 54.3% at 12 months.
This means that 50.3% of the peccaries born at the Project die within the first three
months of age, and that mortality decreases dramatically thereafter, such that only an
additional 4.0% die between 3 and 12 months of age.
Maternity was determined confidently for 58 litters. The age of these sows
ranged from 1.28.3 years. There was no significant different in litter size among
age groups (H=4.89, df=7, P=0.67) (Table 1) or among females (H=1.60, df=6,
P=0.95) (Table 2). Although there was no significant difference in mean litter size
among age groups, it does appear that younger females (<3 years old) generally have
smaller litters (Table 1). Finally, there was no difference in the average time between
litters among females (H=9.66, df=6, P=0.14) (Table 2).

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Fig. 1. Number of births per month of Chacoan peccaries at the Proyecto Tagu captive breeding site.
The solid black bars represent average rainfall from 19901995.

DISCUSSION

These results are the first to demonstrate a clear seasonal peak in births of
Chacoan peccaries (Fig. 1). Previous studies were short term seasonal studies [Mayer
and Brandt, 1982] or had small sample sizes [Unger, 1993]. Mayer and Brandt [1982]
observed 10 neonatal litters in the field between September and November. Further,
TABLE 1. Average litter size among age groups
Age group (yrs)
1
2
3
4
5
6
7
8
All

Mean litter size (SD)

Number of litters

2.2 (0.8)
2.1 (0.8)
2.7 (0.8)
2.4 (0.9)
2.5 (0.6)
2.8 (0.8)
2.5 (0.7)
2.5 (0.7)
2.5 (0.8)

5
13
13
11
6
6
2
2
58

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305

TABLE 2. Age at first litter, average litter size, and average time between litters for the seven
most productive females at Proyecto Tagu
Age of
female (yrs)
9.5
9.3
9.0
8.5
7.2
7.1
6.3

Age at first
litter (yrs)

Number of
litters

Mean litter
size (SD)

Mean time between

litters (SD) (days)

2.4
3.1
2.8
2.1
1.1
2.3
1.3

8
6
7
7
7
5
6

2.6 (0.5)
2.3 (0.8)
2.6 (0.5)
2.7 (0.8)
2.4 (0.8)
2.2 (1.3)
2.7 (0.5)

264.5 (69.8)
295.3 (50.8)
253.3 (70.6)
242.8 (78.2)
245.8 (58.8)
191.2 (22.2)
216.5 (56.4)

they found 11 pregnant sows among 36 sows killed by hunters between July and
September. Unger [1993] recorded 24 litters born at the Proyecto Tagu through
August, 1992. To that time the two most productive months were January (11 newborns) and November (12 newborns). These data are represented graphically in Fig.
1. Of the two most productive months in the present study, October appears to be
more consistent in terms of production with 10 or 11 newborns per year in recent
years. Conversely, 24 young were born in September, 1994 accounting for most of
the births seen in that month (Fig. 1).
Early estimates of average litter size were also based on relatively small sample
sizes. These include average litter size estimates of 2.7 (n=11), 2.5 (n=10) [Mayer
and Brandt, 1982], 2.3 (n=9) [Brooks, 1992], and 2.3 (n=24) [Unger, 1993]. Our
estimate of 2.4, based on 80 litters, falls within the range of these previous estimates.
Although the sex ratio did not differ significantly from 1:1, the higher ratio of
males to females (53:47) corroborates the larger male to female ratio (54:46) found
by Mayer and Brandt [1982], based on 28 fetuses of wild caught Chacoan peccaries.
This contrasts with the collared peccary (Tayassu tajacu) which appears to have more
females than males [Sowls, 1966; Low, 1970; Smith and Sowls, 1975]. However,
Lochmiller et al. [1984] reported a higher ratio of males to females (56:44), based on
57 young from a captive herd in Texas.
There appear to be various causes of infant mortality at the Proyecto Tagu,
and it is not known whether these reflect infant mortality in the wild. We have observed cases of infanticide, usually within the first week after birth. The most common cause of infant death at Proyecto Tagu appears to be pneumonia. In one group,
the mortality has been close to 80% over the past two years. This may explain our
pattern of high mortality in the first three months after birth. However, our 12-month
mortality rate of 54.3% mirrors the result of Day [1985] who found a 55% 12-month
mortality rate in free-ranging juvenile collared peccaries.
Our sample sizes for the oldest age classes are too small to determine whether
fecundity decreased with older age. However, Sowls [1984] noted a litter of four in a
14-year old collared peccary and stated that for most female peccaries, only death or
disease could end the ability to bear young. It is not known how long peccaries live
in the wild. Based on 48 skulls recovered from hunters, Sowls [1984] estimated that
only abut 4% of the population was over nine years in age. Our oldest female, a 10year-old, recently died from unknown causes. She was pregnant with twins and, from
the size of the fetuses, was ready to give birth.
Sows producing the majority of litters at the Proyecto Tagu were captured in

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the wild as babies and raised in captivity. These are represented by the first five
females listed in Table 2, whereas the last two were born at the project. The relatively high age at first litter in the oldest three females may be due to factors related
to acclimating to captive conditions during the time of the projects inception. At this
time there were only 7 - 10 animals, all of about the same age. By early 1996, there
were approximately 70 animals, with an age structure which mirrored that found in
wild populations [Sowls, 1984].
CONCLUSIONS

1. The Chacoan peccary shows a spring birth peak, although births have been
observed in every month.
2. The average litter size is 2.4 with a sex ratio which did not differ from 1:1.
3. The average litter size does not differ among age groups or individuals.

ACKNOWLEDGMENTS

We thank Maria and Domanik Unger for their observations of the Chacoan
peccaries, Jack Allen for logistical support and comments on earlier versions of this
manuscript, and the Center for Reproduction of Endangered Species and the Zoological Society of San Diego for financial support. Finally we thank three anonymous reviewers whose comments greatly improved this manuscript.
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