Mastozoología Neotropical, 21(2):263-274, Mendoza, 2014
Versión impresa ISSN 0327-9383
Versión on-line ISSN 1666-0536
Copyright ©SAREM, 2014
http://www.sarem.org.ar
Artículo
ASSEMBLAGE OF CHIROPTERANS IN A REMNANT
OF SEMIDECIDUOUS SEASONAL FOREST
IN SOUTHERN BRAZIL AND LATITUDINAL PATTERNS
OF SPECIES DIVERSITY IN THE ATLANTIC FOREST
Daniel Paulo de S. Pires1, Rosane V. Marques2, Tiago C. Cabral3,
Camila S. de Lima4, and Cristina V. Cademartori3
Conselho Regional de Biologia 03, Setor de Fiscalização, Coronel Corte Real, 662, Petrópolis, 90630-080 Porto Alegre,
RS, Brasil [correspondence: <[email protected]>].
2
Unidade de Assessoramento Ambiental, Divisão de Assessoramento Técnico, Procuradoria Geral de Justiça do Estado
do Rio Grande do Sul, Rua General Andrade Neves, 106 10° andar, 90210-210 Porto Alegre, RS, Brasil.
3
Laboratório de Conservação e Manejo da Biodiversidade, Pós-Graduação e Pesquisa, Unilasalle, Av. Victor Barreto,
2288, 92010-000 Canoas, RS, Brasil.
4
PPG em Biologia Animal, Depto. de Zoologia, Instituto de Biociências, Universidade Federal do Rio Grande do Sul,
Av. Bento Gonçalves, 9500, Prédio 43435, Agronomia, 91540-000 Porto Alegre, RS, Brasil.
1
ABSTRACT. Observed and expected richness, capture success, diversity, dominance, evenness and the effect of
latitude on the above variables at the regional level were evaluated with regard to the chiropteran fauna of a
semideciduous seasonal forest remnant in the state of Rio Grande do Sul, southern Brazil. Sampling occurred
between July 2008 and June 2010 using mist nets erected at ground level. So that chiropteran fauna could be
described and compared to other studies in the Atlantic Rainforest throughout the 10°–30°S using multivariate
regression. One hundred and sixteen specimens were captured, comprising 9 species and 3 families: Phyllostomidae, Vespertilionidae and Molossidae. Capture success was 0.0008 ind/m²h, with a diversity index (H’) of 1.623;
dominance (BP) of 0.4783; evenness (J’) of 0.7385, and expected richness of 11 species. Sturnira lilium had the
highest capture rate. There was a significant relationship between latitude and capture success and richness
(CS: R²=0.64, p=0.0005; S: R² = 0.28, p=0.0473). Distribution and abundance of bats at the regional scale was
related to latitude as well as the differentiation of assemblages in the Atlantic Rainforest.
RESUMO. Assembleia de quirópteros de um remanescente de Floresta Estacional Semidecidual do sul do
Brasil e gradientes latitudinais na diversidade de espécies. A riqueza observada e esperada, o sucesso de
captura, a diversidade, a dominância e a equitabilidade (evenness), bem como a influência da latitude sobre
estas variáveis em escala regional, foram avaliados, considerando-se a quiropterofauna de um remanescente
de Floresta Estacional Semidecidual no estado do Rio Grande do Sul, Brasil. As amostragens transcorreram de
julho de 2008 a junho de 2010 por meio de oito redes de neblina instaladas ao nível do solo. Assim, foi possível
descrever a quiropterofauna e compará-la, usando-se regressão multivariada, com outros estudos realizados na
Mata Atlântica ao longo de um gradiente de 10° a 30°S. Capturaram-se 116 indivíduos pertencentes a nove
espécies e três famílias, Phyllostomidae, Vespertilionidae e Molossidae. O sucesso de captura foi 0.0008 ind/ m²h.
O índice de diversidade (H’) foi 1.623, dominância (BP), 0.4783, evenness (J’), 0.7385, e a riqueza esperada compreendeu 11 espécies. Sturnira lilium registrou o maior número de capturas. Encontrou-se relação significativa
entre latitude e sucesso de captura e riqueza (SC: R²=0.64, p=0.0005; S: R² = 0.28, p=0.0473). A distribuição e
Recibido 4 marzo 2014. Aceptado 20 agosto 2014. Editor asociado: P Velazco
264 Mastozoología Neotropical, 21(2):263-274, Mendoza, 2014
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a abundância de morcegos, em escala regional, estão relacionadas com a latitude, assim como a diferenciação
das assembleias na Mata Atlântica.
Key words: Abundance. Atlantic Forest. Bats. Species richness.
Palavras-chave: Abundância. Mata Atlântica. Morcegos. Riqueza de espécies.
INTRODUCTION
Knowledge of richness and diversity of species
is a primary source of information for ecological and zoogeographic studies (Townsend et al.,
2006). Further, the study of organisms has been
shown to be a useful tool for the evaluation
of environmental change, whilst information
on diversity provides good indicators of environmental disturbances which, in turn, are
important in conservation biology (Primack
and Rodrigues, 2001).
Bats contribute significantly to the maintenance of forests since they have an important
role in the dispersal of seeds, pollination and
predation of insects (Helversen and Winter,
2003; Jones and Rydell, 2003; Patterson et
al., 2003; Fabián et al., 2008). In fact, the
chiropteran fauna has been shown to be very
suitable for the evaluation of environmental
modifications and as an indicator of its state
of conservation (Medellin et al., 2000; Mikich
and Bianconi, 2005).
Diversity and richness of bat species depend
on numerous factors, such as the habitat’s conservation degree and complexity, availability
of food resources, water or shelter, elevation,
latitude, stratification and seasonality (Kunz
and Kurta, 1988; Pedro and Taddei, 1997; Reis
et al., 2003; Esberard and Bergallo, 2005; Dias
et al., 2008; Kalko et al., 2008; Pacheco et al.,
2010). Examination of latitudinal gradients
on assemblages of chiropterans demonstrated
that species richness and diversity significantly
increased toward lower latitudes, while dominance and evenness did not show the same
trend (Patterson et al., 2003).
Although richness of species is actually affected by latitude gradients (Lyons and Willig,
1999), studies on bats do not include assem-
blages at or below 30°S (Stevens et al., 2006).
The effect of scale and its consequences is a
relevant issue in contemporary ecology while
it becomes more and more evident that results
obtained for a particular space or temporal
scale are not necessarily reproduced at bigger
or smaller scales (Lyons and Willig, 1999). Since
assemblages in southern Brazil do not feature
in publications on the theme, an evaluation that
includes the region and comprises scientific
observations at latitudes between 10°S and
30°S will help to understand whether global
patterns also apply at a regional scale. Empirical
research comprising latitudinal gradients and
diversity contributes towards the understanding
of a universal pattern and contexts in which
exceptions occur (Willig et al., 2003).
Various studies on bats in the forests of
Brazil’s southern region have been published
(e.g., Sipinski and Reis, 1995; Sekiama et al.,
2001; Bianconi et al., 2004; Arnone and Passos, 2007; Carvalho et al., 2009). However,
few have involved sampling longer than one
annual cycle (e.g., Marques and Fabián, 1994;
Fabián and Marques, 1996; Marques and Fabián, 2010; Ortêncio-Filho et al., 2010; Marques
and Fabián, 2011). The state of Rio Grande do
Sul, in particular, still lacks ecological studies
on bats, mainly with regard to their diversity
and species richness (Weber et al., 2011; Pires
and Fabián, 2013). Most studies are important
contributions on reproduction, behavior and
distribution of species (Marques and Fabián,
1994; Fabián and Marques, 1996; Rui and Fabián, 1997; Fabián et al., 1999; Marques and
Pacheco, 1999; Rui et al., 1999; Pacheco et al.,
2007; Weber et al., 2007; Bernardi et al., 2009)
or available ecological information from surveys
that cover all groups of mammals (Kasper et al.,
2007a; Kasper et al., 2007b; Penter et al., 2008;
LATITUDINAL GRADIENTS OF BATS IN ATLANTIC FOREST
Marques et al., 2011). Due to lack of studies
on bat species richness and diversity in the
semideciduous seasonal forest in Rio Grande do
Sul, the current study aims are 1) to determine
the composition of chiropteran species and
dominance relationships in a semideciduous
seasonal forest in Rio Grande do Sul; 2) to
evaluate richness and diversity of species of
the chiropteran fauna and to compare them
with surveys carried out in other forest areas;
3) to investigate whether latitude influences
capture success, richness, diversity, dominance
and evenness of species of bats in the Atlantic
Forest in Brazil.
MATERIAL AND METHODS
The study area comprises the Morro do Coco, a
remnant of semideciduous seasonal forest located
in the municipality of Viamão, Rio Grande do Sul
265
(30°16’15”S, 51°02’54”W), on the margins of Lago
Guaiba (Fig. 1). It is a region comprising private
estates without any status of legally protected area.
The region’s forest vegetation, although secondary,
has been restored and well preserved (Backes, 2000).
Region climate is Cfa type, according to Koppen’s
classification (C = Temperate climate or warm
temperate; f = humid climate without a marked
dry season and well distributed rainfall around the
year; a = hot summer). There are 4 well-defined
seasons throughout the year (Hasenack and Ferraro,
2006; Livi, 2006), with mean annual temperature at
19.5°C, mean temperature of the warmest month
above 22°C, minimum at 3°C in the winter and
maximum at 41°C in the summer. Mean annual
rainfall is approximately 1300 mm throughout the
year, featuring a photoperiod varying between about
10 h on June 21 to 14 h on December 21.
The current study was carried out during two
years, from July 2008 to June 2010, with monthly
expeditions of 3 consecutive nights. Seventy-two
Fig. 1. Localization of Morro do Coco, municipality of Viamão, state of Rio Grande do Sul, Brazil. Right upper corner:
the state of Rio Grande do Sul in Brazil; left upper corner: the circle indicates the study area in Rio Grande do Sul.
266 Mastozoología Neotropical, 21(2):263-274, Mendoza, 2014
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samplings were performed using eight 12 x 3 m
mist nets, placed at a height of 0.5–3.5 m from the
ground. Each net remained open for 7 h per night,
between 19:00 h and 02:00 h, at the edge of or inside
the forest, in clearings, close to fruiting trees and
close to houses, to capture insectivorous species that
potentially take shelter on roofs. Captured specimens
were placed in cotton sacks and later identified with
help of an identification key and specific literature
(Gardner, 2008; Reis et al., 2007; Barquez and Díaz,
2009). Voucher specimens were collected, fluidpreserved with the skull removed and deposited in
the collection of the Museu de Ciências Naturais La
Salle (authorization from SISBIO, #12094-2 of 2008).
Sampling effort was calculated by multiplying
total area of nets by exposure time and number of
repetitions, following Straube and Bianconi (2002).
Sampling effort totaled 145 152 m².h (72 m².h close
to houses). Capture success (CS) was calculated as
the ratio between number of captures and sampling
effort, also calculated by other studies undertaken
in Atlantic Forest of different Brazilian states (Faria,
2006; Althoff, 2007; Faria and Baumgarten, 2007;
Carvalho et al., 2009; Luz et al., 2009; Nobre et al.,
2009; Oprea et al., 2009; Ortêncio-Filho and Reis,
2009; Britto et al., 2010; Rocha et al., 2010; Perachi
et al., 2011; Weber et al., 2011; Pires and Fabián,
2013); the same occurred with species richness (S),
Shannon-Wiener diversity index (H’), Berger Parker
dominance index (BP) and evenness (J’), as estimated by Pielou’s ratio (Pielou 1969). Consequently,
comparisons may be made. These parameters were
obtained exclusively on the basis of total number of
specimens captured per species (absolute frequency)
without taking into consideration habitat distinctions
in species composition and abundance, for example,
to obtain the broadest possible estimate for each
locality. For comparative purposes, the only previous
studies utilized were those that met the following
two criteria: 1) they were conducted in the Atlantic
Forest and 2) mist nets were employed.
Sampling sufficiency was evaluated by species
accumulation curve (C.I. 95%). A second order
Jackknife estimation of species richness determined
expected number of species for the study area since
it provided a reliable assessment of species richness
regardless of sample size (Zahl, 1977).
Each species recorded was included in an ensemble, according to data of its diet obtained from
field observations and in line with the classification
proposed by Patterson et al. (2003).
Multivariate regression analysis tested influence of
latitude on biodiversity on a regional scale, where
the predictor variable (X axis) was latitude and the
response variables (Y axis) were capture success (CS),
richness (S), diversity (H’), dominance (BP) and
evenness (J’), according to Patterson et al. (2003).
Influence of latitude on each separate variable was
also tested by linear regression analysis. Response
is multivariate regression, or rather, y = (y1,…,yp),
and variables y1,…,yp are measured within the same
sampling unit and, as in the univariate case, associated to p sets of non-randomized predictor variables.
When parameters of p regressions are estimated as
a whole by multivariate regression, a gain in the
efficiency of estimators is obtained. However, this
analysis also verifies p of each regression separately.
Multivariate regression, linear regression, capture
success (CS), richness (S), diversity (H’), dominance
(BP) and evenness (J’) were calculated with Past
4.01 (Hammer et al., 2001) and species richness
was estimated with EstimateS 8.1 (Colwell, 2001).
RESULTS
A total of 116 specimens were captured including 9 species and 3 families (Table 1).
Capture success corresponded to 0.0008 bats/
m²h. When compared to other sites in Atlantic
Forest, capture success obtained at Morro do
Coco was always lower (Table 2).
The most represented family was Phyllostomidae, with 5 recorded species and 83%
Table 1
Species of bats captured with mist nets at the
Morro do Coco, municipality of Viamão, state of
Rio Grande do Sul, Brazil, from July 2008 to June
2010. N = Number of captures; E = Ensemble
(F, frugivorous; H, hematophagous; I, insectivorous;
N, nectarivorous).
Species
PHYLLOSTOMIDAE
Sturnira lilium
Artibeus fimbriatus
Artibeus lituratus
Glossophaga soricina
Desmodus rotundus
VESPERTILIONIDAE
Histiotus velatus
Myotis nigricans
MOLOSSIDAE
Molossus molossus
Molossus rufus
N
E
55
18
9
7
7
F
F
F
N
H
4
1
I
I
14
1
I
I
LATITUDINAL GRADIENTS OF BATS IN ATLANTIC FOREST
267
Table 2
Rates of capture success, richness, diversity, dominance, and evenness of chiropteran species in different
locations of the Brazilian forest formations of the Atlantic Forest. L = latitude (°S); CS = capture success; S
= richness; H’ = Shannon-Wiener diversity; BP = Berger Parker dominance; J’ = evenness.
Locality
L
CS
S
H’
BP
J’
Reference
Morro do Coco
30
0.0008
9
1.623
0.4783
0.7385
This paper
Morro São Pedro
30
0.0009
9
1.481
0.4860
0.6760
Pires and Fabián
(2013)
Morro do Elefante
29
0.0060
9
1.553
0.3421
0.7070
Weber et al. (2011)
Jabuticabeira and Morro Bonito
28
0.0095
9
1.75
0.3876
0.8088
Carvalho et al.
(2009)
Rancho do Mono and Terceira Vargem
27
0.0108
23
2.03
0.3144
0.6682
Althoff (2007)
Recanto Marista
23
0.0142
11
1.417
0.4427
0.6014
Britto et al. (2010)
Upper stretch of the Paraná River
22
0.006
17
1.988
0.2433
0.7015
Ortêncio-Filho and
Reis (2009)
Serra Negra
21
0.0111
15
1.773
0.336
0.6549
Nobre et al. (2009)
Praia das Neves
21
0.006
15
1.887
0.4124
0.6970
Luz et al. (2009)
Paulo Cesar Vinha State Park
20
0.0137
14
1.494
0.4097
0.5661
Oprea et al. (2009)
Reserva Natural do Vale
19
0.0138
12
1.668
0.5169
0.6777
Perachi et al.
(2011)
Una and Ilhéus
18
0.0275
44
2.025
0.4268
0.5351
Faria and
Baumgarten (2007)
Una
15
0.0215
39
2.026
0.367
0.5649
Faria (2006)
Campus da Universidade Federal de Sergipe
10
0.0711
18
2.038
0.240
0.7364
Rocha et al (2010)
of captures. Molossidae and Vespertilionidae,
although represented by 2 species each, differed
in number of captures. Sturnira lilium was the
dominant species (Table 1, Fig. 2).
By statistical extrapolation of probability
of occurrence for each species, the second
order Jackknife estimator of species richness
demonstrated that the curve did not reach an
asymptote but showed a slight trend towards
stabilization (C.I. 95% ± 0.54), estimating about
11 species for the study area (Fig. 3). Likewise,
since the species accumulation curve does not
show a complete asymptote, an increase in
sample size implies an increase in the number
of species (C.I. 95% ± 0.49). In fact, there was a
gradual increase in the number of species with
an increase in the number of captures, such
that, with about 86% of the night’s samples,
100% of the species found in this study were
already recorded. Diversity, calculated by the
diversity index (H’), was 1.623, dominance (BP)
0.4783, and evenness (J’) 0.7385.
The comparison of diversity gradients with
regard to latitude (Fig. 4) demonstrated that
latitude was significantly related to the gradients
as a whole (F = 8.94, R² = 0.289, p = 0.003). At
higher latitudes, capture success and richness
were significantly lower (SC: R²= 0.64, p=
0.0005; S: R² = 0.28, p= 0.0473) and there was
an increase in capture success and richness as
latitude decreased. In other words, there was
an inverse relation between capture success and
richness and latitude. Diversity, dominance and
evenness, however, were not significantly influenced by latitude (H’: R² = 0.27, p= 0.0520; BP:
R² =0.11, p = 0.23126; J’: R² = 0.14, p= 0.17).
268 Mastozoología Neotropical, 21(2):263-274, Mendoza, 2014
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50%
Percentage of captures
40%
Fig. 2. Percentages of captures of bats at the Morro
do Coco, municipality of
Viamão, Rio Grande do
Sul, Brazil, between July
2008 and June 2010.
30%
20%
10%
0%
Number of species
12
10
8
6
4
Jackknife 2
2
Observed richness
Fig. 3. Species accumulation curves of chiropterans captured from July
2008 to June 2010 at
the Morro do Coco, municipality of Viamão, Rio
Grande do Sul, Brazil.
0
1
11
21
31
41
51
61
71
Number of nights (samples)
DISCUSSION
The richness observed at the Morro do Coco
included 22.5% of the species recorded in Rio
Grande do Sul and about 48% of the species
expected for the hydrographic basin of Lago
Guaiba (Pacheco et al., 2007; Passos et al.,
2010). However, the collector’s curve suggested
that it was not possible to record all species expected for the study area (Fig. 3). In particular,
the families of insectivores (Vespertilionidae
and Molossidae), which according to Bernard
(2001) are more difficult to sample when mist
nets are employed, could show higher numbers
of recorded species with the utilization of other
sampling techniques. Nevertheless, the use of
mist nets was one of the criteria to compare
data from studies in the Atlantic Forest.
Similar studies on semideciduous seasonal
forests in southern Brazil reported a higher
number of species than that at the Morro do
Coco. Ortêncio-Filho and Reis (2009) found
17 species; Bianconi et al. (2004) demonstrated
the presence of 16 species; Ortêncio-Filho et
al. (2005) recorded 12 species; Gallo et al.
(2008) and Britto et al. (2010) reported 10
species each. Although the sampling effort at
the Morro do Coco was not low and the size
of the forest remnant reached approximately
140 ha, observed richness failed to attain the
minimum number of species registered for the
smallest areas already analyzed in the same
LATITUDINAL GRADIENTS OF BATS IN ATLANTIC FOREST
(a)
0,08
(b)
50
45
40
0,06
35
0,05
RICHNESS
CAPTURE SUCCESS
0,07
269
0,04
0,03
0,02
30
25
20
15
0,01
10
0
5
0
-0,01
0
3
6
9
0
12 15 18 21 24 27 30 33
3
6
9
12 15 18 21 24 27 30 33
LATITUDE
LATITUDE
SHANNON-WIENER DIVERSITY
2,06
1,96
1,86
1,76
1,66
1,56
1,46
1,36
0
3
6
9
12 15 18 21 24 27 30 33
0,5
0,45
0,4
0,35
0,3
0,25
0,2
0
3
6
9
12 15 18 21 24 27 30 33
LATITUDE
LATITUDE
(e)
0,87
(d)
0,55
BERGER PARKER DOMINANCE
(c)
2,16
0,82
Fig. 4. Relationship of latitude with (a) capture success;
(b) richness; (c) diversity; (d) dominance; and (e) evenness
in studies in the Atlantic Rainforest at latitudes 10°-30º S.
EVENNESS
0,77
0,72
0,67
0,62
0,57
0,52
0
3
6
9
12 15 18 21 24 27 30 33
LATITUDE
vegetation formation. However, Weber et al.
(2011), in a survey carried out in a deciduous
seasonal forest, also recorded 9 species, with
differences only in species composition and
number of captures. Pires and Fabián (2013)
reported the same species number and composition to those of the current study in their
research in a seasonal semideciduous forest
close to the Morro do Coco (approximately
10 km). In comparison to similar environments
of other Brazilian regions, lower species richness in Rio Grande do Sul was expected since
the state has the lowest number of bat species
in southern Brazil, especially phyllostomids
(Passos et al., 2010).
The predominance of phyllostomids appears
to be consistent across Neotropical studies. In
fact, it is the family with the greatest richness
and abundance reported in studies carried out
in southern Brazil (Bianconi et al., 2004; Carvalho et al., 2009; Weber et al., 2011). The fact that
most phyllostomids captured were frugivorous
may be due to the sampling method, with nets
270 Mastozoología Neotropical, 21(2):263-274, Mendoza, 2014
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set between 0.5 m and 3 m above the ground,
that is, the height of most fruit trees (Fabián
et al., 2008). The abundance and richness of
species of the families Vespertilionidae and
Molossidae, observed at the Morro do Coco,
did not differ much from other studies in the
southern region of Brazil, where it seemed that
the variety of the number of species depended
on the type of area and method employed in
the study (Reis et al., 2000; Sekiama et al., 2001;
Esberard, 2003; Weber et al., 2011).
Despite the high sampling effort, capture
success was very low when compared to that
registered in other studies. From a biogeographic perspective, the study area is located
in the mid-latitude subtropical region and in
the southern boundary of phyllostomid species.
Phyllostomidae is a tropical and subtropical
family with limited thermo-regulating requirements and physiological restrictions (Willig and
Bloch, 2006). Thus, the possibility that capture
success was low might not be discarded due
to the small population size of the species of
this family in the region. In fact, it is common
that populations are small at their distributional
limits (Begon et al., 2007). An inverse relation
between capture success and latitude, such as
that found on a regional scale, supports this
idea.
The latitudinal gradient of species richness
and diversity of bats in the tropical region is a
known pattern, while dominance and evenness
do not show a correlation with latitude (Patterson et al., 2003). The evaluation of studies
carried out in the Atlantic Forest, between
15° and 30° of latitude south, confirmed, on a
regional scale, the pattern expected for species
richness, but not for diversity, which varied
regardless of latitude, even though results
were only marginally significant. Dominance
and evenness, on a regional scale, also varied
regardless of latitude. Empirical evidence shows
that diversity measures vary neither together
nor identically across latitudes and comparisons
of biodiversity, weighed by species abundance,
may be impaired if not based on identical bases
(Willig et al., 2003). According to Patterson
et al. (2003), temperature, rainfall and insolation are the latitudinal factors that influence
variations in assemblages of tropical bats and
DPS Pires et al.
their capture success. Richness determinants
for New World bat species are defined as a
set of energy availability, environmental and
seasonal heterogeneity, followed by energy and
seasonality and, lastly, heterogeneity with one
of the variables mentioned above (Tello and
Stevens, 2010). The species richness of bats in
the Atlantic Forest primarily varies as a result
of temperature variations (Stevens, 2013). The
scarcity of food supplies and low temperatures
can cause torpor during the winter, which
limits the number of species able to withstand
this stress factor. Tropical ecosystems receive
more sunshine and show seasonality according
to the rainy or dry periods; in a special way,
forests have higher heterogeneity in habitats
and resources. Less species richness is thus
expected in subtropical environments even
though one should not discard the fact that
species evolution and distribution have been
affected by climate and history throughout
millions of years (Stevens, 2011).
In spite of lower capture success, richness and
diversity of species at the Morro do Coco were
equivalent or higher to those in other areas of
the Atlantic Rainforest (Table 2). Bat assemblage at the Morro do Coco generally revealed
higher dominance and evenness rates (Table 2).
Regression analysis between latitude and richness showed that an inverse relationship exists
among these components, with an absence of
association between latitude and diversity, or
rather, different from that expected. Neither
evenness nor dominance exhibits any systematic
variation linked with latitude. At lower latitudes
the rates of increase in the number of rare species outstrip those of common species, with a
greater impact on richness than on evenness,
which varies less as a result (Stevens and Willig,
2002). The arbitrariness and limitations of the
composition of diversity indexes by richness
and evenness, to which different weights are
given, have been pointed out by Melo (2008). In
the case of diversity, the index obtained in this
study was higher than that found by Bianconi
et al. (2004) in fragments of a semideciduous
seasonal forest in the state of Paraná, equal to
1.38, and by Weber et al. (2011) in a fragment
of a deciduous seasonal forest in Rio Grande
do Sul, equal to 1.55.
LATITUDINAL GRADIENTS OF BATS IN ATLANTIC FOREST
The current study showed that S. lilium was
the species with the highest number of records
which overwhelmed the next species by 100%.
Results are not surprising, since the species
is common in places with a greater degree of
anthropogenic influence where the more generalist species tend to increase considerably their
density (Bianconi et al., 2004; Reis et al., 2006).
Although the results of this investigation differ
from those carried out in more preserved areas
(Rex et al., 2008; Dias et al., 2008), evenness
demonstrates that the species are distributed in
a reasonably even way within the community.
Inventories of bats in Brazil usually show
some dominant species coexisting with rarer
ones (Esberard, 2003; Bianconi et al., 2004; Dias
et al., 2008). The locally common S. lilium was
the species with the highest number of captures.
This fact coincides with other investigations in
the state of Rio Grande do Sul (Weber et al.,
2011, Pires and Fabián, 2013) and indicates
that it is also one of the commonest species in
inventories in the state. Artibeus lituratus and
A. fimbriatus also had higher capture frequency
when compared to that of other species. The
3 taxa together comprised 72% of records
known for their high frequency in inventories
in the southern region of Brazil (Bianconi et
al., 2004; Weber et al., 2011). The presence of
these species may be strictly related to their
feeding preferences for Piperaceae, Solanaceae
and Moraceae, respectively (Fabián et al., 2008)
plant species with known occurrence at the
Morro do Coco (Backes, 1999).
All taxa of Vespertilionidae at the Morro do
Coco were considered rare in the sample, as in
other studies conducted in southern Brazil, in
which vespertilionids are few in number, sometimes constituting the second most represented
family in forest surveys that utilize mist nets
(Dias et al., 2008; Carvalho et al., 2009; Britto
et al., 2010; Weber et al., 2011).
The 15 specimens of Molossus molossus captured as well as the only specimen of M. rufus
were trapped with nets set in front of the exit
of a tile-roofed house, a strategy that increased
the probability of sampling the Molossidae
(Esberard, 2003). Despite the low sampling
effort employed for the capture of molossid
bats, M. molossus was the third species with
271
the highest number of specimens. The facts
demonstrated the importance of these animals
in the assemblage of chiropterans in the region.
The species is considered unusual in urban
and periurban environments in Rio Grande
do Sul (Pacheco et al., 2010), especially when
compared to Tadarida brasiliensis which was
not recorded at the Morro do Coco.
There was no capture of piscivorous bats, but
at least one specimen was sighted flying over a
small flooded area between the field and forest
fragment in the study area. Moreover, there is
a record of an occurrence of Noctilio leporinus
in the region (Marques and Pacheco, 1999).
Besides the low number of captures compared
to the high sampling effort, all the ensembles
occupied by chiropterans in the region were
represented in this study. The presence of
frugivorous, nectarivorous, hematophagous,
low-flying insectivorous (vespertilionids),
high-flying insectivorous (molossids) and
piscivorous bats accounted for the 6 different
types of ensembles and demonstrated that,
despite low species richness and capture success, the trophic diversity resembled that of
tropical environments. Comparisons between
bat assemblages in different environments (the
wet tropical forests of Central America and the
Amazon and more open formations such as
the Caatinga and the Cerrado) indicated up to
7 guilds (Stevens and Willig, 2000), although
featuring higher species richness than that at
the Morro do Coco. In spite of low species
richness, 6 trophic groups were reported in
the area under analysis.
The current study surveyed a representative
portion of the fauna of bats for the region.
Since inventories represent the starting point
for ecological studies of communities, many
aspects of the chiropteran fauna still need to
be investigated. Densities, temporal population
fluctuations, roosts, and reproductive and activity patterns may be elucidated in the future with
the application of suitable methods, for a better
understanding of the partitioning of resources
and coexistence of species in the subtropics.
As demonstrated in the current analysis,
latitude is related to the distribution and
abundance of bats on a regional scale too and
thus contributes towards the differentiation of
272 Mastozoología Neotropical, 21(2):263-274, Mendoza, 2014
http://www.sarem.org.ar
assemblages, albeit within the same biome of
Atlantic Rainforest. The proportionally inverse
relationship between latitude, richness and capture success may explain the low number of bat
captures often reported for Rio Grande do Sul.
ACKNOWLEDGMENTS
The authors would like to thank the team that helped
them with field work. Thanks are also due to the Lasalle
Brothers for allowing the authors to conduct the study
on their estate. They are also grateful to the Laboratório
de Geoprocessamento do Unilasalle, especially Diego Medeiros, for the preparation of the map of the area. Thanks
are also due to CNPq for its financial support.
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