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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 However, few studies in the Neotropics have monitored environ-
of game hunting in the tropics is the one known as ‘population mental changes in order to consider this aspect of K and the MSY
growth model’ (Robinson & Redford 1991) or ‘production model’ estimations (Bodmer & Robinson 2004).
(Bodmer & Robinson 2004). This model has been applied in many In fact, the term carrying capacity sometimes causes confusion
case studies (e.g., Fitzgibbon et al. 1995, Alvard et al. 1996, Wilkie (Miller & Wentworth 2000). The most commonly used definition
et al. 1998, Naranjo et al. 2004). The production model requires of K is the maximum number of animals of a given population that
specific information on carrying capacity (K), maximum population can be supported by the available resources (McCullough 1992).
growth rate (λ max ), and density to estimate the maximum sustain- Thus, from a demographic point of view, the term refers to the
able yield (MSY; Robinson & Redford 1991). This model assumes equilibrium density at which population growth stabilizes when
that populations have a density dependent growth and reach max- birth and death rates are equal (Akçakaya et al. 1999). But in terms
imum productivity when at 60 percent of the carrying capacity. of the availability and use of habitat resources, carrying capacity
Therefore, maximum production (P max ) is calculated by multiplying refers to the maximum population that a site can sustain without
the density at 0.6K by the finite population growth rate (λ max ), and suffering deterioration (Groot-Bruinderink & Hazeborek 1995). In
then subtracting the previous year’s density, thus: P max = (0.6K × the case of herbivorous mammals, it has been assumed that a habi-
λ max ) − 0.6K. tat’s carrying capacity depends on the nutritional value of its plants
Usually K is estimated by sampling populations in undisturbed (McCall et al. 1997). There is a difference between ecological and
areas that are considered to be at carrying capacity, in other words economic carrying capacity (Scoones 1993). From an economic or
K = D. However two factors could affect this assumption. First, social point of view, the concept of carrying capacity takes into
population equilibrium does not necessarily occur at carrying ca- consideration the maximum number of animals that can exist with-
pacity, especially in those areas where population growth is strongly out economic repercussions to a specific human interest (West &
regulated by predation (Bodmer & Robinson 2004). Secondly, car- Parkhurst 2002). Another economic definition is the maximum
rying capacity is not a static value but changes depending on pop- number of animals that can be sustainably harvested (DeCalesta
ulation dynamics, seasonal variation, predation, and human needs & Stout 1997). For human communities inhabiting tropical forest,
(McCullough 1987, Hunter & Runge 2004). Thus, in natural sys- carrying capacity is the greatest number of people that the forest can
tems, there are fluctuations in environmental condition and popu- sustain without affecting biodiversity (Robinson & Bennet 1999).
lations fluctuate around K. As a consequence, a single estimation of Although related, these definitions differ and this is reflected in the
D in a little or undisturbed area, could be a biased estimation of K. methods to estimate them.
The white-tailed deer (Odocoileus virginianus) and the collared
Received 25 November 2005; revision accepted 2 November 2006. peccary (Pecari tajacu) are the two most widespread ungulates in the
1 Correspondingauthor; e-mail: salvador.mandujano@inecol.edu.mx 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, group was the unit sample. Each record was classified into one of
and in some areas for trophy hunting (Mandujano & Rico-Gray four perpendicular-distance classes: 0–10 m, 11–20 m, 21–30 m,
1991, Naranjo et al. 2004). Therefore, it is important to evaluate and 31–40 m. Density (D, deer or peccaries/km2 ) was calculated
these ungulates’ potential productivity. In order to produce the using the DISTANCE 4.1 version 2 program (Thomas et al. 2003).
information necessary to employ the Robinson and Redford (1991) To obtain the density of individuals of peccaries, group density was
model, it is important to determine habitat carrying capacity at sites multiplied by the mean group size (Mandujano 1999). Student’s t-
with undisturbed ungulate populations. tests were applied to compare annual estimates of deer and peccary
In this paper, I report estimates of carrying capacity and po- densities. The biomass was estimated by multiplying the annual
tential production of white-tailed deer and collared peccary in a density by the animals’ mean adult weight: 15.7 kg for the collared
protected area on the coast of Jalisco, Mexico. In comparison with peccary and 42.1 kg for the white-tailed deer (Naranjo et al. 2004).
other regions, this is a reserve with little hunting by humans but
high natural predation by species such as puma (Puma concolor), ESTIMATES OF FINITE GROWTH RATE.—The annual finite growth
jaguar (Panthera onca), ocelot (Leopardus pardalis), and coyote (Ca- rate (λ, lambda) was estimated as : λ = N t+1 /N t , where N t+1 is
nis latrans) (Nuñez et al. 2000, Hidalgo-Mihart et al. 2001, Villa- the abundance of individuals per year t + 1 and N t being the
Meza et al. 2002). Here, I defined carrying capacity as equilibrium abundance in the previous year. The abundance was obtained by
density plus the annual number of animals removed by predators. multiplying the annual density by the area sampled. The maximum
Considering interannual variation in food availability depending finite population growth rate was estimated as λ max = ermax . The
on precipitation (Mandujano 2006), it is reasonable to expect that intrinsic rate of natural increase (r max ) can only be calculated for
population density is not static even with an unhunted population. populations not limited by food, space, resource competition, or
Thus, in this study K is not considered as the current density of the predation (Robinson & Redford 1986). I estimated r max using Cole’s
year i, i.e.,: K = D i . Instead, K is estimated considering the density equation (1954, see Robinson & Redford 1991), which has to be
dependent relationships between finite population growth rate and solved iteratively as: 1 = e−rmax + be−rmax(a) −be−rmax(w+1) , where
density using data collected over 5 yr. Also, in this paper I analyze a is the age at the first reproductive event, w is the age at the
the possible variation of K based on data on precipitation over a last reproductive event, and b is the annual birth rate of female
26-yr period, and discuss how this could influence management offspring. The parameters a, b, and w were 1.5, 0.69, and 7; and 1,
strategies. 2, and 7 for deer and peccary, respectively, and were obtained in the
study area (Mandujano & Martinez-Romero 2002, Mandujano &
Gallina 2005).
METHODS
ESTIMATES OF CARRYING CAPACITY.—Carrying capacity was esti-
STUDY AREA.—The study was carried out at the Chamela Bio- mated based on the density dependent relationship between popu-
logical Station of the UNAM (National Autonomous University lation growth and density according to a logistic model (Akçakaya et
of Mexico), located on the coast of Jalisco, Mexico (19◦ 30 N, al. 1999), plus the number of individuals removed by predators. A
105◦ 00 W). The Biological Station (3319 ha) is part of the regression was made using density as the independent variable and
Chamela-Cuixmala Biosphere Reserve (www.ibiologia.unam.mx/ the finite growth rate as the dependent variable, for each ungulate
ebchamela/HIST2.html). The Station is characterized by slopes species. Values of the finite rate greater than one indicate popula-
of 21◦ to 34◦ and altitudinal ranges from 30 m to 580 m tion growth and values less than one indicate population decrease.
asl. The mean annual temperature is 25◦ C (Bullock 1986). The The point at which the regression intercepts the replacement line is
climate is tropical, warm subhumid, with marked seasonality considered as the equilibrium density (Akçakaya et al. 1999). In the
(www.ibiologia.unam.mx/ebchamela/clima.html). The yearly min- study area, predators annually remove 36.7–42.7 percent, and 9–11
imum and maximum average temperatures are 22.1◦ C and 30.3◦ C, percent of the population of white-tailed deer and collared peccary,
respectively. The annual mean rainfall from 1977 to 2003 was 755 respectively (Nuñez et al. 2000, Hidalgo-Mihart et al. 2001, Villa-
± 233 mm. On average, 80 percent of the rain falls between July Meza et al. 2002). These numbers were added to the equilibrium
and October. The dominant vegetation (covering > 80% of the bi- density, under the assumption that removal percentage by preda-
ological station) is a tropical dry forest located on hilly terrain with tors remains constant over the years. The result is an estimation of
shallow soils and low water retention. The station also has tropical carrying capacity.
semi-deciduous forest.
ESTIMATES OF POTENTIAL PRODUCTION.—The maximum potential
ESTIMATES OF UNGULATE DENSITIES.—Direct counts of deer and production (P max ) was estimated as: P max = (0.6K × λ max ) − 0.6K.
peccaries in 5–8 transects were used to estimate population density. (Robinson & Redford 1991). Potential production for human use
Data were gathered in the rainy seasons of 1989 to 1993, and dry was considered to be P RR = 0.40P max . The factor 0.40 was used
seasons of 1990 to 1994. Counts were made two or three times because these species have intermediate life span (final reproductive
a month between 0700–1200 h and 1600–1900 h. A total of 6– event at an age of < 10 yr, according to Slade et al. 1998). In other
11 km was walked each month. For deer, a single observation was words, it was assumed that up to 40 percent of maximum production
considered the unit sample; in contrast, for peccaries, the observed 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 predators.
capacity was estimated at 16.5 to 17.2 deer/km2 and 9.3 to 9.5 f Maximum production was estimated according to Robinson and Redford’s
peccaries/km2 (Table 1). (1991) model.
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 consequence, data show a slightly positively skewed distribution
harvest of 40 percent of maximum productivity, potential produc- (γ 1 = 0.926), while the kurtosis value (γ 2 = 0.431) suggests a
tion for human use was calculated to be 2.1 to 2.2 deer/km2 and slightly leptokurtic distribution. This analysis suggests a moderately
4.4. to 4.6 peccaries/km2 . Because of the deer’s greater weight, the stable, poor environment with occasional good years during the
deer harvest was larger than that for the collared peccary (Table 1). 1977–2002 period.
Annual precipitation during the 26-yr period is presented in
Fig. 1. The precipitation varied notably from year to year from DISCUSSION
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 data suggest that, in the protected tropical dry forest of
the mean. In contrast, only 3 yr presented a precipitation 50–80 Chamela, the current density (15.1 to 24.3 individuals/km2 ) and
percent higher than mean (Fig. 3). Even though the frequency biomass (527 to 741 kg/km2 ) of wild ungulates are relatively high.
distribution fitted a normal curve (K-S d = 0.121, P = 0.20), For example, in tropical dry forest with patches of semi-open habi-
the median value was −5.0 percent with respect to the mean. In 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.

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’
to 275 kg/km2 (Robinson & Bennett 1999). In tropical rain forests or ‘bad’ year was classified depending on the deviation of precipitation of a
with up to five species of ungulates (Tapirus terrestris, Tayassu pecari, specific year with respect to the average precipitation over the 26-yr period. The
P. tajacu, Mazama americana, and M. gouazoubira), there may be deviation was standardized as a percentage with respect to the mean. Thus zero
a combined density and biomass of 28.3 individuals/km2 and 761 value represents the mean, a value < 0 represents a ‘bad’ year, while a value > 0
kg/km2 , respectively (Robinson & Redford 1991), although data represents a ‘good’ year. The continuous line represents the data corresponding
from Panama, Peru, Brazil and Venezuela suggest that the ungulate to a normal distribution.
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 note that these studies varied in sampling design and effort. In con-
9.9 individuals/km2 and 337 kg/km2 (Naranjo et al. 2004). These sequence, the potential production of ungulates for human use in
comparisons suggest that the tropical dry forest of Chamela, with this tropical dry forest is relatively higher than other tropical for-
a lower species richness of ungulates, can maintain a similar or est (e.g., Robinson & Redford 1991, Bodmer & Robinson 2004,
greater density and biomass than other tropical forests. However, 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
FIGURE 2. Relationship between the finite population growth rate (λ) and the
considers the interannual variation in the population change. Ob-
population density of white-tailed deer (full circles) and collared peccary (empty
viously, this procedure needs to monitor the population for several
circles). The horizontal line with value λ = 1 is known as the replacement line.
years. This is important because the dynamics of plant and ungulate
The point where the curve intercepts the replacement line marks the equilibrium
interactions are very complex, and herbivores can influence vege-
point of population growth: 12 individuals/km2 for white-tailed deer and 8.6
tation and the future of herbivore populations, and the potential
individuals/km2 for collared peccary.
for human use (McCullough 1987, Marshal et al. 2002, Ogutu &
Carrying Capacity and Production of Ungulates 523

Owen-Smith 2003). It is important to clarify that equilibrium den- 5.6 individuals/km2 , respectively. This strategy could protect both
sity is not simply the average annual density during the period under ungulate and predator species.
study. Instead, it is calculated according to the density dependent The long-term analysis of precipitation in a specific area could
relationship between annual growth rate and current density. Con- help in the development of management guidelines for ungulates
sequently, the equilibrium density represents the population trend of that area. The rain pattern analysis suggests that the tropical dry
over several years. forest is a moderately stable habitat characterized by poor years with
As pointed out earlier, population equilibrium does not nec- sporadic years of heavy precipitation. Both environmental variance
essarily occur at carrying capacity, especially in those areas where and the frequency distribution of good and poor years influence
population growth is strongly regulated by predation (Bodmer & the precision of management that is possible and the management
Robinson 2004). For the tropical dry forest of Chamela, Nuñez strategy that should be applied (McCullough 1987). In an environ-
et al. (2000) proposed that, in the absence of predators, ungulate ment with random or periodic variation in the carrying capacity,
populations could reach the maximum limit permitted by their populations with a large growth rate (for example small mammals)
habitat. These data seem to corroborate that of other studies that tend to follow the variation, while populations with a small growth
have indicated that predators maintain ungulates below their car- rate tend to average it (May 1976). Due to their large size and long
rying capacity (Skogland 1991). However, to distinguish the equi- life spans, ungulates typically exist in populations close to environ-
librium produced by predators, by resource limitation, and by a mental carrying capacity (McCullough 1992). For example, over
combination of the two, we need to know whether predators or a 5-yr period, the fluctuation of population density was lower in
resources or both are affecting the birth and death rates (Caughley white-tailed deer with a lower growth rate than collared peccary
& Sinclair 1994). Unfortunately, this level of detail is not avail- with a higher population growth rate. McCullough (1987) suggests
able for the study site. Thus, even if predators depress ungulate that, in highly variable environments, management must respond
populations below carrying capacity, the relationship between food to the uncontrollable environment variables on an ad hoc basis. In
availability, population density, and mortality factors is complex contrast, in habitats that have many good years, management can
and requires many years of monitoring to understand better the be precise, but modifications would be needed for the poor years.
underlying mechanisms (McCullough 1992, Sæther 1997). This analysis suggests that, in the tropical dry forest of Chamela,
An important result of the present study is that animals re- management can also be relatively precise and reflect density de-
moved by predators must be added to the equilibrium density, in pendent responses in habitat where most years are poor with a few
order to accurately estimate the K and MSY. This is more important good years. But in a period with consecutive poor years, precaution
for those species with a higher percent of depredation, for example must be taken to avoid overexploitation.
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 ACKNOWLEDGMENTS
Bodmer & Robinson 2004). A sustainable harvest can be realized
at any base population size, but there is only one point at which the This study was supported by the CONACYT and the Department
sustained harvest is at the maximum, or MSY. Harvesting species of Biodiversity and Animal Ecology of Institute of Ecology A. C.
at the theoretical MSY or with small base populations (to the left Xalapa, Mexico. Sonia Gallina, Hector Gomez de Silva, Carolina
of MSY) is a risk-management strategy and should be avoided. If Valdespino and two anonymous reviewers provided useful com-
small misjudgments occur in calculating the sustainable harvest, ments. Judy Shirley checked the final version of the manuscript in
this overhunting would result in a decreased base population in English.
the following year and would quickly lead to overexploitation and
extirpation. In contrast, harvesting species with large base popula-
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