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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- México (Ceballos ~d Miranda, 1986). During
sis) has been studied extensively in the arid a study of white-tailed deer (Odocoileusvirgini-
zones of Arizona and New Mexico (Sowls, anus) in fue aTea (Mandujano and Gallina,
1984). In contrast, there has been little re- 1995a, 1995b),1 algOgathered data on fue col-
search on this subspecies in tropical habitats. lared peccary. The objective of this study was
The collared peccary is resident of tropical for- to analyze fue variation of herd size in relation
ests near Ch~ela in southwestern Jalisco, to seasonal and spatial variation of food, cover,
200 The Southwestern
Naturalist vol. 44, no. 2

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

are no sources of free water during fue dry hillsides in tropical deciduous forest, although
season (Mandujano and Gallina, 1995b). at low densities with limited fruit production
Therefore, fue potential of this tropical forest (Lott et al., 1987; Mandujano et al., 1994).
to support different herd sizes varies spatially Consumption of Spondias fruits by peccaries
(tropical deciduous forest versus tropical semi- depends on fue foraging activity of Chachala-
deciduous forest) and seasonally (rainy versus cas (Ortalis poliocephala), because this bird
dry). Overall, tropical semi-deciduous forest knocks fue fruits to fue ground, making them
supports larger herds of collared peccaries. available to peccaries (Mandujano and Martí-
Diet of collared peccaries in Charnela con- nez-Romero, 1997). A similar association has
sisted of 46% and 50% roots, 43% and 39% been observed between herds of collared pec-
leaves-branches, and 10% and 11 % fruits dur- caries and troops of monkeys foraging in trees
ing fue rainy and dry seasons, respectively (Robinson and Eisenberg, 1985). Large herds
(Martínez-Romero and Mandujano, 1995). were observed eating fruits of Ficus and Brosi-
Particularly, monthly consumption of roots mum trees. Relative abundance of fuese species
and leaves-branches were inversely correlated of trees is high in fue tropical semi-deciduous
(r = -0.93, df= 4, F = 23.8, P = 0.008): more forest (Lott et al., 1987). Therefore, high fruit
roots were consumed in fue dry season, and production and high rate of fruit fall of some
more leaves-branches in fue rainy season. Sim- tree species in the tropical- semi-deciduous for-
ilar patterns of consumption were reported in est could support large herds.
other tropical deciduous forests (McCoy et al., Predators such as jaguar (Panthera onca),
1990; Olmos, 1993), wherein consumption of puma (Puma concolor), ocelot (Leopardus par-
low quality roots increased as high quality dallis), coyote (Canis latrans--Ceballos and Mi-
leaves and fruits decreased. Olmos (1993) pro- randa, 1986) and recentIy bobcat (Lynx rufus--
posed that variability in percentage of roots López-González et al., 1998), inhabit Charnela.
and tubers in fue diet is fue result of pro- Particularly, fue jaguar and puma include pec-
nounced seasonality of tropical deciduous for- caries in their diet (R Nuñez and B. Miller, in
esto In Charnela, availability of leaves is high litt.). In other tropical forests, jaguar is one of
only during fue rainy season;whereas through- fue principal predators of peccaries (Aran da,
outthe year, total root biomass is high (Kum- 1994). Social organization and size of herds of
merów et al., 1990). Despite fue great root bio- large herbivores are influenced also by preda-
mass, fue cost of foraging mar be high because tion risk (Hirth, 1977; Nelson and Mech, 1981;
peccaries must search and scratch for roots at Fryxell, 1995). Thus, small herds and solitary
a depth of 0-10 cm (Olmos, 1993). Foraging individuals are more frequent in forested hab-
for roots in small herds or groups could be a itats, whereas in oren areas, herds tend to be
optimal strategy because fue quantity of roots larger (Mandujano and Gallina, 1996). In fue
of specific plants consumed by peccaries in a Charnela region, fue jaguar and puma use fue
localized area probably is not enough to sup- tropical semi-deciduous forest as pathways in
port a large herd. their daily activities (R. Nuñez, B. Miller, and
During fue dry season, collared peccaries F. Lindzey, in litt.). Thus, when peccaries are
consume fruits of some species like opuntia ex- in the tropical semi-deciduous forest a partic-
celsa,Ficus, Brosimum alicastrum, Spondiaspur- ular individual' s chances of being killed by
purea, and unidentified legumes (Martínez- fuese cats might be lessened by being part of
Romero and Mandujano, 1995). In arid habi- a large herd. Spatial and temporal variation in
tats and other tropical deciduous forests, den- food and cover resources, and predation risk,
sity, herd size, borne range, and movements seem to be fue principal factors determining
depend on abundance of certain forage spe- variation in herd size of collared peccaries at
cies, mainly opuntia in arid habitats and spe- Charnela.
cies of fruit trees in tropical forest (Bissonette,
1985; Robinson and Eisenberg, 1985; McCoyet L. E. Martínez-Romero and A. Hernández gave
al., 1990). In Charnela I commonly observed me information from their field observations on
small groups of peccaries, feeding signs, and collared peccaries in the study area. 1 thank M.
tracks below opuntia and Spondiastrees. The Aranda, A. González-Romero, P. A. Jansen, and B.
greatest number of fuese trees are found on E. Coblentz for their comments and suggestions
June 1999 Mandujano,-CoIlared peccaries in a tropical forest 203

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