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Diversity and Temporal

Variation of Ants
(Hymenoptera:Formicidae)
From Malaise Traps in a
Tropical Deciduous Forest

Article in Sociobiology · January 2009

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 633
Diversity and Temporal Variation of Ants (Hymenoptera:
Formicidae) From Malaise Traps in a Tropical Deciduous Forest
by
Gabriela Castaño-Meneses , Betty Benrey2 & José G. Palacios Vargas3
1

Abstract
The ant community from the shrub layer in a tropical deciduous forest was
sampled for two years with Malaise traps. A total of 1,256 ants belonging to 48
species and 21 genera were collected. The most abundant species were Eciton
burchelli, Crematogaster brevispinosa and Pheidole sp. 1, which represented
more than 60% of the total. A time series analysis was performed to evaluate
the seasonality in the ant abundance. This showed a cyclical pattern, with a
displacement around 15 days between the first rain and the abundance peak
of ants. A significant positive correlation was found between precipitation
and ant abundance, while the temperature did not have a significant relation-
ship. The use of Malaise traps is useful to characterize the structure of the
community of ants in the shrub layer.
Key words: Canopy, collecting methods, diversity, seasonality, shrub layer,
soil, time series analysis.
INTRODUCTION
Various estimates of biodiversity have shown that arthropods comprise a
principle portion of the earth's animals (Erwin 1982; Stork 1993; Odegaard
2000). In addition, understanding the spatial and temporal variations in the dis-
tribution patterns of organisms is very important in order to find differences in
the structures of local faunas (Gaston & Lawton 1988; Frith & Frith 1990).
Due to their rapid response to environmental changes, ants, along with other
invertebrate groups, are very important elements in biodiversity studies (Agosti
1
Ecología y Sistemática de Microartrópodos, Facultad de Ciencias, Universidad Nacional Autónoma
de México, Ciudad Universitaria, Coyoacán 04510, México, D. F. México, e-mail: gcm@hp.fciencias.
una.mx
2
Université de Neuchâtel, Institut de Zoologie, LEAE, 11 rue Emile-Argand, case postale 2, CH-2007
Neuchâtel, Suisse, e-mail : betty.benrey@unine.ch
3
Ecología y Sistemática de Microartrópodos, Facultad de Ciencias, Universidad Nacional Autónoma
de México, Ciudad Universitaria, Coyoacán 04510, México, D. F. México e-mail: troglolaphysa@
hotmail.com
634 Sociobiology Vol. 54, No. 2, 2009

et al. 2000). In tropical areas ants are the dominant group in soil and canopy
(Erwin 1983; Kaspari 1996; Watt et al. 1997; Basset 2001). Ants support the
local species richness significantly and have many functional roles essential to
their ecosystems (Folgarait 1998).
For collecting ants, many diverse methods, like the use of pitfall traps, as-
pirators, webs, Berlese funnels, fogging and litter sieves have been used, and
in order to obtain better results a combination of two or more methods are
usually applied (Romero & Jaffe 1989; Majer & Delabie 1994; Longino et
al. 2002). Malaise traps are usually used for flying insects; nevertheless, these
traps provide knowledge about the general composition and insect commu-
nity structure in a specific place (Hansen 1988; Brigham & Saunders 1990).
In large sampling areas, Malaise traps are very efficient for their capacity to
collect a great number of tourist and wandering species (Clark & Samways
1997). This method collects specimens from soil, shrub layers and also canopy
species (Longino & Colwell 1997), and is very useful for studying the spatial
and temporal variation in arthropod communities (Knizeski & Sullivan 1984;
Hutcheson & Jones 1999; Kaspari et al. 2001).
In the following work we study the ant community in a tropical deciduous
forest focusing on the spatial and seasonal changes in its structure.
Material and Methods
Study area
The Chamela Biology Station is located in the coast of Jalisco State, Mexico
(19oC 30'N, 105o 03'W). The station is larger than 3,300 ha in area, covered
mainly by tropical deciduous forest (Bullock 1988). There are more than 780
botanical species, with Leguminosae and Euphorbiacea the most diverse families
(Lott et al. 1987). The soils are entisols, sandy, with neutral pH and low organic
matter content (Solís 1993).
The weather of Chamela is Aw(x)i, according to the Köppen classification
modified by García (1981).This is one of the driest of the sub-humid hot
types, with rain in summer and low thermal variations. The annual average
precipitation is 707 mm, and more than 80% of the rain falls between July
and October. The temperature is nearly constant, with maximum monthly
averages of 28.8 to 32.2oC during May to July, and minimum averages of
15.9 to 22.6oC from December to February (Bullock 1988). Fig. 1 shows
 Meneses, G.C. et al. — Diversity of Ants in a Tropical Deciduous Forest 635

the precipitation and temperature values recorded during the study time
(August 1991- July 1993).
The sampling area is located in a hydrological watershed system in the
occidental slope of the Cerro Colorado hill. The system comprises five wa-
tersheds ranking from 10 to 30 ha in area. The sampling sites were into the
watersheds named 1 and 4 (Cervantes et al. 1988). Those watersheds are very
similar in size (12 ha) and average litter production (7,642 kg ha-1, annual
average, following Patiño, 1990) according to the studies about ecosystem
function development in Chamela (Martínez-Yrizar & Sarhukán 1990;
Patiño 1990). In addition, sampling was done in a watershed located to the
north of the watershed 4, named 4A. That watershed, with approximately
9 ha in size, had been used for corn crops, but that agricultural practice was
abandoned about 30 years ago, which allowed for the regeneration of the
original vegetation.
Sampling
We made a biweekly sampling using Malaise traps (Townes 1962), for

Fig. 1. Precipitation and temperature recorded during the study in the Chamela Biological Station,
Jalisco, Mexico.
636 Sociobiology Vol. 54, No. 2, 2009

two years, from August 1991 to July 1993. During the first year (August
1991 to July 1992) we collected samples in watersheds 1 and 4, and in the
second year (August 1992 to July 1993) we sampled in 1 and 4A. Five traps
were located in each watershed by random selection. The traps were active
for one week at 15-day intervals. No attractant was used, only 80% alcohol
to preserve the specimens collected. The specimens were quantified and
isolated by taxonomic groups. Ants were identified by genus and isolated
in morphospecies. An estimation of Shannon diversity index, evenness and
species richness was performed.
Statistical analysis
The effect of watershed and collection date on ant abundance was evalu-
ated by a two way analysis of variance (ANOVA). The association degree
between the ant abundance and precipitation variations during the study
were estimated by time series analysis, according to Pearson & Derr (1986).
In order to verify the cycling or randomness of the distribution we used a
significance test according to the tables proposed by Anderson (Yamane
1979). The time series were measures of the ant abundance along with the
biweekly accumulated precipitation. The time series models were selected by
the Auto Regressive Integrated Moving Average (ARIMA). The original data
were expressed as the difference between the observed data and the values
predicted by the ARIMA model. The correlations between the time series
of the abundance during the biweekly t and the time series of the precipita-
tion during the biweekly t-1 were estimated. The correlation coefficient was
used to evaluate the association between the abundance and precipitation
fluctuations during the study time.
Results
General Faunistic Analysis
During the two years of sampling, we collected 1,256 ants belonging to
48 species and 21 genera (Table 1). The most abundant species were Eciton
burchelli (43.4%), followed by Crematogaster brevispinosa (11.4%) and
Pheidole sp. 1 (6.6%).
According to the ANOVA results, there are no significant differences in
the ant abundance of watersheds 1 and 4 (F1, 228=1.12; p>0.05), nor in 1 and
4A (F1, 228=0.02; p>0.05).
 Meneses, G.C. et al. — Diversity of Ants in a Tropical Deciduous Forest 637

Table 1. Ant abundance collected by Malaise traps in Chamela Biological Station, Jalisco, Mexico.
Data from watershed 1 included collections from 1991 and 1992. In watershed collecting was
only performed during 1991, and in watershed 4A only in 1992.

TAXA Watershed 1 Watershed 4 Watershed 4A

PONERINAE
Ectatomma ruidum (Roger) 21 1
Odontomachus sp. 1 1
Pachycondyla sp. 3
Platythyrea sp. 3 1
ECITONINAE
Eciton burchelli (Westwood) 380 133 32
Labidus coecus (Latreille) 1
Neivamyrmex chamelensis Watkins 2

PSEUDOMYRMECINAE
Pseudomyrmex sp. 1 4 2 1
P. sp. 2 2 1
P. sp. 3 1
P. sp. 5 16 11 2
P. sp. 6 1
P. sp. 7 3 10 4
MYRMICINAE
Acromyrmex sp. 1 1 1
Atta mexicana (Smith) 17 6
Crematogaster brevispinosa Mayr 84 53 6
C. sumichrasti Mayr 4 2 1
Cyphomyrmex sp. 6 5 2
Temnothorax sp. 1 5 7
T. sp. 2 3 1
Pheidole sp. 1 34 53 5
Ph. sp. 2 1 15
Ph. sp. 3 1
Ph. sp. 4 9 15 1
Ph. sp. 5 23
Ph. sp. 6 22
Ph. sp. 7 1
Solenopsis sp. 1 16 60 1
Strumigenys sp. 1 1 1
Trachymyrmex sp. 1
Cephalotes sp. 1 2
C. sp. 3 2 4
C. sp. 4 26 14 3
C. sp. 5 1
C. sp. 6 2 21

DOLICHODERINAE
Forelius sp. 1 4 2
Tapinoma sp. 3 3 3
638 Sociobiology Vol. 54, No. 2, 2009

Table 1. Ant abundance collected by Malaise traps in Chamela Biological Station, Jalisco, Mexico.
Data from watershed 1 included collections from 1991 and 1992. In watershed collecting was
only performed during 1991, and in watershed 4A only in 1992 (continued).
TAXA Watershed 1 Watershed 4 Watershed 4A
FORMICINAE
Camponotus sp. 1 16 1
Cm. sp. 2 18 6 3
Cm. sp. 3 7
Cm. sp. 4 2 2
Cm. sp. 5 1
Cm. sp. 7 1
Cm. sp. 8 1 8 1
Cm. sp. 9 1 1 1
Cm. sp. 11 1
Cm. sp. 14 9 3
Cm. sp. 17 1

Table 2. Parameters of ant community sampling by Malaise traps in the tropical deciduous forest in
Chamela, Jalisco, Mexico. S=Species richness; A=Abundance; H’=Shannon diversity index; J’=
Pielou’s evenness.

Watershed S A H’ J’
Watershed 1 (91-92) 30 438 1.81 0.53
Watershed 4 (91-92) 35 461 2.51 0.71
Watershed 1 (92-93) 35 266 1.95 0.55
Watershed 4A (92-93) 24 91 2.41 0.76
General 48 1256 2.32 0.60

The highest value of species richness was recorded in watershed 1 during

the second year and in watershed 4, with 35 species, while the highest diversity
was found in general in watersheds 4 and 4A (Table 2).
The genus with the most species was Camponotus (11 species), followed by
Pheidole (6 species). The species richness recorded in the Malaise traps is high,
considering that this method is not usually recommended for ant collection.
Nevertheless, this method allows us to find species with arboricolous nests
and soil foraging habits, such as Crematogaster sumichrasti, some species of
Camponotus and Cephalotes, and those with preferences for the shrub layer,
like Platythyrea.
Temporal Variation
The species richness and Shannon diversity index values had variations
along the study period (Fig. 2). In general, the diversity values in watershed
1 are above the values of 4 and 4A. The species richness monthly average for
 Meneses, G.C. et al. — Diversity of Ants in a Tropical Deciduous Forest 639

watershed 1 during the first year was 8.1±4.5, while in the second year was
6.6±1.8. The average for the second year was 7.3±3.4. In watershed 4 the
species richness monthly average was 8.2±3.0, and in watershed 4A it was
3.3±3.0.
The distribution pattern of variables, according to the table proposed by
Anderson, was cyclical (p>0.01). Fig. 3 shows the correlation between the
abundance and precipitation from the time series analysis. Fig. 4 shows the

Fig. 2. Temporal variation of ant Species Richness (A) and Diversity (B) collected by Malaise traps in
the Chamela Biological Station, Jalisco, Mexico.
640 Sociobiology Vol. 54, No. 2, 2009

Fig. 3. Correlation between ant abundance in Malaise traps and biweekly accumulated precipitation in
Chamela Biological Station, Jalisco, Mexico.

Fig. 4. Time series analysis by ARIMA model between ant abundance in Malaise traps and biweekly
accumulated precipitation in Chamela Biological Station, Jalisco, Mexico.
 Meneses, G.C. et al. — Diversity of Ants in a Tropical Deciduous Forest 641

ARIMA model obtained from the time series analysis for the two variables,
and we can see a diphase between the precipitation and abundance peaks in
some points. We found a positive and significant correlation between the
ant abundance and precipitation during the biweekly t-1 (r460=0.25; p<0.05).
We did not identify a significant correlation with temperature (r460=0.04;
p>0.05).
Trophic guilds
The ants collected by Malaise traps belonged to six trophic guilds, in order of
importance: omnivorous, predator, granivorous, detritophagous, herbivorous
and nectarivorous, as shown in Fig. 5. The best represented trophic guilds
in the three watersheds were the omnivores and the predators. In general,
watershed 4A shows the lowest diversity of trophic guilds. During the rainy
season ( July- October), we found the highest diversity of trophic guilds, while
in the dry season the dominant group was the omnivores (Fig. 6).
DISCUSSION
Malaise traps are a very useful tool in the study of insect communities.
Research in homogeneous habitats for beetles shows that only one Malaise
trap can represent the main characteristics of the community (Hutchesen

Fig. 5. Distribution of trophic guilds of ants collected by Malaise traps in Chamela Biological Station,
Jalisco, Mexico. N=48 species.
642 Sociobiology Vol. 54, No. 2, 2009

& Jones 1999). The ant diversity recorded by Malaise traps is high compared
with those found in other habitats, like soil and canopy in the same study area
(Castaño-Meneses 1997). In the shrub layer it is possible to find species from the
soil and litter as well as canopy, because it is a transitional habitat for those ant
species due their foraging habits, adding to species that prefer the shrub layer.
In relation to the spatial distribution of the ants, we did not find significant
differences between watersheds in abundance, but the composition is different
in each one. There are more species shared between watersheds 1 and 4 (24 spe-
cies; Table 1), than 1 and 4A (19 species), probably as result of the modification
in the original vegetation in the watershed 4A.
The time series analysis is useful to understand and model the population
variations, allowing the best interpretation of the observed data in the field

Fig. 6. Temporal variation of trophic guilds of ant species collected by Malaise trap at Chamela Biological
Station, Jalisco, Mexico.
 Meneses, G.C. et al. — Diversity of Ants in a Tropical Deciduous Forest 643

(BjØrnstad & Grenfell 2001). The observed displacement between ant abundance
and precipitation can be explained because the arthropod populations do not
have a rapid response to increased rain amounts; they take approximately three
weeks from the beginning of the rain to their increase in abundance (Pearson
& Derr 1986). In our research we found that the difference is shown on the
15th day.
Temporal distribution of trophic guilds is related to resource availability,
as are mutualistic and predatory interactions that ants establish with other
organisms, whose presence can be very seasonal (Cushman & Whitham 1989;
Rico-Gray 1993).
Our results show that the use of Malaise traps to collect ants can be of great value
in studies of the community structure of epiedaphic and shrub layer ants.
ACKNOWLEDGMENTS
The present research was supported by Project IN2078/91 DGAPA-UNAM.
The field work was developed with the help of M Sc. Alicia Rodríguez Palafox.
Drs. Zenón Cano Santana, Alfonso Pescador Rubio, Francisco Villalobos,
Norma García Calderón and Victor Rico-Gray gave invaluable suggestions on
the manuscript. Lawrence Ross reviewed the spelling of the manuscript and Luis
Parra (ENAP-UNAM) made valuable grammatical and style corrections.
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