Journal of Mammalogy, 83(1):207–217, 2002

COEXISTENCE OF PECCARIES AND FERAL HOGS IN THE

BRAZILIAN PANTANAL WETLAND:
AN ECOMORPHOLOGICAL VIEW

FERNANDO L. SICURO AND LUIZ FLAMARION B. OLIVEIRA*

Seção de Mastozoologia, Departamento de Vertebrados, Museu Nacional, Universidade Federal do

Rio de Janeiro, Quinta da Boa Vista, 20.940-040, Rio de Janeiro, Rio de Janeiro, Brazil

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Niche partitioning among sympatric populations of white-lipped peccary (Tayassu pecari),
collared peccary (Pecari tajacu), and feral hog (Sus scrofa) is evaluated using an ecomor-
phological approach. Masticatory apparatus and head-elevation lever system of the 3 spe-
cies are compared by skull measures related to moment arms of associated muscles. Force-
rates based on the equation of static equilibrium are proposed as a basis of comparison
between species-specific cranial designs related to mastication and digging ability. Mor-
phofunctional data suggest that feral hogs have a powerful bite and are able to feed on
seeds of different degrees of resistance. With an optimized lever system of head elevation,
feral hogs are more efficient than peccaries at rooting. These facts, along with the capacity
of feral hogs to explore a wide range of habitats, support the view that the species acts as
a potential competitor of native peccaries.

Key words: cranial morphology, ecomorphology, feral hogs, Pecari tajacu, peccary, Tayassuidae,
Tayassu pecari, Sus scrofa

Populations of white-lipped peccary March 1993). Foraging behavior of both

(Tayassu pecari) and collared peccary (Pe-
cari tajacu) widely coexist over the Neo-
tropics. Peccaries occur from arid regions
of the southwestern United States and Mex-
ico to South America. They are widespread,
ranging from the northern and eastern trop-
ical rain forest, central savanna vegetation
(Cerrado), dry regions in northeastern Bra-
zil (Caatinga), and western Paraguayan
Chaco, to other subtropical areas of South
America (Bodmer and Sowls 1993; Corn
and Warren 1985; Eddy 1961; March 1993;
Osgood 1914). Food habits of both species
have been reported as being associated pri-
marily with fruits, hard seeds, and roots
(Barreto et al. 1997; Bodmer 1990, 1991;
Kiltie 1982; March 1993; Olmos 1993). In-
sects, annelids, and small vertebrates are
cited as a complementary diet for the 2 spe-
cies (Bodmer 1990, 1991; Kiltie 1982;
* Correspondent: melfo@terra.com.br
species includes digging to access subter-
ranean food items (Kiltie 1982; Kiltie and
Terborgh 1983).
Craniomandibular characteristics and
masticatory functional performance of pec-
caries are important in niche partitioning
when species are sympatric (Kiltie 1982,
1985). According to Kiltie (1982), the dif-
ference in force used by a species to crack
hard food items is the basis for niche dif-
ferentiation. This is also reported by Olmos
(1993) for Brazilian northeastern dry areas
(caatinga), where white-lipped peccaries are
able to forage on harder food than collared
peccary. Data presented by Barreto et al.
(1997) also suggest this dichotomy. Nev-
ertheless, collared peccary seems to be
more of a generalist, having a high capacity
to change diet according to resources of-
fered. Despite the bite force of white-lipped
peccaries, which allows them to crack food

207
208 JOURNAL OF MAMMALOGY
items of a wide range of hardness, they tend
to be more selective. White-lipped pecca-
ries prey mainly on roots and hard seeds
that are not eaten by collared peccaries
(Bodmer 1991; Kiltie 1982; Olmos 1993).
Feral morphotype of hogs (Sus scrofa)
and effects of their presence on the envi-
ronment have been widely studied (Hone
1988, 1990; Hone and Stone 1989; Mayer
and Brisbin 1991; McKnight 1976; Oliver
and Brisbin 1993). At present, feral hogs
have spread worldwide, occurring com-
monly in Argentina, Australia, Brazil,
Chile, Ecuador (including Galapagos), New
Zealand, and United States (including Ha-
waii). The diet of feral hogs is composed
of fruits, seeds, roots, grass, and small- to
medium-sized vertebrates (Ilse and Hell-
gren 1995a; Oliver and Brisbin 1993). Be-
cause of their generalist profile and a high
capacity to spread over new habitats, feral
hogs are eminent competitors of native fau-
na in those countries (Gabor and Hellgren
2000; Hone and Stone 1989; Ilse and Hell-
gren 1995b; Lacher et al. 1986; McKnight
1976; Oliver and Brisbin 1993). Notwith-
standing, Ilse and Hellgren (1995a) and Ga-
bor and Hellgren (2000) suggest that water
availability and high temperatures are con-
straining factors to the success of feral hog
populations in hot and dry environments.
The Pantanal is a 140,000-km2 alluvial
area in the upper Paraguay basin; composed
of a complex mosaic of vegetation, it is one
of the richest wetland areas in the world
(Adamoli 1981; Alho et al. 1987). Intro-
duced hogs possibly became feral in the
2nd half of the 18th century and now occur
in all regions of the Pantanal, coexisting
with native peccaries (Alho and Lacher
1991). Lacher et al. (1986) ask if such co-
existence could be the cause of the decrease
of populations of peccaries. Feral hogs are
major contributors to biomass of mammals
in the Pantanal, reflecting the ecological im-
portance of this species to the region (Lach-
er et al. 1986).
This study, through an analysis of cranial
morphology, considers the potential capac-
Vol. 83, No. 1
ity of each species to access and to use food
resources. A functional analysis of the jaw
lever system compares the performance of
the masticatory apparatus of white-lipped
peccaries, collared peccaries, and feral
hogs. The system of head elevation is also
compared with and associated with soil-dig-
ging display of foraging. The 3 species
have morphological adaptations for masti-
cating resistant food (Kiltie 1981, 1982;
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Herring 1972) and specializations related to
digging, such as a robust rhinarium (Ewer
1958; Herring 1972) and obvious hypertro-
phy of the nuchal muscles (Herring 1972).
An ecomorphological analysis was per-
formed to assess the possibility of niche
overlap based on food resources. Scientific
names follow Grubb (1993).
MATERIALS AND METHODS
Skulls of collared peccaries (n 5 99), white-
lipped peccaries (n 5 80; Figs. 1A and 1B), and
feral hogs (n 5 27; Fig. 1C), including speci-
mens of several regions of Brazil, were studied
in the following Brazilian collections: the Museu
Nacional (MN), Rio de Janeiro; the Museu de
Zoologia of the University of São Paulo
(MZUSP), São Paulo; the collection of the Cen-
tro de Pesquisa Agropecuária do Pantanal (Em-
presa Brasileira de Pesquisa Agropecuária)
(MML), Corumbá; and the Herrera private col-
lection (CH) in Corumbá, Mato Grosso do Sul.
All individuals studied were adults or subadults
according to the age classification of Mayer and
Brandt (1982) for peccaries and Mayer and Bris-
bin (1991) for hogs. A list of specimens is avail-
able from the authors.
Hypotheses presented here are based primar-
ily on field activities of collecting and observing
free-ranging peccaries and feral hogs along the
Pantanal wetland (subregion of Nhecolandia,
Brazil, 198049S, 568369W) during the dry season
(July–August) of 1995.
Seventeen skull measures described below
were selected to reflect significant functional as-
pects of cranial morphology. These measures
were used to evaluate the performance of prin-
cipal masticatory and head-elevating muscle
groups (Fig. 2), according to species-specific
cranial morphology (Figs. 3A, 3B, and 4). Two
of these measures, width of complex muscle scar
February 2002 SICURO AND OLIVEIRA—PECCARIES AND FERAL HOGS
FIG. 1.—Skulls of peccaries and feral hogs. A)
Pecari tajacu (MN 37014); B) Tayassu pecari
(MN 42908); and C) Sus scrofa in its feral mor-
photype (MN 42886). This male S. scrofa has
the typical cranial pattern of feral hogs found in
Pantanal Mato-Grossense region of Brazil.
(CSW) and width of lambdoidal crest (LCW)
were taken as an indicator of the transverse sec-
tion area of the complex muscle in the occipital
region (Fig. 4).
The following 17 measurements, based par-
tially on Radinsky (1981a, 1981b) and Kiltie
(1982, 1984, 1989), were used to evaluate skull
morphology relative to functional implications:
Basal condyle length (BCL), measured from
209
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FIG. 2.—Muscle groups analyzed. Open arrow
indicates action of masseter group; solid arrow
indicates action of the temporal group; and the
hatched arrow indicates action of the complex
group.
posterior end of occipital condyles to anterior
end of premaxilla, estimates resistance moment
arm when an animal is digging the soil. Condyle
to canine length of jaw (CCL), measured from
posterior end of condyles to posterior end of ca-
nine alveolus, measures resistance moment arm
when an animal is biting with canine teeth. Con-
dyle to M1 length of jaw (CM1L), measured from
posterior end of condyles to anterior end of M1
crown, measures the resistance moment arm
when an animal is biting with the teeth. Com-
plex muscle scar width (CSW), measured at the
widest point of complex muscle scar on occipi-
tal, estimates size of this muscle. Jaw height at
M1 (JHM1), measured about mid point of den-
tary between M1 and P4, estimates the 2nd mo-
ment of area of jaw (SMA). Jaw width at M1
(JWM1), measured near point of JHM1, esti-
mates cross-section of dentary bone. Lambdoi-
dal crest width (LCW), measured at widest point
of lambdoidal crest in occipital, estimates size
of complex muscle. Masseter muscle moment
arm (MMA), measured from dorsal surface of
condyle to ventral border of angular process, es-
timates moment arm of this muscle. Masseter
muscle scar length (MSL), measured in ventral
face of zygomatic arch, from anterior limit of
muscle scar in jugal to anterior end of glenoid
fossa (temporomandibular fossa). Masseter mus-
cle scar width (MSW), widest point of masseter
scar in jugal bone, an estimator, with anterior
measurement, of area of cross-section of the
masseter. Occipital region height (ORH), mea-
sured from ventral border of foramen magnus to
lowest limit of middle of complex muscle scar,
210 JOURNAL OF MAMMALOGY
FIG. 3.—Diagram showing 15 measurements
of peccaries and feral hogs skulls defined in text.
A) Frontolateral view of a skull of a peccary. B)
Ventral view of skulls of a peccary (left) and
feral hog (right) with cranial measures. Masse-
teric scar width (MSW) in feral hog was calcu-
lated as mean of measures MSW (from the an-
terior region of the zygomatic arch, where the
masseteric scar is well marked) and MSW9 (at
the widest region of the zygomatic arch). This
drawing shows general differences of skulls of
the 2 families. C) Lateral view of a mandible of
a peccary.
Vol. 83, No. 1
FIG. 4.—Posterior view of skulls of A) a pec-
cary and B) a feral hog with 2 cranial measures
depicting general differences between 2 fami-
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lies. Complex muscle scar width (CSW), mea-
sured at the widest point of complex muscle scar
in the occipital (a–a9), is an estimator of the size
of this muscle. The lambdoidal crest width
(LCW), measured at the widest point of the
lambdoidal crest in the occipital (b–b9), esti-
mates size of the complex muscle.
estimates moment arm of this muscle. Posttem-
poral fossa width (PTW), narrowest posterior
width across parietals, estimates temporal mus-
cle width (TMW). Rostral width at the 2nd pre-
molar P2 (RWP2), defined as width between ex-
ternal limits of maxillary bones about P2, indi-
cates robustness of rostrum. Skull length (SL),
measured from posterior end of the middle point
of lambdoidal crest to anterior end of premax-
illa, estimates body size of individual. Temporal
fossa length (TFL), measured from most poste-
rior point of temporal fossa to supraorbital pro-
cess, estimator of temporal muscle size. Tem-
poral muscle moment arm (TMA), measured
from posterior end of condyle to apex of coro-
noid process, estimates moment arm of temporal
muscle. Zygomatic arches internal width (ZIW),
widest point across zygomatic arches taken from
its inner faces, estimates temporal muscle width
(TMW).
The use of fresh muscle samples or recon-
structions based on osteological analysis to es-
timate the contraction force of a muscle is a sim-
plification of the muscular mechanic system
(Bryant and Russell 1995; Bryant and Seymour
1990; Hildebrand 1988). Nevertheless, this
method is very useful when employed in mor-
phofunctional comparisons between morpholog-
ically or phylogenetically related individuals
(Bryant and Russell 1995; Bryant and Seymour
1990; Eisenberg 1985; Hildebrand 1988; Kiltie
1982, 1984, 1989; Radinsky 1981a, 1981b).
Muscle scars and dimensions of fossae measured
February 2002 SICURO AND OLIVEIRA—PECCARIES AND FERAL HOGS
on the skull were associated with original mus-
cle volume or cross-sectional area (e.g., length
and width of the zygomatic scar of the masse-
ter). The distances between the mandibular con-
dyle and point of insertion of the temporal mus-
cle (coronoid process of the jaw) and deep and
superficial masseters (angular process of the
jaw) were associated with the in-force moment
arm of those muscles (Fig. 3C). The out-force
moment arm of the jaw is defined as distance of
the mandibular condyle to a bite point in the
tooth row (e.g., canines, M1). This assumption
is acceptable for bidimensional models of orthal
movement. The lever-system model of head el-
evation follows the same criterion: the in-force
lever arm is the distance from the occipital con-
dyle to origin scar of the complex muscle in the
occipital bone. The distance from the occipital
condyle to the premaxilla, which defines the out-
force moment arm, presupposes that the resis-
tance point, during the digging movement, is the
rhinarium.
The pterygoid muscle was not included in the
model, as the origin of this muscle is not clearly
seen on the ventral surface of skulls of the 3
species. The way of action and mass of this mus-
cle are very similar to the masseter (Herring
1972; Turnbull 1970). Thus, pterygoid muscle,
with the masseter are treated as a single func-
tional unit.
The equation of static equilibrium (sum of
moments equal to zero) was used to predict the
theoretical maximum force performed by jaws
and nuchal muscle of the animals. The force-rate
presumes the resistance torque of a food item
between tooth rows or of soil against the rhi-
narium as the limit that the animal can crush or
displace. In force-rate equations, linearized area
replaces in-force magnitude or volume estimated
for a muscle; therefore, force-rate is an index of
the maximum force that an individual can exert
according to its specific cranial morphology. The
force-rate general equation is FR 5 LS or LV 3
L1/L2, where LS or LV are the linearized (1/2
or 1/3 powers) muscle scar area (S) or volume
(V), L1 is the muscle moment arm, and L2 is
the resistance moment arm. Force-rates of the
masticatory muscles are corrected by the SMA
about the dentary bone. SMA is an estimator of
resistance of the dentary to the bending forces
that act in this bone during the bite. According
to Alexander (1988), no biological structure is
harder than the load it needs to bear. Thus, there
211
is a correlation between variation in the robust-
ness of the jawbone of the species and loads
involved in mastication. The corrected force-
rates (CFR) are derived variables that bring
more detail to the bidimensional model.
The following equations based on criteria
used by Kiltie (1981) and Radinsky (1981a,
1981b) were calculated, using variables de-
scribed above: force-rate of complex muscle
(FCM), ([LCW 3 CSW]0.5 3 ORH)/BCL.
Force-rate of masseter muscle at canines (FMC),
([MSL 3 MSW 3 MMA]1/3 3 MMA)/CCL.
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Force-rate of masseter muscle at M1 (FMM1),
([MSL 3 MSW 3 MMA]1/3 3 MMA)/CM1L.
Temporal muscle width (TMW), (ZIW 2 PTW)/
2. Force-rate of temporal muscle at canines
(FTC), ([TFL 3 TMW]0.5 3 TMA)/CCL. Force-
rate of temporal muscle at M1 (FTM1), ([TFL 3
TMW]0.5 3 TMA)/CM1L. Force-rate of temporal
and masseter muscles at M1 (FTMM1), (FMM1
3 FTM1)0.5. Second moment of area of dentary
bone at M1 (SMA), ((p 3 [JWM1/2] 3 [JWM1/
2]3)/4)0.25. Corrected force-rate of masseter mus-
cle at M1 (CFMM1), (SMA 3 FMM1)0.5. Cor-
rected force-rate of temporal muscle at M1
(CFTM1), (SMA 3 FTM1)0.5. Corrected force-
rate of temporal and masseter muscles at M1
(CFTMM1), (SMA 3 FTMM1)0.5.
The real values of skull measures and the
force-rates of the 3 species were compared by
means of analysis of variance (ANOVA). Force-
rates were examined through factor analysis
(principal component extraction method), and
functional performance of each species was
compared through the analysis of variance of the
factor scores of the individuals. All statistical
analysis was performed using the program Sta-
tistica (StatSoft 1999). Results of the analysis
were related to field observations in Pantanal
and to some ecological aspects described in the
literature on the 3 species.
RESULTS
Morphological analysis.—Divergences in
cranial morphology of suids and peccaries
are broadly stressed in the literature. The
main characteristics of suids are Turnbull’s
ungulate-grinding cranial-mandibular pat-
tern (Turnbull 1970) with a high glenoid
cavity and consequently higher position of
the jaw condyle (well above the toothrow
line); more specialized molar teeth, and a
212 JOURNAL OF MAMMALOGY Vol. 83, No. 1

TABLE 1.—Means and morphological differences in skulls of peccaries (Tayassu pecari, Pecari
tajacu) and feral hogs (Sus scrofa). Differences between species were significant in all cases (P ,
0.001). Differences between groups were significant (according to Tukey HSD test for unequal n)
except for 3 cases (see text).

Mean (mm) ANOVA

Skull
variable P. tajacu T. pecari S. scrofa F d.f.
BCL 198.42 242.84 284.35 576.56 2, 185
CCL 130.22 162.17 208.18 702.82 2, 185
CM1L 80.79 99.92 149.05 962.59 2, 185
CSW 7.61 10.50 22.27 350.15 2, 184

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JHM1 33.51 39.65 46.08 153.25 2, 185
JWM1 11.78 16.32 24.71 619.66 2, 185
LCW 33.40 42.44 66.62 438.27 2, 181
MMA 67.72 87.34 123.44 741.83 2, 185
MSL 55.21 65.07 90.08 529.93 2, 195
MSW 11.34 14.49 10.72 98.34 2, 195
ORH 72.73 85.54 103.74 290.36 2, 185
PTW 23.66 32.28 52.16 735.25 2, 195
RWP2 34.66 53.90 51.93 928.25 2, 194
SL 228.53 271.39 313.70 467.50 2, 195
TFL 78.82 90.51 86.71 136.67 2, 195
TMA 28.74 35.45 41.91 224.50 2, 184
ZIW 96.04 109.03 119.18 185.65 2, 195

masseter muscle with a high degree of pen-
nation. In peccaries, the leading features are
the hinge-like jaw joint with pre- and post-
glenoid processes; interlocking canines that
constrain lateral movement of the jaw; and
FIG. 5.—Distribution of the specimens accord-
ing to scores for each factor in multivariate anal-
ysis. Ellipses depict 95% confidence limits for
the species centroids based on bivariate means.
Arrows indicate the relationships of the vari-
ables with axes. Factor 1 indicates masticatory
performance and size of individuals. Factor 2
characterizes mechanical optimization of the
temporal system and superiority of the system
of head elevation. Acronyms defined in text.
enamel reinforcement in molar teeth
(Greaves 1978, 1980; Herring 1972, 1985;
Janis 1995; Kiltie 1981, 1989).
Almost all skull measures distinguished
the 3 species (Table 1). Skulls of collared
peccaries are significantly smaller than
those of white-lipped peccaries in all 17
measures (P , 0.0001). Tukey’s P was
,0.001 for comparisons of each species
with each other, for each skull variable,
with 3 exceptions. The exceptions: collared
peccaries did not differ significantly from
feral hogs in masseter muscle scar width
(MSW—P 5 0.45), white-lipped peccaries
did not differ significantly from feral hogs
in rostral width at P2 (RWP2—P 5 0.11),
and white-lipped peccaries and feral hogs
differed in length of temporal fossa (TFL)
only at the level of P 5 0.03. Both sexes
of feral hogs have a large skull with a ro-
bust aspect. Nevertheless, white-lipped pec-
caries also have a powerful skull, whereas
the skulls of both peccaries seem to be
more compact than that of hogs.
Morphofunctional multivariate analy-
sis.—Factor scores (Fig. 5) indicate contri-
February 2002 SICURO AND OLIVEIRA—PECCARIES AND FERAL HOGS 213

and 2nd factors. P values are .0.0001 in all cases. Significant differences between groups (according to Tukey HSD test for unequal n) are
TABLE 3. Results of ANOVA of 2 species of peccary (Pecari tajacu and Tayassu pecari) and feral hogs (Sus scrofa) based on scores of 1st
TABLE 2.—Loadings of each original force

T. pecari 3 S. scrofa
rate related to the first 2 axes of the factor anal-
ysis. Loads higher then 0.650 are indicated by an

,0.0001*
asterisk.

0.99
Variable Factor 1 Factor 2
FTM1 0.596 20.795*
FTC 0.686* 20.670*
FMM1 0.924* 0.280
FMC 0.862* 0.464

P. tajacu 3 S. scrofa
FTMM1 0.945* 20.258
FCM 0.569 0.696*

,0.0001*
,0.0001*

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Tukey P
CFTM1 0.932* 20.288
CFMM1 0.932* 0.341
CFTMM1 0.985* 0.075
Eigenvalue 6.346 2.128
Total variation (%) 70.51% 23.65%

P. tajacu 3 T. pecari
butions of each individual to the axis in fac-

,0.0001*
tor analysis. Loading of force-rates indi-

0.36
cates relative association of each variable
with factors (Table 2). Factor 1 can be con-
sidered as the total masticatory perfor-
mance of the species according to specific
muscular mechanical systems and size of
individuals. Factor 2 characterizes mechan-

2, 154
2, 154
ical optimization of the temporal system
d.f.
ANOVA
(FTM1, FTC) of peccaries and superiority
of the system of head elevation (FCM) of 108.62
feral hogs that is related to ability to root. 127.88
F

According to factor 1, masticatory perfor-

mance of white-lipped peccaries and feral
hogs is similar (Table 3), although the 2
S. scrofa

species have different morphofunctional

0.717
2.855

patterns. Potential bite force of collared

peccaries seems to be less powerful than
that of the other 2 species. Difference in
Mean score

potential bite force of collared peccaries

T. pecari
0.751
20.136

and white-lipped peccaries was also ob-

served by Kiltie (1982), who associated it
with niche differentiation between species.
Our analysis suggests that bite force achiev-
indicated by an asterisk.

P. tajacu
20.778
20.280

able by feral hogs is at least equivalent to

that of white-lipped peccaries and superior
to that of collared peccaries. This functional
modeling is not sensitive to differences in
specific muscle architecture such as the de-
Factors

gree of pennation. Notwithstanding, the

1
2

highly pennated masseter of suids (Herring

1972) may increase the final bite force of
214 JOURNAL OF MAMMALOGY Vol. 83, No. 1

TABLE 4.—Results of ANOVA for force-rates comparisons between peccaries and feral hogs. P values are .0.0001 in all cases. Significant
feral hogs or give them fine control of the

T. pecari 3 S. scrofa
jaw during lateral mastication (Gans and

,0.0001*
,0.0001*

,0.0001*
Bock 1965; Herring 1972; Hildebrand

,0.001*

,0.001*
,0.001*
0.27

0.10
0.87
1988). Whatever hypothesis is adopted, it

differences between groups (according to Tukey HSD test for unequal n) are indicated by an asterisk. Force-rates are described in text.
does not change substantially the essence of
the results: feral hogs are able to display a
bite force comparable or superior to that of
white-lipped peccaries and stronger than
that of collared peccaries.

P. tajacu 3 S. scrofa
Factor 2 emphasizes the dichotomy be-

,0.0001*
,0.0001*
,0.0001*

,0.0001*

,0.0001*

,0.0001*

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tween the mechanical systems of the tem-

Tukey P

0.15

0.46

0.03
poral and the complex muscles (Fig. 5; Ta-
ble 3). Enhancement of the temporal system
in peccaries is related to some morpholog-
ical skull features rather than to size of the
specimen. Although peccaries are smaller
than feral hogs, the coronoid process in the

P. tajacu 3 T. pecari
former looks more robust than in suids. The
anatomical space filled by this muscle in

,0.0001*
,0.0001*
,0.0001*
,0.0001*
,0.0001*
,0.0001*
,0.0001*
,0.0001*
,0.0001*
peccaries skulls is bigger than in feral hogs.
Conversely, despite the long snout (i.e.,
a long resistance moment arm), the FCM of
Pantanal feral hogs is higher than that of
peccaries. This improvement in the head-
elevation system is directly related to per-
172
178
178
177
177
177
178
177
177
formance in soil digging. In fact, rooting
d.f.
2,
2,
2,
2,
2,
2,
2,
2,
2,
areas of feral hogs measured in the field
ANOVA

(mean depth 5 248.0 mm 6 100 SD, n 5

469.69
222.15
180.24
26.39
53.37
105.47
253.01
83.37
166.49
10) are substantially deeper than those
F

made by the peccaries (mean depth 5 77.2

6 21.4 mm, n 5 18; P , 0.0001). The area
of soil excavation by peccaries is limited to
S. scrofa

about 1 m2, whereas excavation by feral

14.02
28.90
40.24
10.76
14.98
24.51
26.63
16.25
20.80

hogs extends from 2 to .50 m2.

Univariate analysis of force-rates.—The
3 species differed in most of the original
values of force-rate (Table 4). The similar-
T. pecari
Mean

7.41
23.45
38.06
12.89
20.93
28.19
23.83
17.67
20.51

ity observed between collared peccaries and

hogs at the FTMM1 (5[FMM1 3 FTM1]0.5,
not corrected by the SMA) is associated
with superior performance of temporal
P. tajacu

muscle system of collared peccaries and

5.79
18.03
29.06
11.75
18.94
23.41
18.76
15.15
16.85

with values of masseter force-rates of feral

hogs. The equivalence between total bite
force based on corrected force-rate of tem-
Force-rate

poral and masseter muscle at M 1

CFTMM1

(CFTMM1) of white-lipped peccaries and

CFMM1
FTMM1

CFTM1
FMM1

FTM1
FCM
FMC

feral hogs as seen with multivariate analysis

FTC

is confirmed by analysis of variance results

February 2002 SICURO AND OLIVEIRA—PECCARIES AND FERAL HOGS
of force-rates. There is no significant dif-
ference in total bite force between the 2
species. Despite the morphofunctional spe-
cialization of masseter mechanics and the
great size of hogs skull, mean values of
FMM1 of white-lipped peccaries and feral
hogs are comparable.
DISCUSSION
Ecological implications of morphology.—
Omnivory and tolerance to environmental
variation are the most conspicuous ecolog-
ical characteristics of the 3 species. Al-
though there are other foragers on seeds and
fruit in the neotropics, Kiltie (1982) pointed
out that the 2 peccary species are the most
direct competitors of one another. Kiltie
suggested that the specific adaptation for
consuming food items of different hardness
is an evolutionary response that reduces
competition. This proposition is reasonable
for species with a long history of coexis-
tence. Because of the powerful bite force,
white-lipped peccaries can access a wide
range of food items, including those in the
fundamental niche of collared peccaries.
The more selective white-lipped peccaries
forage mainly on hard seeds and roots,
whereas collared peccaries have a diversi-
fied diet of softer food items, minimizing
overlap of the realized niche of each species
and thus reducing possibilities for compe-
tition.
Hogs, in their feral morphotype, are rec-
ognized as a pest in many countries (Hone
and Stone 1989; Mayer and Brisbin 1991;
McKnight 1976). Relations between feral
hogs and native fauna are not clear in most
cases. According to Oliver and Brisbin
(1993), feral hogs are typically generalists
and effective foragers. Our analysis sug-
gests that feral hogs are able to bite with a
final force comparable to that of white-
lipped peccaries. They are, therefore, able
to crack hard food items as well as exploit
a broad-spectrum diet. As a result, the fun-
damental niche of feral hogs can overlap
the fundamental niche of peccaries. Indeed,
similarities were found in the food items
215
identified in the stomachs of individuals of
the 3 species (Herrera 1995) in the Pantan-
al. Moreover, access to underground food
items (i.e., roots, annelids, mollusks, and
crustaceans) is related to an efficient soil-
digging performance of the 3 species. Mor-
phofunctional analysis of the complex mus-
cle suggests an optimized pattern in the
head elevation system of feral hogs. Di-
mensions of feral hogs excavations indicate
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they perform better, accessing resources in
wider areas and deeper strata than pecca-
ries.
If 2 or more species share similar eco-
logical resources but niche overlap is very
small, or if resources are abundant, then
species can coexist in essentially separate
and almost identical fundamental niches. If
niches overlap to a greater extent, and
available resources in the overlap zone can-
not meet demand, then the less-efficient
species will be subject to competitive ex-
clusion (Giller 1984). Therefore, coexis-
tence of such species is a function of equi-
librium between supply and demand of re-
sources. Ilse and Hellgren (1995a) found
niche overlap between collared peccaries
and feral hogs in southern Texas to be lower
than they expected. They also hypothesize
that low availability of free water in that
region gives collared peccaries some advan-
tage over feral hogs. Notwithstanding, Ilse
and Hellgren (1995a) document dietary
overlaps between those species in winter
and spring, overlap of activity patterns dur-
ing spring and especially summer, and high
habitat overlap in winter. In the Pantanal,
we observed feral hogs and white-lipped
peccaries foraging together near water bod-
ies in the dry period.
The Pantanal Mato-Grossense is a region
of great abundance and diversity of ecolog-
ical resources. This environmental condi-
tion may reduce the potential impact caused
by presence of feral hogs. However, the re-
gion is stressed by seasonal flooding and
dry periods, which may reduce food supply.
In such conditions or in areas where re-
sources are limited, the process of compet-
216 JOURNAL OF MAMMALOGY
itive exclusion may be intensified by the
overlap of realized niches. In this case, by
interference or by exploitation of resources,
the peccary populations will be competi-
tively inferior and in a position to be po-
tentially excluded from resources.
RESUMO
A partição de nichos de populações sim-
pátricas de queixadas (Tayassu pecari), cai-
titus (Pecari tajacu) e porcos-monteiros
(Sus scrofa) é avaliada através de um en-
foque ecomorfológico. Índices de força, ba-
seados na equação de equilı́brio estático,
são propostos para inferir o momento de
força dos músculos associado à mastigação
e à elevação da cabeça. Esses ı́ndices, ger-
ados a partir de medidas cranianas relacion-
adas aos complexos biomecânicos ósteo-
musculares, comparam o design craniano
das espécies. As análises sugerem que os
porcos-monteiros possuem uma maior força
de oculasão mandibular, que permite a estes
incluı́rem itens alimentares com diferentes
graus de dureza na sua dieta. Devido a um
sistema otimizado para a elevação da ca-
beça, os porcos ferais são mais eficientes do
que os taiassuı́deos na atividade de revolver
o solo durante o forrageamento. Este fato,
associado à sua capacidade de explorar uma
variada gama de hábitats, dá suporte à hi-
pótese de que a espécie possa atuar como
um potencial competidor com os taiassuı́-
deos.
ACKNOWLEDGMENTS
We thank the Empresa Brasileira de Pesquisa
Agropecuária (EMBRAPA) for cooperation at
the research farm Nhumirim in Pantanal Mato-
Grossense during field work. We are grateful to
R. Herrera and H. Herrera for sharing their field
experience with peccaries and feral hogs. T.
Lacher, M. E. Araujo de Oliveira, and M. Velho
provided helpful revisions of the manuscript.
Conselho Nacional de Desenvolvimento Cientı́-
fico e Tecnológico (CNPq) and Coordenadoria
de Apoio de Pessoal de Ensino Superior
(CAPES) afforded financial support.
Vol. 83, No. 1
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