BIOTROPICA 38(5): 695–699 2006 10.1111/j.1744-7429.2006.00184.

Preliminary Evidence of the Importance of ENSO in Modifying Food Availability

for White-tailed Deer in a Mexican Tropical Dry Forest1

Salvador Mandujano2
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
The influence of El Niño/Southern Oscillation (ENSO) on rainfall and its possible effect on availability of food for white-tailed deer (Odocoileus virginianus) in a
tropical dry forest in the Pacific coast of Mexico was studied. From 1977 to 2003 there were three significant El Niño and La Niña events. During El Niño years
rainfall decreased during the wet season ( June to October) and increased during the dry season (November to May), with the opposite effect during La Niña years.
Plant diversity was monitored in permanent plots during the wet and dry seasons of 1989–1993. The results provide evidence that ENSO events affect deer food
availability, particularly in the dry season.

RESUMEN
Este trabajo analiza la influencia de los eventos de El Niño/Oscilación del Sur (ENSO) sobre el patrón de precipitación y su posible influencia sobre la disponibilidad
de alimento para el venado cola blanca (Odocoileus virginianus) en un bosque tropical seco de la costa Pacı́fica de México. En el periodo de 1977–2003 se produjo tres
eventos claros de El Niño y de La Niña. Durante los años del El Niño la lluvia en la época húmeda (junio a octubre) disminuyó, y aumentó durante la época seca
(noviembre a mayo); mientras que durante La Niña se observó un efecto contrario. La diversidad de plantas en el sotobosque fue monitoreada en parcelas permanentes
durante las épocas de lluvia y sequı́a de 1989–1993. Los resultados sugieren que ENSO podrı́a tener un efecto adicional sobre la disponibilidad de alimento para el
venado, principalmente en la época seca.

Key words: El Niño/Southern Oscillation; food availability; Mexico; Odocoileus virginianus; rainfall; tropical dry forest.

CLIMATIC VARIATION ASSOCIATED WITH THE NORTH ATLANTIC OS-
CILLATION (NAO) and El Niño/Southern Oscillation (ENSO) has
a widespread influence on biosphere primary production and on
the population dynamics of many organisms worldwide (Behren-
feld et al. 2001, Stenseth et al. 2003). These large-scale climate
fluctuations affect local weather through teleconnection patterns
(Stenseth et al. 2003). For example, ENSO episodes are associated
with rainfall patterns in the Pacific region, which affect both marine
and terrestrial ecosystems ( Jaksic 2001). Weather and ENSO effects
can have important consequences for the demography of terrestrial
herbivores (e.g., Owen-Smith 1990, Langvatn et al. 1996, Sæther
1997, McKinney et al. 2001, Marshal et al. 2002, Georgiadis et al.
2003, Oguto & Owen-Smith 2003), because they influence plant
phenology, forage quality, and biomass production, which in turn
affect habitat carrying capacity (Coe et al. 1976, Sinclair et al. 1985,
Langvatn et al. 1996, Sæther 1997, Post & Stenseth 1999). Evidence
for these effects on ungulate populations has been generated prin-
cipally in temperate regions and in tropical Africa; in contrast, few
studies exist in Neotropical forests (e.g., Wright et al. 1999, Ticktin
2003).
In the Neotropical region, the white-tailed deer (Odocoileus
virginianus) is used by indigenous and rural people to complement
their diet and, in some places, as a commercial and game trophy
(Mandujano & Rico-Gray 1991, Escamilla et al. 2000, Naranjo
1 Received 28 April 2005; revision accepted 22 November 2005.
2 Corresponding author; e-mail: mandujan@ecologia.edu.mx

C 2006 The Author(s)
Journal compilation 
C 2006 by The Association for Tropical Biology and Conservation
et al. 2004). Thus, white-tailed deer is an important species from
the management perspective. This study was carried out in a trop-
ical dry forest in Chamela, located on the Mexican central Pacific
coast, with densities of 10–14 deer/km2 (Mandujano et al. 2002).
The seasonality of rainfall is the main factor that influences this
ecosystem’s structure and dynamics (Murphy & Lugo 1986). Plant
communities in Chamela are affected by interannual variation in
trade winds, Pacific hurricanes, and ENSO events (Bullock 1986,
Garcia-Oliva et al. 1991). Along with rainfall, topography influ-
ences the availability of water on the ground, producing extensive
areas of tropical dry forest on slopes and mountaintops while narrow
patches of semideciduous tropical forest establish on riverbanks and
near streams (Lott et al. 1987, Bullock & Solis-Magallanes 1990).
Therefore, this forest is characterized by both temporal and spatial
variations in food availability, nutrients, and water for white-tailed
deer. The first part of this paper analyzes the effect of ENSO on
rainfall patterns in the Chamela study area using data collected over
26 yr. The second part of the paper provides preliminary evidence
for the importance of ENSO in modifying temporal food availabil-
ity for white-tailed deer using data collected over 4 yr.
The study was carried out at the Chamela Biological
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 (http://www.ibiologia.unam.
mx/ebchamela/HIST2.html). The Station is characterized by slopes
of 21◦ to 34◦ and altitudinal ranges from 30 to 580 m above
695
696 Mandujano
sea level. Mean annual temperature is 25◦ C (Bullock 1986).
The climate is tropical, warm subhumid, with a marked season-
ality (http://www.ibiologia.unam.mx/ebchamela/clima.html). The
yearly minimum and maximum average temperatures are 22.1◦ C
and 30.3◦ C, respectively. The average number of days with sub-
stantial rain is 52, with approximately five strong rains per year
(Bullock 1986). The average annual rainfall from 1977 to 2003 was
755±233 mm, with a total of 1394 mm in the rainiest year (1992)
and 392 mm in the driest (2001). On average, 80 percent of the
rain falls between July and October, whereas 20 percent falls during
the dry season from November to June. Around 90 percent of all
plant species begin to foliate with a minimum of 100 mm of rain
(Bullock & Solis-Magallanes 1990).
The dominant vegetation (covering >80% of the biological
station) is tropical dry forest located on hilly terrain with shallow
soils and low water retention. Many tree and shrub species lose
their leaves during the dry season. Forest height varies between 4
and 15 m and there is a well-developed understory (Lott et al.
1987). Common trees are Cordia alliodora [Ruiz y Pav.] Oken,
Lonchocarpus lanceolatus Benth., and Caesalpinia eriostachys Benth.
The station also has tropical semideciduous forest, ranging from
10 m to 25 m in height. This type of forest is found along the
streams in deep soils with high water retention. A lower percentage
of tropical semideciduous forest tree species lose their leaves during
FIGURE 1. Time series of Southern Oscillation Index values and annual rainfall (wet season plus dry season) for 1977–2003.
the dry season. The most common trees are Astronium graveolens
Jacq., Brosimum alicastrum Sw., and Sideroxylon capiri A. DC.
In this study, the year was considered as beginning in the
month of the first important rain, typically around the third week
of June (Bullock 1986). The probability of having 100 mm of rain
defines a 5-mo wet season ( June to October) and 7-mo dry sea-
son (November to May; Garcia-Oliva et al. 1991). Rainfall data
from 1977 to 2003 were obtained from the station’s weather sta-
tion. ENSO information was obtained from the US Department
of Commerce data base (http://www.pmel.noaa.gov/tao/elnino/el-
boy-story.html), as well as from the Climate Diagnostics Cen-
ter (http://www.cdc.noaa.gov./ENSO/enso.different.html) and the
University of Illinois (http://www.ww2010.atmos.uiuc.edu/(G1)/
wwhlpr/guides/mtr/eln/def.rxml). From these data bases, a tempo-
ral pattern of the Southern Oscillation Index (SOI) was obtained
for the period 1977–2003, and SOI was calculated grouping data
from June to May. Negative values of SOI are associated with the
intrusions of El Niño (warm phase), and positive values with those
of La Niña (cold phase; Stenseth et al. 2003). The relationship be-
tween the SOI and annual and seasonal rainfall was analyzed with
linear correlations using the least squares procedure.
To estimate food availability, 25 permanent plots (10×10 m
each) were randomly established in tropical dry forest and 25 in
tropical semideciduous forest. During each sampling period, a 1 m2
 Short Communications 697

quadrant was randomly established; only the young leaves and

branches of all species on the understory were collected. Samplings
were done in October for the wet seasons of 1989–1992 and in
May for the dry seasons of 1990–1993. Only the species consumed
by deer at this site according to Arceo et al. (2005) were consid-
ered. Information on food availability was estimated as the standing
crop biomass (g/m2 ) and species richness (number of species/m2 ).
However, because these variables were significantly correlated both
in tropical dry (df 7 , R2 = 0.79, P = 0.02) and semideciduous forest
(df 7 , R2 = 0.88, P = 0.004), the Shannon–Wiener diversity index
(H ) was used as an estimate of the food available to the deer. The
index was estimated for each quadrant and then averaged to obtain
a seasonal index for each forest type.
Data analysis showed that from 1977 to 1984, annual rain-
fall was near to the average of 755 mm; this was followed by two
dry periods in 1985–1988 and in 2001–2003. The decade from
1989 to 2000 showed high variation in annual rainfall (Fig. 1).
The average rainfall during the wet seasons ( June to October) was
665 ± 222 mm (range: 368–1288 mm), while in the dry seasons
(November to May) it was 90±148 mm (range: 0–721 mm). The
rain that fell during the dry season was independent of the rain-
fall from the previous wet season (df 1,24 , R2 = 0.07, F = 1.73,
P = 0.20).
During the 26 yr analyzed, there were three strong El Niño
events (SOI >−2): 1982–83, 1991–92, and 1997–1998; and five
less evident (SOI: from −1.5 to −1.9) events during 1977–78,
1992–93, 1993–94, 1994–95, and 1998–99 (Fig. 1). La Niña (pos-
itive SOI) was evident during 1988–89, 1998–99, and 1999–2000.
There was no relationship between SOI and annual rainfall for the
1977–2003 period (Fig. 2; df 1,24 , R2 = 0.07, F = 1.83, P = 0.19).
The SOI was positively associated with rainfall during the wet sea- FIGURE 2. Relationship between annual, wet season and dry season rainfall
son (Fig. 2; df 1,24 , R2 = 0.40, F = 16.2, P = 0.0005). During the and Southern Oscillation Index (SOI) in the tropical dry forest of Chamela
El Niño years, rainfall was low in the wet season, while during La during the 1977–2003 period.
Niña years rainfall was high in this season. During the dry season,
the SOI was negatively associated with the rainfall (Fig. 2; df 1,24 ,
R2 = 0.28, F = 9.4, P = 0.005). In particular, during the El Niño et al. (1999) found a similar pattern when analyzing a 99-yr period
years (1982–83 and 1991–92) rainfall was high in the dry season, in the semiarid region of Chile, as did Marshal et al. (2002) in
while during La Niña years there was no rain in this season. the Sonoran desert for a period of 47 yr. Thus, ENSO events
Monthly rainfall distribution and plant species diversity esti- increase the variation of rainfall in the study region and have an
mates during the wet and dry seasons of the 1989–1993 period important consequence in the ecosystem function (Garcia-Oliva
appear in Figure 3. Species diversity was higher in the wet sea- et al. 2002).
son than in the dry season with the exception of 1992 (df 1,12 , Rain falling during the wet season controls vegetation growth
F = 30.9, P = 0.0001). Species diversity was similar in both forests and hence annual food production for large ungulates (Coe et al.
during the wet season, while in the dry season diversity was higher in 1976, Owen-Smith 1990, McKinney et al. 2001). The preliminary
the tropical semideciduous forest than in tropical dry forest (df 1,12 , evidence of this study suggests that similar to other sites (e.g., Ogutu
F = 5.22, P = 0.04). During the wet season diversity increased & Owen-Smith 2003), ENSO could have an additional effect on
slightly in the La Niña year 1989, while during the dry season plant species. This effect was evident during the 1992 dry season
the species diversity increased notably in the El Niño year 1992 when a record 649 mm rain fell in 15 d in January (Garcia-Oliva
(Fig. 3). et al. 2002). However, this effect depends on the month when
The analysis of 26 yr suggests a significant association between rain falls. For example, during the ENSO years 1983 and 1997,
ENSO and seasonal rainfall in the tropical dry forest of Chamela. 177 and <100 mm fell in May, respectively. Thus, considering
Unusually high rainfall during the dry season occurred when the the notable differences in floristic composition and life-forms, be-
SOI was strongly negative (El Niño years), whereas unusually high tween the tropical dry and semideciduous forest in the study area
rainfall during the wet season and no rainfall during the dry season (Lott et al. 1987), we could expect a complex relationship between
occurred when the SOI was strongly positive (La Niña years). Lima ENSO, precipitation, and plant responses, as has been shown in
698 Mandujano
FIGURE 3. Monthly rainfall from June 1989 to October 1993 and seasonal
variation in plant diversity (Shannon index, H ) of tropical dry forest (•) and
tropical semideciduous forest (◦).
others studies (e.g., Wright et al. 1999, Connell & Green 2000,
Williamson & Ickes 2002, Rolim et al. 2005).
During the dry season the response to hydric stress depends on
the uptake and control of water loss by plants (Reich & Borchert
1984). For example, in African savannah ecosystems the period of
retention of leaves into the dry season by deciduous trees and shrubs
is dependent upon the preceding rainfall (Owen-Smith 1990). At
the study site, leaffall normally occurs at the beginning of the dry
season, but leaves may remain longer in response to rain at the
beginning of the period (Bullock & Solis-Magallanes 1990). Thus,
in some years ENSO events increase rainfall during the dry sea-
son and the likelihood of plants staying leafy for a longer period
increases the food available to deer. ENSO conditions enhance
fruit production in tropical wet forests (e.g., Wright et al. 1999,
Williamson & Ickes 2002). Considering that white-tailed deer in-
creased the consumption of fruits during the dry season (Arceo et al.
2005), a possible additional effect of ENSO is the increase of food
availability. However, the temporal dynamics among rain, ENSO,
plant growth, and fruit production are complex and need further
monitoring.
ACKNOWLEDGMENTS
This study was supported by CONACYT and the Department
of Biodiversity and Animal Ecology of Institute of Ecology A. C.
Xalapa, Mexico. F. Garcia-Oliva, S. Gallina, V. Arroyo-Rodriguez,
H. Gomez de Silva, and two anonymous reviewers provided useful
comments on the paper.
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