BIOTROPICA 39(4): 519–524 2007 10.1111/j.1744-7429.2007.00305.

Carrying Capacity and Potential Production of Ungulates for Human Use in a Mexican
Tropical Dry Forest

Salvador Mandujano1
Departamento de Biodiversidad y Ecologı́a Animal, Instituto de Ecologı́a A. C., km 2.5 Camino Antiguo a Coatepec No. 351,
Xalapa 91070, Ver., México

ABSTRACT
Data are provided on the carrying capacity and potential production for sustainable human use of white-tailed deer (Odocoileus virginianus) and collared peccary
(Pecari tajacu) in a protected tropical dry forest at Chamela on the Pacific coast of Mexico. In this paper, the carrying capacity was defined as the equilibrium density
plus the number of animals removed by predators. The equilibrium point was estimated from the density dependent relationship between the finite population growth
rate and the current density according to a logistic model. Annual density was estimated using the line transect method. Carrying capacity estimates were 16.5 to
17.2 deer/km2 and 9.3–9.5 peccaries/km2 , representing a combined biomass of 841–874 kg/km2 . A potential production for human use of 2.1 deer/km2 and 4.4
peccaries/km2 was estimated employing the model of Robinson and Redford (1991). The data suggest that, in the protected tropical dry forest of Chamela, the density
and biomass of wild ungulates can maintain a similar or greater density and biomass than other Neotropical forests. To obtain an accurate estimation of the maximum
sustainable yield (MSY), it is necessary to consider predation. From a management point of view, it is important to consider that carrying capacity varies as a function
of the rainfall pattern.

Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp.

Key words: density; environment variation; Mexico; Odocoileus virginianus; Pecari tajacu.

THE MOST COMMON MODEL FOR EVALUATING THE SUSTAINABILITY
of game hunting in the tropics is the one known as ‘population
growth model’ (Robinson & Redford 1991) or ‘production model’
(Bodmer & Robinson 2004). This model has been applied in many
case studies (e.g., Fitzgibbon et al. 1995, Alvard et al. 1996, Wilkie
et al. 1998, Naranjo et al. 2004). The production model requires
specific information on carrying capacity (K), maximum population
growth rate (λ max ), and density to estimate the maximum sustain-
able yield (MSY; Robinson & Redford 1991). This model assumes
that populations have a density dependent growth and reach max-
imum productivity when at 60 percent of the carrying capacity.
Therefore, maximum production (P max ) is calculated by multiplying
the density at 0.6K by the finite population growth rate (λ max ), and
then subtracting the previous year’s density, thus: P max = (0.6K ×
λ max ) − 0.6K.
Usually K is estimated by sampling populations in undisturbed
areas that are considered to be at carrying capacity, in other words
K = D. However two factors could affect this assumption. First,
population equilibrium does not necessarily occur at carrying ca-
pacity, especially in those areas where population growth is strongly
regulated by predation (Bodmer & Robinson 2004). Secondly, car-
rying capacity is not a static value but changes depending on pop-
ulation dynamics, seasonal variation, predation, and human needs
(McCullough 1987, Hunter & Runge 2004). Thus, in natural sys-
tems, there are fluctuations in environmental condition and popu-
lations fluctuate around K. As a consequence, a single estimation of
D in a little or undisturbed area, could be a biased estimation of K.
Received 25 November 2005; revision accepted 2 November 2006.
1 Correspondingauthor; e-mail: salvador.mandujano@inecol.edu.mx
However, few studies in the Neotropics have monitored environ-
mental changes in order to consider this aspect of K and the MSY
estimations (Bodmer & Robinson 2004).
In fact, the term carrying capacity sometimes causes confusion
(Miller & Wentworth 2000). The most commonly used definition
of K is the maximum number of animals of a given population that
can be supported by the available resources (McCullough 1992).
Thus, from a demographic point of view, the term refers to the
equilibrium density at which population growth stabilizes when
birth and death rates are equal (Akçakaya et al. 1999). But in terms
of the availability and use of habitat resources, carrying capacity
refers to the maximum population that a site can sustain without
suffering deterioration (Groot-Bruinderink & Hazeborek 1995). In
the case of herbivorous mammals, it has been assumed that a habi-
tat’s carrying capacity depends on the nutritional value of its plants
(McCall et al. 1997). There is a difference between ecological and
economic carrying capacity (Scoones 1993). From an economic or
social point of view, the concept of carrying capacity takes into
consideration the maximum number of animals that can exist with-
out economic repercussions to a specific human interest (West &
Parkhurst 2002). Another economic definition is the maximum
number of animals that can be sustainably harvested (DeCalesta
& Stout 1997). For human communities inhabiting tropical forest,
carrying capacity is the greatest number of people that the forest can
sustain without affecting biodiversity (Robinson & Bennet 1999).
Although related, these definitions differ and this is reflected in the
methods to estimate them.
The white-tailed deer (Odocoileus virginianus) and the collared
peccary (Pecari tajacu) are the two most widespread ungulates in the
New World (Hall 1981, Eisenberg 1989). In Mexico, they are widely

C 2007 The Author(s) 519
Journal compilation

C 2007 by The Association for Tropical Biology and Conservation
520 Mandujano
used in indigenous and rural settings for subsistence, meat for sale,
and in some areas for trophy hunting (Mandujano & Rico-Gray
1991, Naranjo et al. 2004). Therefore, it is important to evaluate
these ungulates’ potential productivity. In order to produce the
information necessary to employ the Robinson and Redford (1991)
model, it is important to determine habitat carrying capacity at sites
with undisturbed ungulate populations.
In this paper, I report estimates of carrying capacity and po-
tential production of white-tailed deer and collared peccary in a
protected area on the coast of Jalisco, Mexico. In comparison with
other regions, this is a reserve with little hunting by humans but
high natural predation by species such as puma (Puma concolor),
jaguar (Panthera onca), ocelot (Leopardus pardalis), and coyote (Ca-
nis latrans) (Nuñez et al. 2000, Hidalgo-Mihart et al. 2001, Villa-
Meza et al. 2002). Here, I defined carrying capacity as equilibrium
density plus the annual number of animals removed by predators.
Considering interannual variation in food availability depending
on precipitation (Mandujano 2006), it is reasonable to expect that
population density is not static even with an unhunted population.
Thus, in this study K is not considered as the current density of the
year i, i.e.,: K = D i . Instead, K is estimated considering the density
dependent relationships between finite population growth rate and
density using data collected over 5 yr. Also, in this paper I analyze
the possible variation of K based on data on precipitation over a
26-yr period, and discuss how this could influence management
strategies.
METHODS
STUDY AREA.—The study was carried out at the Chamela Bio-
logical Station of the UNAM (National Autonomous University
of Mexico), located on the coast of Jalisco, Mexico (19◦ 30 N,
105◦ 00 W). The Biological Station (3319 ha) is part of the
Chamela-Cuixmala Biosphere Reserve (www.ibiologia.unam.mx/
ebchamela/HIST2.html). The Station is characterized by slopes
of 21◦ to 34◦ and altitudinal ranges from 30 m to 580 m
asl. The mean annual temperature is 25◦ C (Bullock 1986). The
climate is tropical, warm subhumid, with marked seasonality
(www.ibiologia.unam.mx/ebchamela/clima.html). The yearly min-
imum and maximum average temperatures are 22.1◦ C and 30.3◦ C,
respectively. The annual mean rainfall from 1977 to 2003 was 755
± 233 mm. On average, 80 percent of the rain falls between July
and October. The dominant vegetation (covering > 80% of the bi-
ological station) is a tropical dry forest located on hilly terrain with
shallow soils and low water retention. The station also has tropical
semi-deciduous forest.
ESTIMATES OF UNGULATE DENSITIES.—Direct counts of deer and
peccaries in 5–8 transects were used to estimate population density.
Data were gathered in the rainy seasons of 1989 to 1993, and dry
seasons of 1990 to 1994. Counts were made two or three times
a month between 0700–1200 h and 1600–1900 h. A total of 6–
11 km was walked each month. For deer, a single observation was
considered the unit sample; in contrast, for peccaries, the observed
group was the unit sample. Each record was classified into one of
four perpendicular-distance classes: 0–10 m, 11–20 m, 21–30 m,
and 31–40 m. Density (D, deer or peccaries/km2 ) was calculated
using the DISTANCE 4.1 version 2 program (Thomas et al. 2003).
To obtain the density of individuals of peccaries, group density was
multiplied by the mean group size (Mandujano 1999). Student’s t-
tests were applied to compare annual estimates of deer and peccary
densities. The biomass was estimated by multiplying the annual
density by the animals’ mean adult weight: 15.7 kg for the collared
peccary and 42.1 kg for the white-tailed deer (Naranjo et al. 2004).
ESTIMATES OF FINITE GROWTH RATE.—The annual finite growth
rate (λ, lambda) was estimated as : λ = N t+1 /N t , where N t+1 is
the abundance of individuals per year t + 1 and N t being the
abundance in the previous year. The abundance was obtained by
multiplying the annual density by the area sampled. The maximum
finite population growth rate was estimated as λ max = ermax . The
intrinsic rate of natural increase (r max ) can only be calculated for
populations not limited by food, space, resource competition, or
predation (Robinson & Redford 1986). I estimated r max using Cole’s
equation (1954, see Robinson & Redford 1991), which has to be
solved iteratively as: 1 = e−rmax + be−rmax(a) −be−rmax(w+1) , where
a is the age at the first reproductive event, w is the age at the
last reproductive event, and b is the annual birth rate of female
offspring. The parameters a, b, and w were 1.5, 0.69, and 7; and 1,
2, and 7 for deer and peccary, respectively, and were obtained in the
study area (Mandujano & Martinez-Romero 2002, Mandujano &
Gallina 2005).
ESTIMATES OF CARRYING CAPACITY.—Carrying capacity was esti-
mated based on the density dependent relationship between popu-
lation growth and density according to a logistic model (Akçakaya et
al. 1999), plus the number of individuals removed by predators. A
regression was made using density as the independent variable and
the finite growth rate as the dependent variable, for each ungulate
species. Values of the finite rate greater than one indicate popula-
tion growth and values less than one indicate population decrease.
The point at which the regression intercepts the replacement line is
considered as the equilibrium density (Akçakaya et al. 1999). In the
study area, predators annually remove 36.7–42.7 percent, and 9–11
percent of the population of white-tailed deer and collared peccary,
respectively (Nuñez et al. 2000, Hidalgo-Mihart et al. 2001, Villa-
Meza et al. 2002). These numbers were added to the equilibrium
density, under the assumption that removal percentage by preda-
tors remains constant over the years. The result is an estimation of
carrying capacity.
ESTIMATES OF POTENTIAL PRODUCTION.—The maximum potential
production (P max ) was estimated as: P max = (0.6K × λ max ) − 0.6K.
(Robinson & Redford 1991). Potential production for human use
was considered to be P RR = 0.40P max . The factor 0.40 was used
because these species have intermediate life span (final reproductive
event at an age of < 10 yr, according to Slade et al. 1998). In other
words, it was assumed that up to 40 percent of maximum production
can be used without adversely affecting the population.
 Carrying Capacity and Production of Ungulates 521

ANALYSIS OF RAINFALL VARIATION.—In order to know the possible

TABLE 1. Parameters of white-tailed deer and collared peccary populations used
variation of K from 1977 to 2002, the rainfall pattern was an-
to estimate carrying capacity and potential production for human use
alyzed assuming that food availability depends on precipitation
in the tropical dry forest of Chamela, Mexico.
(Mandujano 2006). McCullough (1987) proposed a schematic
model of the frequency distribution of the occurrence of good and White-tailed Collared
bad years in different environments. The classification of a year as Parameters deer peccary
‘good’ or ‘bad’ depended on the deviation in precipitation of that
year compared with the average precipitation from the 26-yr period. Current population (individual/km2 )
The deviation was standardized as a percentage of the mean. Thus mean 11.8 7.5
zero value represents the mean, a value of < 0 represents a ‘bad’ range 11.0–13.6 4.1–10.7
year, while a value of > 0 represents a ‘good’ year. The mean, me- Biomass (kg/km2 ) 463–573a 64–168
dian, skewed and kurtosis values were used as an index to evaluate Maximum finite population growth rateb 1.52 3.00
where the distribution frequency is according to a ‘stable–unstable’ Equilibrium density (individual/km2 )c 12.0 8.6
and ‘good–poor’ environment. According to McCullough (1987), Animals removed by predators (individual/km2 )d 4.5–5.2 0.7–0.9
a leptokurtic (γ 2 > 0) and platykurtic (γ 2 < 0) frequency distri- Carrying capacitye
bution represents highly stable and highly unstable environments, individual/km2 16.5–17.2 9.3–9.5
respectively; while a positively skewed (γ 1 > 0) and negatively kg/km2 695–724 146–150
skewed (γ 1 < 0) distribution represents a poor environment with Maximum sustainable yield (MSY)
occasional good years, and a good environment with occasional bad individual/km2 10.3 5.6
years, respectively. Data analysis was based on the STATISTICA kg/km2 434 89
Program for Windows, version 5.5 (Statsoft Inc. 2000).
Maximum production (P max )f
individual/km2 5.3 11.3
kg/km2 223 178
RESULTS Production for human use (0.40 × P max )
individual/km2 2.1 4.5
I walked 817 km and observed 363 deer and 44 peccary groups
kg/km2 88 71
(106 individuals). The mean annual densities were estimated as
11.8 deer/km2 and 7.5 peccaries/km2 (Table 1). In a 5-yr period, a Weight values used for the estimate:15.7 kg for the peccary and 42.1 kg for
the fluctuation of population density was lower for white-tailed the deer.
deer than for collared peccary (Fig. 1). In all years, deer density was b Maximum finite population growth rate (λ
max ) was obtained from Cole’s
significantly greater than for peccaries (t = 2.86, df = 8, P = 0.02). equation.
The biomass of deer was estimated to be 463 to 573 kg/km2 , while c Equilibriumdensity was estimated according to Fig. 2.
for peccaries the figure was 64 to 168 kg/km2 (Table 1). d Considering that in this study area predators annually remove 36.7–42.7
The estimated maximum finite population growth rate (λ max ) percent, and 9–11 percent of the population of white-tailed deer and collared
was 1.52 and 3.0 for white-tailed deer and collared peccary, respec- peccary, respectively (Nuñez et al. 2000, Hidalgo-Mihart et al. 2001, Villa-Meza
tively. Annual population growth rate varied positively and nega- et al. 2002).
tively in both species. The equilibrium density was estimated as e Carrying capacity is the sum of equilibrium density plus animals removed by

12.0 deer/km2 and 8.6 peccaries/km2 (Fig. 2), while the carrying
capacity was estimated at 16.5 to 17.2 deer/km2 and 9.3 to 9.5
peccaries/km2 (Table 1).
MSY was estimated as 10.3 deer/km2 and 5.6 peccaries/km2
(Table 1), while maximum production was estimated as 5.2 to 5.4
deer/km2 and 11.1 to 11.4 peccaries/km2 (Table 1). Considering a
harvest of 40 percent of maximum productivity, potential produc-
tion for human use was calculated to be 2.1 to 2.2 deer/km2 and
4.4. to 4.6 peccaries/km2 . Because of the deer’s greater weight, the
deer harvest was larger than that for the collared peccary (Table 1).
Annual precipitation during the 26-yr period is presented in
Fig. 1. The precipitation varied notably from year to year from
1989 to 1994 when the density of ungulates was estimated. Over
26 yr, 14 yr had a precipitation from 1 to 50 percent lower than
the mean. In contrast, only 3 yr presented a precipitation 50–80
percent higher than mean (Fig. 3). Even though the frequency
distribution fitted a normal curve (K-S d = 0.121, P = 0.20),
the median value was −5.0 percent with respect to the mean. In
predators.
f Maximum production was estimated according to Robinson and Redford’s
(1991) model.
consequence, data show a slightly positively skewed distribution
(γ 1 = 0.926), while the kurtosis value (γ 2 = 0.431) suggests a
slightly leptokurtic distribution. This analysis suggests a moderately
stable, poor environment with occasional good years during the
1977–2002 period.
DISCUSSION
The data suggest that, in the protected tropical dry forest of
Chamela, the current density (15.1 to 24.3 individuals/km2 ) and
biomass (527 to 741 kg/km2 ) of wild ungulates are relatively high.
For example, in tropical dry forest with patches of semi-open habi-
tat in Venezuela and Brazil, wild ungulate biomass ranges from 224
522 Mandujano
FIGURE 1. Time series of the annual rainfall from 1977 to 2002 (gray bars),
and population density of white-tailed deer (full circles) and collared peccary
(empty circles) during the period under study from 1989 to 1993.
to 275 kg/km2 (Robinson & Bennett 1999). In tropical rain forests
with up to five species of ungulates (Tapirus terrestris, Tayassu pecari,
P. tajacu, Mazama americana, and M. gouazoubira), there may be
a combined density and biomass of 28.3 individuals/km2 and 761
kg/km2 , respectively (Robinson & Redford 1991), although data
from Panama, Peru, Brazil and Venezuela suggest that the ungulate
biomass varies from 270 and 542 kg/km2 (Robinson & Bennett
2004). In the Selva Lacandona, Mexico, the combined density and
biomass of five ungulate species in areas with little hunting are
9.9 individuals/km2 and 337 kg/km2 (Naranjo et al. 2004). These
comparisons suggest that the tropical dry forest of Chamela, with
a lower species richness of ungulates, can maintain a similar or
greater density and biomass than other tropical forests. However,
FIGURE 2. Relationship between the finite population growth rate (λ) and the
population density of white-tailed deer (full circles) and collared peccary (empty
circles). The horizontal line with value λ = 1 is known as the replacement line.
The point where the curve intercepts the replacement line marks the equilibrium
point of population growth: 12 individuals/km2 for white-tailed deer and 8.6
individuals/km2 for collared peccary.
FIGURE 3. Frequency distribution of the occurrence of good and bad years
in the tropical dry forest of Chamela from 1977 to 2002. In this study a ‘good’
or ‘bad’ year was classified depending on the deviation of precipitation of a
specific year with respect to the average precipitation over the 26-yr period. The
deviation was standardized as a percentage with respect to the mean. Thus zero
value represents the mean, a value < 0 represents a ‘bad’ year, while a value > 0
represents a ‘good’ year. The continuous line represents the data corresponding
to a normal distribution.
note that these studies varied in sampling design and effort. In con-
sequence, the potential production of ungulates for human use in
this tropical dry forest is relatively higher than other tropical for-
est (e.g., Robinson & Redford 1991, Bodmer & Robinson 2004,
Naranjo et al. 2004). Species with large baseline populations and
faster reproduction, such as peccaries and deer, are less vulnerable
to overhunting (McCullough 1987, Bodmer & Robinson 2004).
However, because in tropical habitats it is difficult to monitor pop-
ulations and habitat over a long period, most harvesting of wildlife
has been managed largely by trial and error (Caughley & Sinclair
1994). This approach works well when the species has a high in-
trinsic rate of increase, a density dependent relationship, and the
population size is greater than 60 percent of K.
The model of Robinson and Redford (1991) assumed that K =
D in sampled populations in little or undisturbed areas. The data
provided for the protected tropical dry forest of Chamela, showed
that the current density of both ungulates varied during the study
period. Thus, a single estimation of D could be a biased estima-
tion of K. To avoid this, Bodmer and Robinson (2004) suggested
monitoring the population, habitat, and hunting to incorporate the
variation into the models used to assess the sustainability of hunting.
The estimation of the equilibrium density obtained in this study
considers the interannual variation in the population change. Ob-
viously, this procedure needs to monitor the population for several
years. This is important because the dynamics of plant and ungulate
interactions are very complex, and herbivores can influence vege-
tation and the future of herbivore populations, and the potential
for human use (McCullough 1987, Marshal et al. 2002, Ogutu &

Owen-Smith 2003). It is important to clarify that equilibrium den-
sity is not simply the average annual density during the period under
study. Instead, it is calculated according to the density dependent
relationship between annual growth rate and current density. Con-
sequently, the equilibrium density represents the population trend
over several years.
As pointed out earlier, population equilibrium does not nec-
essarily occur at carrying capacity, especially in those areas where
population growth is strongly regulated by predation (Bodmer &
Robinson 2004). For the tropical dry forest of Chamela, Nuñez
et al. (2000) proposed that, in the absence of predators, ungulate
populations could reach the maximum limit permitted by their
habitat. These data seem to corroborate that of other studies that
have indicated that predators maintain ungulates below their car-
rying capacity (Skogland 1991). However, to distinguish the equi-
librium produced by predators, by resource limitation, and by a
combination of the two, we need to know whether predators or
resources or both are affecting the birth and death rates (Caughley
& Sinclair 1994). Unfortunately, this level of detail is not avail-
able for the study site. Thus, even if predators depress ungulate
populations below carrying capacity, the relationship between food
availability, population density, and mortality factors is complex
and requires many years of monitoring to understand better the
underlying mechanisms (McCullough 1992, Sæther 1997).
An important result of the present study is that animals re-
moved by predators must be added to the equilibrium density, in
order to accurately estimate the K and MSY. This is more important
for those species with a higher percent of depredation, for example
the white-tailed deer in the area under study. An underestimation
of MSY could put the ungulate population at risk (McCullough
1987). To clarify this point, consider the following arguments (see
Bodmer & Robinson 2004). A sustainable harvest can be realized
at any base population size, but there is only one point at which the
sustained harvest is at the maximum, or MSY. Harvesting species
at the theoretical MSY or with small base populations (to the left
of MSY) is a risk-management strategy and should be avoided. If
small misjudgments occur in calculating the sustainable harvest,
this overhunting would result in a decreased base population in
the following year and would quickly lead to overexploitation and
extirpation. In contrast, harvesting species with large base popula-
tions (to the right of MSY) is a safe management strategy that can
be used for long-term sustainable use of a species. As McCullough
(1987) showed, if small misjudgments occur with harvesting with
a large base population, this slight overhunting would result in a
decreased base population in the following year. But the population
would stabilize at this new sustained harvest level and would not de-
crease further. In consequence, harvesting a species with large base
populations is a safer management strategy, since it is less likely to re-
sult in extirpation. This is especially important for predator-limited
species in which prey densities are held below K by predation mor-
tality. An underestimate of K would lead to an underestimate of
MSY and a misrepresentation in the relationship between N and
the actual MSY. Thus, the results suggest that a safe management
strategy in the studied region will be hunting deer and peccary
for human use only when population density is above 10.3 and
Carrying Capacity and Production of Ungulates 523
5.6 individuals/km2 , respectively. This strategy could protect both
ungulate and predator species.
The long-term analysis of precipitation in a specific area could
help in the development of management guidelines for ungulates
of that area. The rain pattern analysis suggests that the tropical dry
forest is a moderately stable habitat characterized by poor years with
sporadic years of heavy precipitation. Both environmental variance
and the frequency distribution of good and poor years influence
the precision of management that is possible and the management
strategy that should be applied (McCullough 1987). In an environ-
ment with random or periodic variation in the carrying capacity,
populations with a large growth rate (for example small mammals)
tend to follow the variation, while populations with a small growth
rate tend to average it (May 1976). Due to their large size and long
life spans, ungulates typically exist in populations close to environ-
mental carrying capacity (McCullough 1992). For example, over
a 5-yr period, the fluctuation of population density was lower in
white-tailed deer with a lower growth rate than collared peccary
with a higher population growth rate. McCullough (1987) suggests
that, in highly variable environments, management must respond
to the uncontrollable environment variables on an ad hoc basis. In
contrast, in habitats that have many good years, management can
be precise, but modifications would be needed for the poor years.
This analysis suggests that, in the tropical dry forest of Chamela,
management can also be relatively precise and reflect density de-
pendent responses in habitat where most years are poor with a few
good years. But in a period with consecutive poor years, precaution
must be taken to avoid overexploitation.
ACKNOWLEDGMENTS
This study was supported by the CONACYT and the Department
of Biodiversity and Animal Ecology of Institute of Ecology A. C.
Xalapa, Mexico. Sonia Gallina, Hector Gomez de Silva, Carolina
Valdespino and two anonymous reviewers provided useful com-
ments. Judy Shirley checked the final version of the manuscript in
English.
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