DOI: 10.13102/sociobiology.v60i3.222-228Sociobiology 60(3): 222-228 (2013)
Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology
ISSN: 0361-6525
Ant Fauna on Cecropia pachystachya Trécul (Urticaceae) Trees in an Atlantic Forest Area,
Southeastern Brazil
PCJ Reis, WD DaRocha, LAD Falcão, TJ Guerra & FS Neves
Introduction
Cecropia genus comprises pioneer trees which are
characteristic elements of forest borders and gaps in the
Neotropics (Vasconcelos & Casimiro, 1997; Sposito & Santos,
2001). Most species are inhabited by mutualistic ants that
nest inside hollow internodes where they store eggs, larvae
and pupae (Harada & Benson, 1988), and feed on glycogen-
rich Müllerian bodies called trichilium located at the base of
leaf petioles (Yu & Davidson, 1997). Although Cecropia can
host a variety of resident ant genera, Azteca spp are the most
common ant inhabitants (Longino, 1991a). Usually only one
mature Azteca colony inhabits a Cecropia tree, however up
to five Azteca species may be found in a Cecropia population
(Longino, 1991a). In addition to a resident Azteca colony,
other ant species can be found living or foraging on the same
Cecropia tree (Vieira et al., 2010). However, the diversity
of non-Azteca ants living on Cecropia trees or using them as
foraging substrates is still poorly known, as well as the factors
that might affect ant richness and abundance on these trees.
Abstract
Cecropia are pioneer successional trees frequently associated with ants. Generally a
single dominant colony of Azteca ant inhabits each mature Cecropia tree, but other
ant species may be found living or foraging on the same tree. In this study, we asses-
sed the diversity of ant species on Cecropia pachystachya trees in two sites in the Bra-
zilian Atlantic Forest: a dust-free roadside and a dusty roadside. We also investigated
the influence of tree architecture on ant species richness. We found a total of 24 ant
species distributed in 11 genera and five subfamilies on C. pachystachya trees; 18 in
the dust-free roadside and 14 in the dusty roadside. We found up to five ant species
on a single tree, but only Azteca alfari was frequently encountered. Ant species rich-
ness per tree did not differ significantly between sites and was related to tree architec-
tural traits. On the other hand, ant species composition on trees differed significantly
between sites. Our study indicates that heavy dust deposition on Cecropia trees may
affect associated ant communities, not by changing ant species richness, but by cau-
sing different species to live and forage on trees under different dust exposure.
Sociobiology
An international journal on social insects
Universidade Federal de Minas Gerais (UFMG), Belo Horizonte, Minas Gerais, Brazil.
RESEARCH ARTICLE - ANTS
Article History
Edited by
Kleber Del-Claro, UFU, Brazil
Received 07 June 2013
Initial acceptance 29 June 2013
Final acceptance 25 July 2013
Keywords
Azteca, road dust, roadside,
plant architecture
Corresponding author
Paula Campos Junqueira Reis
Prog. de Pós-Graduação em Ecologia,
Cons. e Manejo da Vida Silvestre
Universidade Federal de Minas Gerais
Belo Horizonte, MG, Brazil, 36580-000
E-Mail: paulacjr@gmail.com
Complex tree architecture is an indicative of resource
diversity and availability, such as nesting sites, sites for ovipo-
sition, and food (Lawton, 1983). Therefore, tree architecture
might be an important factor affecting the diversity of ant
fauna associated to trees as reported for other insects (Hay-
som & Coulson, 1998; Espírito-Santo et al., 2007; Neves et
al., 2013). Cecropia trees are known for their great varia-
tion in size and architectural traits (Sposito & Santos, 2001),
but the way this variation affects the associated ants remains
unexplored.
Distribution of ants on Cecropia trees is also likely
influenced by external factors such as changes of environmental
conditions caused by human activities, which can directly or
indirectly affect both plants and ants. The Brazilian Atlantic
Forest range largely coincides with the most populated areas
of Brazil (Tabareli et al., 2005), thus this forest ecosystem suffers
with impacts of a dense road network. Roads are known to
have many ecological effects on surrounding biota (Spellerberg,
1998), mainly by altering landscape spatial patterns and proces-
ses and modifying abiotic and biotic conditions (Forman &
Sociobiology 60(3): 222-228 (2013) 223
Alexander, 1998; Trombulak & Frissell, 2000; Barbosa et al.,
2010). Besides, production of dust either by cars exhausts or
road surfaces may have important potential impacts on ve-
getation in roadsides including plant mortality (Spellerberg,
1998), especially on unpaved roads where dust production is
higher than on paved roads (Farmer, 1993). Dust deposition
on leaves may increase water loss, cause leaf injuries, also
leading to changes in associated invertebrate communities
(Farmer, 1993). Nevertheless, the effects of roadside dust on
ant fauna associated to Cecropia trees have never been studied
so far.
In this context, our aim was to assess the diversity of
ant fauna on Cecropia pachystachya Trécul (Urticaceae) trees
in a Brazilian Atlantic Forest area in Southeastern Brazil.
We evaluated ant species richness and composition on trees
located in two roadside forest sites: one where plants were
heavily covered by road dust and another where dust on
vegetation was negligible. We also evaluated the relationship
between ant species richness and the architectural traits of C.
pachystachya trees. We hypothesized that ant species richness
on trees should be positively related to higher complexity in
plant architecture and should be lower in plants disturbed by
heavy dust deposition. We also hypothesized that contrasting
dust deposition should affect the composition of ant fauna
associated to C. pachystachya trees.
Material and Methods
Study area
This study was conducted in the Rio Doce State Park
(Parque Estadual do Rio Doce-PERD hereafter) (19°40’S;
42°33’W) and its surroundings in Minas Gerais State, Southe-
astern Brazil (Fig 1). PERD is the largest continuous remnant
of Atlantic Forest in the state (c. 36,000 ha) (Instituto Estadual
de Florestas de Minas Gerais [IEF-MG], 2012) and thus has
a major importance to the regional conservation of this biome.
It is surrounded by agriculture, pasturelands, large areas
of Eucalyptus plantations, metallurgical plants, and a subs-
tantial road network (Brito et al., 1997). Cecropia trees are
abundant in early successional stages of forest both inside
and outside the park, and at the roadsides of unpaved roads
Cecropia trees are often heavily covered by dust (Fig 2).
Fig 1. Rio Doce State Park in Southeastern Brazil. The circle shows
the dust-free sampling area inside the park. Adapted from IBGE/
Brasil topographical map by Philippe Maillard - Instituto de Geoci-
ências-IGC, Universidade Federal de Minas Gerais.
Fig 2. Cecropia pachystachya leaves. A - Inside Rio Doce State Park, not covered by road dust. B - Covered by unpaved road dust in the
surroundings of the park.
PCJ Reis, WD da Rocha, LAD Falcão, TJ Guerra & FS Neves - Ant fauna on Cecropia pachystachya trees224
Sampling
Data were collected in July 2012 in two Atlantic Fo-
rest early successional sites: a dust-free roadside nearby Lake
Carioca inside PERD and a dusty roadside in the surroun-
dings of the park. Although both sites are roadsides, the one
inside PERD is located in a rarely used airplane landing field
while the road outside the park connects small municipalities
and is frequently used by cars. To sample the associated ant
fauna, we selected 25 mature Cecropia pachystachya trees
(Circumference at Breast Height (CBH) ≥ 20 cm) at least 30
m away from each other along a transect in each site. Ants
found on leaves and trunks were manually captured with the
assistance of a ladder and a pole pruner, and additional arbo-
real pitfall traps with sardine as bait were used to catch ants
that we did not capture manually (Bestelmeyer et al., 2000;
Ribas et al., 2003). Pitfalls were set up at approximately 1.50
m height in C. pachystachya trunks and were removed 48
hours later. In order to avoid that ground-living ants were
caught, we used grease in the trunks below pitfall traps. All
ants caught were taken to the Myrmecology Laboratory of
CEPEC/CEPLAC in Ilhéus, Bahia. In this laboratory, ant
species were identified after preparation and deposited in the
collection referred by CPDC acronym (#5687). For each C.
pachystachya tree we measured the following architectural
parameters: CBH, total height, number of branches and leaves,
and first branch height (FBH) following Sposito & Santos
(2001). We used ground-based methods of access (Von Matter
et al., 2010) because of the difficulty of sampling very tall
and fragile trees. Total height of trees was measured with a
12 m pole, and branches and leaves were counted by eye.
Statistical Analyses
In order to evaluate the effectiveness of our sampling
effort, we constructed rarefaction curves for each site using
Mao Tao observed richness based on 100 randomizations. A
generalized linear model (GLM) was constructed to inves-
tigate the effect of area (dusty and dust-free roadsides;
explanatory variable) on ant richness (response variable).
Difference in ant species composition between sites was
tested using Non-metric multidimensional scaling (NMDS)
with Raup-Crick index for incidence data and Analysis of
similarities (ANOSIM). ANOSIM provides a way to check
whether there is a significant difference between groups or
not (Clarke, 1993). Similarity percentage (SIMPER) analysis
was used to determine which ant species mostly contributed
to differentiation between groups (dust-free and dusty sites).
In order to determine which architectural variables (CBH, total
height, number of branches and leaves, or FBH) affected ant
species richness, we used Hierarchical Partitioning. Then,
generalized linear models (GLMs) were built to estimate the
effect of these pre-selected architectural variables (explana-
tory variables) on ant species richness (response variable).
Residual analyses were conducted to check data adequacy to
the probability distribution used as well as error distribution
(Crawley, 2007). Species rarefaction curves were constructed
on Estimates 8.0 software (Colwell, 2006); NMDS, ANOSIM
and SIMPER were carried out on PAST (Paleontological Sta-
tistics) version 2.15 (Hammer et al., 2001); Hierarchical par-
titioning and GLMs were performed using software R 2.6.2
(R Development Core Team, 2008).
Results and Discussion
A total of 24 ant species distributed in 11 genera and
five subfamilies were found on Cecropia pachystachya trees
(Table 1). The absence of stabilization of the rarefaction
curves (Fig 3) indicated that ant species richness associated
to C. pachystachya trees is probably underestimated in both
study sites. Therefore, additional studies are needed to deter-
mine more accurately the richness of ant species associated to
these trees.
Formicinae and Myrmicinae were the most representa-
tive subfamilies with eight and nine species respectively (71%
of all ant species found); and Camponotus Mayr, 1861 was
the most representative genus with seven species. Although
more than one Azteca species can inhabit a single popu-
lation of Cecropia trees (Longino, 1991a), we only found
Azteca alfari Emery, 1893 on the trees we analyzed. Azteca
alfari is known to inhabit Cecropia trees located in open and
disturbed habitats such as the early successional sites we stu-
died while other Azteca species such as Azteca constructor
and Azteca xanthochroa are more common in older forest
stages (Longino, 1991b). The presence of various non-Azteca
ant genera such as Camponotus Mayr, 1861, Crematogaster
Lund, 1831 and Pseudomyrmex Lund, 1831 (Table 1) on trees
of Cecropia was also reported by Longino (1991a) and their
presence indicates low aggressiveness of the Azteca resident
colony. As we observed on the field, some C. pachystachya
trees inhabited by A. alfari also housed other ant genera nests
(e.g. Cephalotes species). Indeed, A. alfari along with Azteca
ovaticeps Forel, 1904 are the least aggressive species among
all Cecropia-inhabiting Azteca ants (Longino, 1991a).
Eighteen ant species were found on C. pachystachya
trees in the dust-free site while 14 were found on the road
dust covered trees. Only eight ant species were common to
both sites, six species were found exclusively in the dusty ro-
adside and 10 were exclusive of the dust-free roadside inside
the park (Table 1). Azteca alfari Emery, 1893 and Camponotus
senex (Smith, F., 1858) were the most frequent species on C.
pachystachya trees in the dusty roadside, while more species
were frequent in the dust-free area - e.g. Camponotus balzani
Emery, 1894. Unlike the expected, each C. pachystachya
tree under heavy dust deposition held an average number
of ant species similar to each tree in the dust-free roadside
(dusty roadside: 2.12±0.24; dust-free roadside: 2.25±0.21;
Deviance1,47 = 0.094; p = 0.687).
Sociobiology 60(3): 222-228 (2013) 225
to CBH (Fig 5; p = 0.001). Low FBH and large CBH are indi-
catives of structural complexity, since low FBH trees usually
have long and dense crowns, and high CBH trees are gene-
rally large. Larger crowns and trees probably support higher
resource availability and higher diversity of conditions than
smaller trees (Campos et al., 2006) allowing more ant species
to live or forage on the same tree. In fact, other studies found
similar results of increase in ant and/or other insects’ richness
with increase in tree or crown size (e.g. Campos et al., 2006;
Costa et al., 2011; Neves et al., 2013).
Table 1. Ant species captured on Cecropia pachystachya trees in
Rio Doce State Park and surroundings, southeastern Brazil. Sampled
sites were a dusty roadside (in the surroundings of the park) and a
dust-free roadside (inside the park).
Fig 3. Rarefaction curves of ant species captured on Cecropia
pachystachya trees in the dusty roadside and in the dust-free road-
side in Rio Doce State Park and surroundings, southeastern Brazil.
Bars denote 95% CI.
Among the architectural variables evaluated, only
first branch height (FBH) and circumference at breast height
(CBH) were significantly related to ant species richness. Ant
species richness on trees ranged from zero to five, being nega-
tively related to FBH (Fig 4; p = 0.02) and positively related
Fig 4. Effect of first branch height (FBH) of Cecropia pachystachya
trees on ant species richness in Rio Doce State Park and surroundings,
southeastern Brazil.
Composition of ant species found on C. pachystachya trees
was significantly different between sampling sites (RANOSIM = 0.039;
p = 0.04; Fig. 6). SIMPER analysis revealed that Camponotus
senex (Smith, F., 1858), Crematogaster curvispinosa Mayr,
1862, Camponotus balzani Emery, 1894 and Azteca alfari
Emery, 1893 were the species that mostly contributed to
differentiation of groups and together explained 40.21% of
the dissimilarity between sites (Table 2). These ant genera
(Table 2) are commonly associated with myrmecophytes
and are classified as dominant arboreal ants (Brandão et al.,
2011). Therefore, ant fauna on C. pachystachya trees diffe-
red between sampling sites mainly due to dominant species
that live on the trees, while opportunistic ants had a smaller
contribution to differentiation between dusty and dust-free
roadsides. Camponotus senex is conspicuous and can occur
both in mature and disturbed forests, nesting in high regions
of canopy including dead branches of Cecropia trees (Lon-
gino, 2002). The greater frequency of C. senex in the dusty
roadside (Table 1) might be related to the abundance of dead
branches on C. pachystachya trees in this site, as observed
by us in the field. In turn, the abundance of dead branches
in the dusty roadside could possibly be caused by road dust
effects on C. pachystachya trees such as increased water loss.
The second species that most contributed to differentiation
between sites - Crematogaster curvispinosa (Myrmicinae) -
is a very common but inconspicuous ant that feeds on extra-
floral nectaries and inhabits initial secondary forests as well
as highly disturbed areas such as roadsides (Longino, 2003).
Subfamily/Species
Number of C. pachystachya
trees with given ant species
Dusty roadside
(n=25)
Dust-free roadside
(n=25)
Dolichoderinae
Azteca alfari Emery, 1893 24 22
Dolichoderus diversus Emery, 1894 1
Dolichoderus lutosus (Smith, F., 1858) 1
Ectatomminae
Ectatomma tuberculatum (Olivier, 1791) 3
Formicinae
Brachymyrmex sp.01 1
Camponotus balzani Emery, 1894 2 4
Camponotus cingulatus Mayr, 1862 1
Camponotus gp. Fastigatus sp.01 2
Camponotus renggeri Emery, 1894 3 3
Camponotus rufipes (Fabricius, 1775) 2
Camponotus senex (Smith, F., 1858) 9 3
Camponotus vittatus Forel, 1904 1
Myrmicinae
Atta rubripilosa Forel, 1908 1
Cephalotes goeldii (Forel, 1912) 1
Cephalotes minutus (Fabricius, 1804) 1
Cephalotes pusillus (Klug, 1824) 1 4
Crematogaster brasiliensis Mayr, 1878 1
Crematogaster curvispinosa Mayr, 1862 2 4
Nesomyrmex costatus (Emery, 1896) 1
Solenopsis sp.01 2 2
Solenopsis sp.02 1
Pseudomyrmecinae
Pseudomyrmex gracilis (Fabricius, 1804) 3 1
Pseudomyrmex oculatus (Smith, F., 1855) 1
Pseudomyrmex schuppi (Forel, 1901) 1
Total species 14 18
PCJ Reis, WD da Rocha, LAD Falcão, TJ Guerra & FS Neves - Ant fauna on Cecropia pachystachya trees226
It can tolerate other ant species nests nearby it (e.g. Cam-
ponotus and Cephalotes) and its colonies can occupy Cecropia
internodes that are located between resident Azteca nests
(Longino, 2003).
for instance, or it could indirectly affect ants through effects
on vegetation that generate avoidance of dusty trees by some
ant species and preference by others (e.g. dead branches for
Camponotus senex). Cecropia pachystachya trees under
heavy dust deposition may also have impaired development
and growth which may reduce the quantity and quality of
essential resources for resident and opportunistic ants. In
addition, dust may have effects on associate invertebrate
communities and these changes likely affect ant fauna espe-
cially through competition for resources. Finally, our results
indicated that effects of road dust might transcend impacts
on vegetation and might have important impacts on animal
associate communities especially ants.
Taxon
Av.
dissim.
Contrib. % Cumulative %
Mean
abund. 1
Mean
abund. 2
Camponotus senex 8.202 15.51 15.51 0.36 0.125
Crematogaster
curvispinosa
4.816 9.105 24.61 0.08 0.167
Camponotus balzani 4.205 7.949 32.56 0.08 0.167
Azteca alfari 4.047 7.651 40.21 0.96 0.917
Table 2. Similarity Percentage (SIMPER) analysis outcome. Av.
Dissim.: average dissimilarity of taxon between sites; Contrib. %:
relative contribution of taxon to dissimilarity between sites; Cumu-
lative %: cumulative contribution of taxon to dissimilarity between
sites; Mean adund. 1: mean abundance of taxon in the dusty roa-
dside; Mean abund. 2: mean abundance of taxon in the dust-free
roadside.
Despite its high tolerance in disturbed areas, C. curvis-
pinosa was more abundant inside the park possibly because of
the abundance of shrubs in the dust-free roadside in contrast
to the scarcity of these in dusty site. Abundant shrubs are indi-
catives of higher availability of extrafloral nectaries which are
important food sources to C. curvispinosa. Dust deposition
might limit the abundance of shrubs outside the park through
negative effects on plants as well as block access to the re-
maining extrafloral nectaries in the dusty roadside.
As shown by our results, composition of ant commu-
nities on Cecropia pachystachya trees differed between sites
with distinct exposure to dust. The substantial difference in
Fig 5. Effect of circumference at breast height (CBH) of Cecropia
pachystachya trees on ant species richness in Rio Doce State Park
and surroundings, southeastern Brazil.
Acknowledgments
This study was conducted during the Ecology Field
course fully supported by the graduate program in Ecolo-
gia, Conservação e Manejo da Vida Silvestre (UFMG) and
IEF-MG (Instituto Estadual de Florestas de Minas Gerais).
PCJR and WDDR were granted CNPq (Conselho Nacional
de Desenvolvimento Científico e Tecnológico) scholarships
and LADF was granted a CAPES (Coordenação de Aper-
feiçoamento de Pessoal de Nível Superior) scholarship. We
thank two other anonymous referees for their comments and
criticisms on the manuscript.
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