THE SoUTHWESTERN NATURALIST 44(2):199-204 jUNE 1999

VARIATION IN HERD SIZE OF COLLARED PECCARIES IN A MEXICAN

TROPICAL FOREST

SALVADOR MANDUjANO

Departamentode Ecologíay Comportamiento

Animal, Instituto deEcologíaA. C., APartadoPostal 63, Xalapa,
CP 91000, Veraauz,México.email: mandujan@ecologia.edu.mx

ABSTRAcr-Collared peccaries (Pecan tajacu sonorensis)inhabit the tropical deciduous and semi-
deciduous forests of Charnela Biological Station on the coast of Jalisco, located in the southern-
most pan of the geographical distribution of this subspecies. These vegetative communities are
difIerent in fIoristic composition, phenology, aerial biomass, productivity, nutritive value, and root
biomass. From 1989 to 1994, I analyzed the variation of herd size of collared peccaries in relation
to seasonal and spatial variation of food, cover, water, and predation risk. Nl,1mber of individuals
per herd ranged from 1 to 12, with groups of 1 to 4 individuals being the most common. In
tropical semi-deciduous forest pec{;aries usually formed large herds, but in tropical deciduous
forest they most commonly formed small herds. Number of individuals per herd was similar during
rainy and dry seasons. Collared peccaries subdivided into small groups to forage in tropical de-
ciduous forest, and aggregated into herds in semi-deciduous tropical forest. Density and herd size
of P. t. sonorensisin Charnela were more similar to those found in other tropical forests than those
in northern arid habitats. Peccaries consumed a high percentage of roots thorough the year. From
rainy to dry seasons consumption of low quality roots increased as high quality leaves-branches
decreased. During the dry season, variation in fruit production and rate of fruit fall support
difIerent herd sizes.

RESUMEN-El pecari de collar (Pecan tajacu sonorensis)habita en los bosques tropicales caduci-
folios y subperennifolios de la Estación Biológica "Chamela" en la costa de Jalisco, localizada en
la parte más sureña de la distribución geográfica de estasurespecie. Estascomunidades vegetativas
son diferentes en su composición flonstica, fenología, biomasa aérea, productividad, valor nutri-
tivo y biomasa de raíz. Se analiza la variación del tamaño de las manadas del pecari de collar en
relación a los cambios estacionales y espaciales del alimento, cobertura, agua y riesgo de depre-
dación durante el período 1989 a 1994. El número de individuos por manada varió de 1 a 12,
siendo los grupos de 1 a 4 individuos los más comunes. En el bosque tropical subeperennifolio
los pecaries formaban manadas grandes, mientras que en el caducifolio comunmente formaban
manadas pequeñas. El número de individuos por manada fue similar entre las épocas de lluvia y
la seca. El pecarí de collar se dividió en pequeños grupos para pastar en el bosque tropical
caducifolio, y se reunieron en manadas en el bosque tropical subperennifolio. En Charnela, la
abundancia y el tamaño de las manadas de P. t. sonorensisfueron más similares a lo encontrado
en otros bosques tropicales que lo reportado para zonas áridas norteñas. Los pecaries consumen
altos porcentajes de raíces durante todo el año. De la época de lluvias a la época seca, el consumo
de raíces de baja calidad se incrementó mientras que disminuyó el consumo de hojas y ramas de
alta calidad. Durante la época seca la variación en la producción de frutos y la tasa de caída de
frutos, sostienen manadas de diferente tamaño. .

The collared peccary (Pecan tajacu sonoren-
sis) has been studied extensively in the arid
zones of Arizona and New Mexico (Sowls,
1984). In contrast, there has been little re-
search on this subspecies in tropical habitats.
The collared peccary is resident of tropical for-
ests near Ch~ela in southwestern Jalisco,
México (Ceballos ~d Miranda, 1986). During
a study of white-tailed deer (Odocoileusvirgini-
anus) in fue aTea (Mandujano and Gallina,
1995a, 1995b),1 algOgathered data on fue col-
lared peccary. The objective of this study was
to analyze fue variation of herd size in relation
to seasonal and spatial variation of food, cover,
200 The Southwestern
and water in fue tropical forest located in fue
southernmost part of fue geographical distri-
bution of P. t. sonorensis(Hall, 1981).
METHOOSANO MATERIALS-This study was con-
ducted in fue Charnela Biological Station on fue
southern Pacific coast of jalisco. The station covers
3,200 ha with elevations ranging from 30 to 500 m.
Mean annual temperature and precipitation are
25°C and 748 mm (SD = 119 mm), respectively.
Eighty percent ofthe raiD falls betweenjuly and Oc-
tober, fue dry season lasts from November to june
(Bullock, 1986). Dominant vegetation is tropical de-
ciduous forest located on bilIs. Tree heights wry
from 4 to 15 m and there is a well-developed under-
story (Lott et al., 1987). The station also has semi-
deciduous forest along large streams, with trees
ranging from 10 to 25 m in height.
For each sample month, five to eight transects
were located along existing roads. Though place-
ment of transects should be random (Burnham et
al., 1980), dense understory at the Charnela sta-
tion prevented placement of long random tran-
sects. Therefore, 1 used existing dirt roads. Total
transect length varied from 6 to 11 km (X = 9.1
km) each month. Length of these transects was
proporcional to fue afea of each vegetation type,
74% crossed tropical deciduous forest, the rest
crossed tropical semi-deciduous forest. 1 walked
along transects at 1 to 2 km per hour from 0700
h to ,1200 h and 1600 h to 2000 h, two to four
times per month. AlI observations were made
from O to 30 m into the forest, perpendicular to
the transect. Fieldwork was done in the rainy sea-
son from 1989 to 1993 and the dry season from
1990 to 1994. The observation rate was defined as
the number of herds observed per kilometer of
transect covered. This rate was calculated by sea-
son using data from all years; the nonparametric
Mann-Whitney U-test was used to see differences
between seasons (Sokal and Rohlf, 1995). The dis-
tribution of the number of individuals in small
herds (1-4 peccaries per herd) or large herds (>5
peccaries per herd) by season (rainy and dry) and
vegetation type (tropical deciduous forest and
tropical semi-deciduous forest) , were compared
by Chi-square Goodness of Fit tests. The strip tran-
sect method was employed to estimate annual
density (D) of herds per km2. The formula applied
was D = ni L2w, where n is the number of herds
observed, L is the totallength of the transect, and
w is one half of the total transect width (w = 0.03
km in this study). Once herd density was obtained,
the number of individuals per km2 was estimated
by multiplying herd density by average herd size
in this afea. Results are expressed as a X :t 1 Sr.'.
Naturalist vol. 44, no. 2
RESULTS ANDDISCUSSION-Atotal of 44 herds
was observed along fue 816 km walked in Cha-
mela. Due to fue dense vegetation cover, I
could not count fue numbe~ of individuals in
10 of fue herds. In fue remaining 34, a total of
139 individuals wasseen. AlI individuals prob-
ably were not counted, because sightings typi-
cally occur at road crossings and some individ-
uals mar have already gone by. Herd observa-
tion Taleswere similar between rainy (0.071 :!:
0.04 herds per km) and dry seasons (0.078 :!:
0.06) (Mann-Whitney U-test,U= 12, P= 0.84).
The number of individuals per herd ranged
from 1 to 12, with groups of 1 to 4 individuals
being fue most common (Fig. 1). In fue trop-
ical semi-deciduous forest, peccaries usually
formed large herds (5-12 individuals per
herd), whereas in fue tropical deciduous forest
they most commonly formed small herds (1-4
individuals per herd-}(I. = 4.6, di = 1, P =
0.03). The number ofindividuals ofboth small
and large herds were similar during fue rainy
and dry seasons (}(I. = 0.002, df= 1, P = 0.66).
Independent of season, average herd size in
tropical deciduous forest was 3.3 :!: 0.6 individ-
uals per herd, and 4.5 :!: 0.6 in tropical semi-
deciduous forest. In contrast, independent of
forest type, average size of herd during fue
rainy season was 3.9 :!: 0.6, and 4.4 :!: 0.7 in
fue dry season. Density of herds per year varied
from 0.3 to 2.5 herds per km2 depending on
fue year, with an average of 1.2 :!: 0.9 herds per
km2. Average density of individuals was 4.9 :!:
1.6 peccaries per km2.
Density and herd size of P. t. sonorensisin
Charnela were more similar to those found in
other tropical forests than those in northern
arid habitats. In Peru, Kiltie and Terborgh
(1983) estimated a density of 1.2 herds per
km2; in Panarna, Eisenberg and Thorington
(1973) calculated 7 individuals per km2. In
some forests in Venezuela, Panarna and Peru,
47%, 75%, and 87% of fue herds, respectively,
were made up of 1 to 4 individuals (Kiltie and
Terborgh, 1983; Robinson and Eisenberg,
1985). In contrast, in Arizona, mean popula-
tion density was estimated as 11.8 :!: 4.4 pec-
caries per km2, average herd size fluctuated be-
tween seven and 16 animals, and maximum
herd sizevaried from 20 to 30 peccaries (Sowls,
1984). Therefore, population density and herd
size of this subspecies in fue Charnela region
june 1999 Mandujano-CoIlared peccaries in a tropical forest 201

16

14

12
>-
()
c
Q) 10
:J
O-
Q)
.::: 8
Q)
:J
-O 6
CJ)
..c
ro
4

o
1-2 3-4 5-6 7-8 9-10 11-12
number of animals / herd

FIG. 1.-Frequency of fue nurnber of individuals per herd in two types of tropical forest near Charnela,
jalisco, Mexico.

depends more on habitat features than on tax- in fue tropical deciduous forest throughout
onomic affinity. fue year. Aggregation of herbivores is expected
Th'e continuous observations and mapping to increase in habitats with substantial variation
of two particular herds suggest that subdivision in forage quality and productivity, and signifi-
of a herd into small groups occurs in Charnela. cant predation (Fryxell, 1991). Vegetative com-
This pattern has been reported in other trop- munities in tropical semi-deciduous forest and
ical forests (Castellanos, 1983; Robinson and tropical deciduous forest are different in their
Eisenberg, 1985) and brushland in Texas floristic composition (Lott et al., 1987), phe-
(Green et al., 1984), where a herd ofcollared nology (Bullock and Solis-Magallanes, 1990),
peccaries subdivided into small groups in areas aerial biomass (Martínez-Yrizar et al., 1992),
with few resources. In contrast, fue en tire herd productivity (Martínez-Yrizar et al., 1996), nu-
and sometimes aggregations of distinct herds, tritive value (Silva-Villalobos, 1996), and root
occurred at sites of abundant food (Green et biomass (Castellanos et al., 1991). During fue
al., 1984). During fue entire year, herds com- rainy season,speciesrichness in the understory
monly comprised juvenile and adult individu- is higher in tropical deciduous forest than in
als in fue tropical semi-deciduous forest;whereas tropical semi-deciduous forest, but net foliage
in fue tropical deciduous forest it was production (NFP) is similar between fuese veg-
more common to see groups of one to four etation types (S. Mandujano and S. Gallina, in
adults and juveniles foraging together. New- litt.); and fue quality (protein : fibre index) oí
born peccaries were seen only in tropical semi- some species is higher in tropical deciduous
deciduous forest during October and Novem- forest than in tropical semi-deciduous forest
ber, but there is not enough data to test if pec- (Silva-Villalobos, 1996). During fue dry season
caries prefer fue semi-deciduous forest for speciesrichness and NFP are higher in tropical
rearing young. semi-deciduous forest than in tropical decidu-
Collared peccaries form large herds in fue ous forest, and quality is low and similar in
tropical semi-deciduous forest and small ones both vegetative comunities. In addition, there
202 The SouthUle5temNaturalist
are no sources of free water during fue dry
season (Mandujano and Gallina, 1995b).
Therefore, fue potential of this tropical forest
to support different herd sizes varies spatially
(tropical deciduous forest versus tropical semi-
deciduous forest) and seasonally (rainy versus
dry). Overall, tropical semi-deciduous forest
supports larger herds of collared peccaries.
Diet of collared peccaries in Charnela con-
sisted of 46% and 50% roots, 43% and 39%
leaves-branches, and 10% and 11 % fruits dur-
ing fue rainy and dry seasons, respectively
(Martínez-Romero and Mandujano, 1995).
Particularly, monthly consumption of roots
and leaves-branches were inversely correlated
(r = -0.93, df= 4, F = 23.8, P = 0.008): more
roots were consumed in fue dry season, and
more leaves-branches in fue rainy season. Sim-
ilar patterns of consumption were reported in
other tropical deciduous forests (McCoy et al.,
1990; Olmos, 1993), wherein consumption of
low quality roots increased as high quality
leaves and fruits decreased. Olmos (1993) pro-
posed that variability in percentage of roots
and tubers in fue diet is fue result of pro-
nounced seasonality of tropical deciduous for-
esto In Charnela, availability of leaves is high
only during fue rainy season;whereas through-
outthe year, total root biomass is high (Kum-
merów et al., 1990). Despite fue great root bio-
mass, fue cost of foraging mar be high because
peccaries must search and scratch for roots at
a depth of 0-10 cm (Olmos, 1993). Foraging
for roots in small herds or groups could be a
optimal strategy because fue quantity of roots
of specific plants consumed by peccaries in a
localized area probably is not enough to sup-
port a large herd.
During fue dry season, collared peccaries
consume fruits of some species like opuntia ex-
celsa,Ficus, Brosimum alicastrum, Spondiaspur-
purea, and unidentified legumes (Martínez-
Romero and Mandujano, 1995). In arid habi-
tats and other tropical deciduous forests, den-
sity, herd size, borne range, and movements
depend on abundance of certain forage spe-
cies, mainly opuntia in arid habitats and spe-
cies of fruit trees in tropical forest (Bissonette,
1985; Robinson and Eisenberg, 1985; McCoyet
al., 1990). In Charnela I commonly observed
small groups of peccaries, feeding signs, and
tracks below opuntia and Spondiastrees. The
greatest number of fuese trees are found on
vol. 44, no. 2
hillsides in tropical deciduous forest, although
at low densities with limited fruit production
(Lott et al., 1987; Mandujano et al., 1994).
Consumption of Spondias fruits by peccaries
depends on fue foraging activity of Chachala-
cas (Ortalis poliocephala), because this bird
knocks fue fruits to fue ground, making them
available to peccaries (Mandujano and Martí-
nez-Romero, 1997). A similar association has
been observed between herds of collared pec-
caries and troops of monkeys foraging in trees
(Robinson and Eisenberg, 1985). Large herds
were observed eating fruits of Ficus and Brosi-
mum trees. Relative abundance of fuese species
of trees is high in fue tropical semi-deciduous
forest (Lott et al., 1987). Therefore, high fruit
production and high rate of fruit fall of some
tree species in the tropical- semi-deciduous for-
est could support large herds.
Predators such as jaguar (Panthera onca),
puma (Puma concolor), ocelot (Leopardus par-
dallis), coyote (Canis latrans--Ceballos and Mi-
randa, 1986) and recentIy bobcat (Lynx rufus--
López-González et al., 1998), inhabit Charnela.
Particularly, fue jaguar and puma include pec-
caries in their diet (R Nuñez and B. Miller, in
litt.). In other tropical forests, jaguar is one of
fue principal predators of peccaries (Aran da,
1994). Social organization and size of herds of
large herbivores are influenced also by preda-
tion risk (Hirth, 1977; Nelson and Mech, 1981;
Fryxell, 1995). Thus, small herds and solitary
individuals are more frequent in forested hab-
itats, whereas in oren areas, herds tend to be
larger (Mandujano and Gallina, 1996). In fue
Charnela region, fue jaguar and puma use fue
tropical semi-deciduous forest as pathways in
their daily activities (R. Nuñez, B. Miller, and
F. Lindzey, in litt.). Thus, when peccaries are
in the tropical semi-deciduous forest a partic-
ular individual' s chances of being killed by
fuese cats might be lessened by being part of
a large herd. Spatial and temporal variation in
food and cover resources, and predation risk,
seem to be fue principal factors determining
variation in herd size of collared peccaries at
Charnela.
L. E. Martínez-Romero and A. Hernández gave
me information from their field observations on
collared peccaries in the study area. 1 thank M.
Aranda, A. González-Romero, P. A. Jansen, and B.
E. Coblentz for their comments and suggestions
June 1999 Mandujano,-CoIlared peccaries in a tropical forest
regarding an early version of this rnanuscript. This
research was supported by the Charnela Biological
Station. The study was also enhanced by a sirnul-
taneous study of the white-tailed deer financed by
CONACYT (P220CCOR-892154, PO20CCOR-903703
y 0327N9107).
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Submitted9 june 1997. Accepted24 November1998.
AssociateEditor was Mari¡ D. Engstrom.