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Austral Ecology (2007) 32, 254–260 doi:10.1111/j.1442-9993.2007.01682.x

Ecological parameters of the leaf-litter frog community of an

Atlantic Rainforest area at Ilha Grande, Rio de Janeiro state,
Brazil
M. Van SLUYS,* D. VRCIBRADIC, MAS ALVES, HG BERGALLO AND CFD ROCHA Department of
Ecology, Roberto
Alcantara Gomes Institute of Biology, State University of Rio de Janeiro, Rua São Francisco Xavier 524, CEP 20550-013,
Rio de Janeiro Janeiro, RJ, Brazil (Email: vansluys@uerj.br)

Abstract The Atlantic Rainforest originally covered much of the Brazilian coast and is now reduced to approximately
only 7% of its original area. Data on abundance distribution and microhabitat characteristics of anuran amphibians living
on the forest floor leaf litter in the Atlantic Rainforest are scarce. In this study, we analyzed the effect of litter depth and
structure on the abundance and species richness of leaf-litter frogs in an area of Atlantic Rainforest at Ilha Grande, Rio
de Janeiro State, south-eastern Brazil. We performed monthly samples (nocturnal and diurnal) from August 1996 to
October 1997 using small (2 m ¥ 1 m) plots. We sampled 234 plots, totaling 468 m2 of forest leaf litter. We estimated leaf-
litter depth and the proportion of leaves in the plot and tested their effect on the total abundance of frogs and species
richness using multiple regression analysis. We found 185 frogs from eight species: Brachycephalus (=Psyllophryne)
didactylus (Izecksohn, 1971) (Brachycephalidae), Dendrophrynis cus brevipollicatus Jiménez de la Espada 1871
(Bufonidae), Adenomera marmorata Steindachner 1867, Eleutherodac tylus parvus (Girard 1853), Eleutherodactylus
guentheri (Stei ndachner 1864), Eleutherodactylus binotatus (Spix 1824) and Zachaenus parvulus (Girard 1853)
(Leptodactylidae), and Chiasmocleis sp. (Microhylidae). Brachycephalus didactylus was the most abundant species, with
91 individuals, whereas Dendrophryniscus brevipollicatus was the rarest, with two individuals. Mean litter depth and the
proportion of leaves in the leaf litter were significantly related to frog abundance (R2 = 0.17; F2.107 = 10.779; P = 0.0001)
and species richness (R2 = 0.11; F2.107 = 6.375; P = 0.002 ) indicating that microhabitat characteristics may affect local
distribution and abundance of frogs in the forest floor.

Keywords: abundance, frog community, leaf-litter depth, microhabitat, species richness.

INTRODUCTION having less rich litter frog communities than other areas
(Watanabe et al. 2005).
Amphibians are important constituents of leaf-litter Most estimates of litter frog density and diversity in
communities in tropical forests (eg Fauth et al. 1989; tropical forest areas have been based on data gathered
Allmon 1991; Vitt & Caldwell 1994). In general, tropical using plot methods (eg Allmon 1991; Jaeger & Inger
leaf-litter amphibians are more abundant in the NewWorld 1994; Giaretta et al. 1997, 1999; Rocha et al. 2001 ;
tropics than in the OldWorld (Inger 1980; Duellman 1988; Watanabe et al . 2005 ). Estimates of litter frog densities
Fauth et al. 1989; Allmon 1991) although the reason for have been obtained in tropical forest areas of different
this difference is unclear (Huang & Hou 2004) . Several continents, and have indicated significant between-site
studies have dealt with patterns of abundance of leaf- differences in abundance and diversity of litter frogs (eg
litter amphibians, and litter frog communities from tropical Brown & Alcala 1961; Lloyd et al .
regions are relatively well known (Inger 1980; Toft 1968; Scott 1976, 1982; Inger 1980; Allmon 1991;
1980a; Fauth et al. Watanabe et al. 2005). The differences in frog densities
1989; Allmon 1991; Rodriguez 1992; Giaretta et al. may be explained by differences in environmental
1997, 1999; Rocha et al. 2000, 2001). Recent data show conditions among sites (Toft 1982) and also in the sam
that South-East Asia tropical forests may harbor very pling protocol used (Rocha et al. 2001, 2004b).
dense litter frog populations, although Frog distribution and abundance in the forest floor is
certainly not homogeneous but rather may depend on
local conditions of the leaf litter, such as humidity and
*Corresponding author. litter depth (Toft 1980a,b; Giaretta et al. 1997 ) .
Accepted for publication June 2006. Medium to large-scale studies have analyzed the effect

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L ITTER FROGS AT AN ATLANT IC RA INFOREST AREA 255

of landscape characteristics on the richness and abun south-eastern Brazil. They found that frog abundance
dance of amphibian species (Fauth et al. 1989; Toral et was related to soil cover depth, altitude, leaf-litter mass
al. 2002; Herrmann et al. 2005). Data on microhabi tat and fallen trunk area.
use by anurans are mostly limited to breeding sites for In the present study, we analyzed the species richness
adults and pond habitats for larvae and there are not and abundance of frogs inhabiting the leaf-litter floor in
much data on how microhabitat characteristics of an area of Atlantic Rainforest in Rio de Janeiro State,
the forest floor leaf litter may affect frog assemblages. south-eastern Brazil and tested how litter depth and
Fauth et al. (1989) analyzed the effect of litter depth and proportion of leaves in the leaf litter might affect these
elevation on herpetofauna density, richness and evenness parameters of the frog assemblage within the leaf litter.
in different sites in Costa Rica. Neither vari able affected We also evaluated how different species might respond
the density of the herpetofauna, but only leaf-litter depth to the environmental variables analyzed.
significantly affected species richness.
Allmon (1991) tested for the effect of litter moisture,
depth and average dry litter mass on the number of frogs METHODS
in the Brazilian Amazon and did not find any significant
results, although plots with frogs had higher average dry
litter mass than plots without frogs. study area
Ovaska (1991) found that quality and quantity of retreat
sites accounted for differences between sites in The study was carried out in a remnant patch of relatively
frog abundance and Herrmann et al. (2005) analyzed the undisturbed Atlantic Rainforest near Vila Dois Rios
effect of landscape and wetland characteristics on the (23°12ÿS, 44°13ÿW), Ilha Grande, a large (19 000 ha)
frog species richness and density. However, these continental island on the southern coast of Rio de Janeiro
studies used relatively large plots. The paucity of studies State, southeastern Brazil. Ilha Grande is covered by
on microhabitat characteristics and their effects on leaf- Atlantic Rainforest with different levels of regeneration
litter frogs prevents us from under standing as more due to disturbances caused by human activities over the
subtle and fine-grained landscape characteristics may last five centuries. Remnants of undisturbed (or less
affect communities and populations of these vertebrates. disturbed) forest occur in the less accessible central
areas of the island. Annual rain fall in the area is about
The Brazilian Atlantic Rainforest consists of a strip of 1700 mm with a mean annual temperature of 23°C
evergreen tropical rainforest that extends along the (Araújo & Oliveira 1988).
Atlantic coast of Brazil, from north-east to south and, in Field work was carried out in an area of approximately
the past, covered much of the Brazilian coast (Morellato 14 ha of relatively undisturbed Atlantic Rain forest,
& Haddad 2000). as one of the world's biodiversity located at an elevation of 220–240 m above sea level.
hotspots because of its high biological diversity, number We performed monthly samplings (nocturnal and diurnal)
of endemic species, and serious deforestation threat from August 1996 to October 1997 using small (2 m2 )
(Myers et al. plots. We delimited patches of forest floor using a
2000). Now it is reduced to approximately only 7% of its previously prepared 2 m ¥ 1 m cotton string having at its
original area (Rocha et al. 2003) and is considered to be corners small bamboo sticks that were fixed to the floor.
one of the most threatened tropical forests in the world All leaf litter inside the plot was removed by compressing
(Oliveira-Filho & Fontes 2000). The Atlantic rainforest portions of leaf litter and rapidly transferring it into a
harbors the world's greatest diversity of anuran species plastic bag (see Rocha et al. 2001 for details of the
and anuran reproductive modes (Duellman 1999; Haddad methodology). After all leaf litter was removed, we
& Prado 2005). Many of the Atlantic Forest frog species carefully checked the ground within the plot for any
are endemic, some with very restricted distributions remaining frogs. We sampled 234 plots, totaling 468 m2
(Duellman 1999). Despite the high species richness, data of forest leaf litter. Each plot was an independent
on community compo sition and abundance distribution sampling unit and was at least 20 m apart from adjacent
of anurans living on the forest floor leaf litter in Atlantic plots and no forest patch was sampled more than once.
Rainforest areas (including dense shoulderphilous forest All frogs encountered were identified to species level and
and restinga habitats) are scarce and restricted to a few quantified.
localities in south-eastern Brazil ( Duellman 1988; Heyer Before removing the leaf litter, we took five measures
et al. (at each corner and at the center of the plot) of the litter
nineteen ninety; Haddad & Sazima 1992; Giaretta et al. depth with a caliper to the nearest mm. We estimated
1997, 1999; Rocha et al. 2000, 2001, 2004b; Pombal & leaf-litter depth for each plot as the mean of these five
Gordo 2004; Van Sluys et al. 2004). The study of Giaretta measures.We also weighed the leaf litter (g) of each plot
et al. (1999) was the first to try to compare the relative and estimated the proportion of litter mass that was
importance of litter elements on the com munity structure represented by leaves, separated from twigs, branches
of frogs in the Atlantic Forest of and stones.We tested the simultaneous effect

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256 M. Van SLUYS ET AL.

130 influence of the proportion of leaves in the leaf litter

(Table 2; Fig. 3a). Litter depth also explained an
104 additional part of the variation in species richness, after
taking out the influence of the proportion of leaves in the
leaf litter (Table 2; Fig. 4).
78
number
plots
of
The three most abundant species (B. didactylus, E.
parvus and Z. parvulus) showed different trends in
52 abundance in relation to the leaf-litter characteristics
measured (Table 3). For B. didactylus, neither litter depth
26 nor proportion of leaves in the leaf litter influenced local
frog abundance (Fig. 3b), whereas for E. parvus and Z.
parvulus, the multiple regression yielded significant
0 -1 0 1 2 3 4 5 6 7
results. The depth of the leaf litter explained an additional
total abundance
part of the abundance of E. parvus (Fig. 3c) and both
Fig. 1. Abundance distribution of leaf-litter frogs species in 234 variables explained the abundance of Z. parvulus (Fig.
plots of 2 m2 in an Atlantic Rainforest area, at Ilha Grande, Rio 3d and Table 3), with greater abundance where there
de Janeiro State, Brazil. was deeper litter, and a lower proportion of leaves in the
litter.

of litter depth and proportion of leaves by mass (arcsine

transformed) on the total abundance of frogs and species
richness using multiple regression analysis (Zar 1999), DISCUSSION
considering plots with at least one frog.
Partial correlations were used to determine the relative
The eight species found in this study have all been
additional contribution of each of these variables on
previously reported from the same area in the study of
the total abundance of frogs and species richness. We Rocha et al. (2001), whose data for small plot sampling
performed the same statistical analysis for each of the comprise part of the data of the present study. Those
three most abundant species to evaluate how individual species are typical constituents of anuran communities
species numbers responded to these environmental in the leaf-litter floor in other Atlantic Forest areas (Lutz
variables of the leaf litter. All analysis and graphs were 1944; Lynch 1976; Heyer et al. 1990; Haddad & Sazima
made in Systat 11.0 (Systat Software Inc., San Jose, 1992; Giaretta et al. 1997, 1999; Pombal & Gordo 2004) .
USA). Only D. brevipollicatus has been suggested to be more
commonly found in bromeliads, not being considered as
a typical leaf-litter species (Peixoto 1995). All of the
RESULTS sampled species are endemic to the Brazilian Atlantic
Rainforest, with most of them being relatively widely
Approximately half (53%; 124/234) of the sampled plots distributed within this biome (Frost 2004). Exceptions are
contained in no frogs and 64 plots (27.3%) had only one B. didactylus, which is endemic to Rio de Janeiro State
individual (Fig. 1). The maximum number of frogs (Frost 2004; Rocha et al. 2004a) and Chiasmocleis sp.,
encountered in a given plot was six and the maximum which is apparently an undescribed species currently
number of species found in a plot was three. known only from Ilha Grande. Brachycephalus didactylus
We found a total of 185 individuals from eight species in was previously known from only two other areas in the
the remaining 110 plots (Table 1). All species are State (Tinguá and Teresópolis) and Rocha et al. (2000,
endemic to the Atlantic Rainforest biome. Brachyceph 2001) extended its range to Ilha Grande, ca 100 km
alus didactylus was the most abundant species, whereas south of its type locality.
Dendrophryniscus brevipollicatus was the least abundant.
The three most abundant species in the area (B. The plots surveyed on the forest floor patches with the
didactylus, Eleutherodactylus parvus and Zachaenus deepest leaf-litter layer tended to contain more frogs and
parvulus) accounted for 89.7% of all frogs found in the more species, independent of the contribution of leaves
plots (Table 1, Fig. 2). in that patch. This result indicated that microhabitat
Our predictor variables (leaf-litter depth and pro characteristics affect local distribution and abundance of
portion of leaves in the leaf litter) significantly affected frogs in the forest floor of Ilha Grande. Fauth et al. (1989)
overall abundance of frogs (R2 = 0.17; F2,107 = 10.779; and Allmon (1991) also found that litter depth affected
P = 0.0001; Table 2) as well as species richness (R2 = frog density in the areas they studied. However, the
0.11; F2.107 = 6.375; P = 0.002; Table 2). Only litter relative amount of leaves in the litter did not, alone,
depth explained an additional part of the variation in influence local pat terns of abundance.
total abundance of frogs, after taking out the
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L ITTER FROGS AT AN ATLANT IC RA INFOREST AREA 257

Table 1. Abundance (no. of individuals) and density (no. of individuals/100 m2 ) of leaf-litter frog species found in an Atlantic Rainforest area at Ilha Grande,
Brazil

Species abundance Density

Brachycephalidae
Brachycephalus didactylus 91 19.4
Leptodactylidae
Eleutherodactylus parvus 35 7.5
Eleutherodactylus guentheri 6 1.3
Eleutherodactylus binotatus 3 0.6
Zachaenus parvulus 40 8.5
Adenomera marmorata 4 0.8
Bufonidae
Dendrophryniscus brevipollicatus two 0.4
Microhylidae
Chiasmocleis sp. 4 0.8

Sample size = 234 plots (468 m2 ).

70 Table 2. Statistics of the Multiple Regression analyzes between total

abundance of frogs (R2 = 0.17; F2.107 = 10.779; P < 0.001) and species
60 richness (R2 = 0.11; F2.107 = 6.375; P = 0.002) and leaf- litter depth and
proportion of leaves (arcsine transformed) in the litter, for an area of Atlantic
50 Rainforest at Ilha Grande, Brazil

40
number
plots
of

effect Coefficient P-value

30
frog abundance
20 constant 1.1755 0.085
litter depth 0.0299 <0.001
10
Proportion of leaves in the litter -0.5883 0.319

0 species richness
Bd Ep Zp Eg Am Csp Eb Db constant 1,059 0.011
Species litter depth 0,014 0.001
Proportion of leaves in the litter -0,230 0.520
Fig. 2. Frequency of occurrence of leaf-litter frog species in 234 plots of 2
m2 in an Atlantic Rainforest area, at Ilha Grande, Rio de Janeiro State,
Brazil. Am, Adenomera marmorata; Bd, Brachycephalus didactylus; Csp,
Chiasmocleis sp.; Eb, Eleutherodactylus binotatus; Ep, Eleutherodactylus
parvus; Eg, Eleutherodactylus guentheri; Db, Dendrophryniscus brevipollica species (B. didactylus) was not associated with any of
tus; Zp, Zachaenus parvulus. the microhabitat characteristics, another (E. parvus) was
explained by leaf-litter depth and the third (Z. parvulus)
was positively related to litter depth and negatively with
the proportion of leaves . For this last species, the
Litter depth may enhance abundance by providing a negative relationship between abundance and proportion
wider range of microhabitats and refuges against of leaves in the leaf litter may result from the fact that the
predators, as well as greater densities of arthropod preys highest abundance in any one plot (four individuals) was
(Fauth et al. 1989). Also, for terrestrial breeding frogs (as found in only one plot, which did not have a high
are most of the species found in our study), the leaf layer proportion of leaves in the litter. These results indicate
may provide adequate conditions for egg laying. This that, although, in general, litter depth affected patterns of
may facilitate the coexistence of more individuals and frog abundance in the litter floor at Ilha Grande, each
more species in the leaf litter. Scott (1976) argued that species may have an individual response to local
the reduced amount of litter pro duced in South-East environmental conditions.
Asia may account for the reduced herpetofaunal Other abiotic variables not measured in our study may
abundances there. But this author and Fauth et al. (1989) also affect frog abundance such as leaf-litter moisture.
studied the herpetofauna as a whole, not only anurans, Elevation is a major factor which affects herpetofaunal
which may restrict comparing our results with theirs. species richness (Fauth et al. 1989; Giaretta et al.
1999). Because our samples were all performed at about
The analysis for the three most abundant frog species the same altitude (200–240 m) we believe this factor did
gave different results. abundance of one not affect our results.

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258 M. Van SLUYS ET AL.

The)

5 5

4
4

3
3

two

abundance
frog abundance
frog

two

1
0

0 -1

-2
-1 -30 -20 -10 0 10 20 30 40 50 -1.0 -0.5 0.0 Proportion 0.5
litter depth of leaves

B)
3 3

two two

abundance
frog

1 abundance
frog

0 0

-1 -30 -20 -10 0 10 20 30 40 50 -1 -1.0 -0.5 0.0 Proportion 0.5

litter depth of leaves

w)
1.0 1.0

0.5 0.5

abundance
frog abundance
frog

0.0 0.0

-0.5 -0.5
-30 -20 -10 0 10 20 30 40 Litter depth -1.0 -0.5 0.0 Proportion 0.5
of leaves

d)
3 3

two two

abundance
frog
1 abundance
frog
1

0 0

-1 -20 -10 0 10 20 30 -1 -0.3 -0.2 -0.1 0.0 0.1 0.2 0.3 Proportion of leaves
litter depth

Fig. 3. Relationships (partial regressions) between mean litter depth (mm) and proportion of leaves in the leaf litter (%) and total
abundance of (a) overall litter frogs community, (b) Brachycephalus didactylus, (c) Eleutherodactylus parvus , ( d) Zachaenus parvulus
in an Atlantic Rainforest area, at Ilha Grande, Rio de Janeiro State, Brazil.

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L ITTER FROGS AT AN ATLANT IC RA INFOREST AREA 259

two two

1 1

richness
species richness
species

0 0

-1 -30 -20 -10 0 10 20 30 40 50 -1 -1.0 -0.5 0.0 0.5

litter depth Proportion of leaves

Fig. 4. Partial regression plots between mean litter depth (mm) and proportion of leaves in the leaf litter (%) and species richness of litter frogs in an Atlantic
Rainforest area, at Ilha Grande, Rio de Janeiro State, Brazil.

Table 3. Statistics of the multiple regression of litter depth and proportion of leaves in the leaf litter on the abundance of the three most abundant frog species in the
leaf litter at Ilha Grande, Rio de Janeiro, Brazil

Total Total independent Partial correlation partial correlation

Species R2 F p-value variable coefficient p-value

Brachycephalus didactylus (n = 0.05 1,686 0.194 constant 0.049

66) litter depth 0.222 0.076
proportion of leaves -0.042 0.731
Eleutherodactylus parvus (n = 0.20 3,715 0.037 constant 0.129
32) litter depth 0.437 0.013
proportion of leaves 0.102 0.542
Zachaenus parvulus (n 0.31 6,354 0.005 constant 0.002
= 31) litter depth 0.530 0.005
proportion of leaves -0.533 0.005

At this point, we do not know the causes of the relatively low from National Council for Scientific Development
richness of frogs in the leaf litter of Ilha Grande. However, because fico e Tecnológico (CNPq, Processes no. 301401/04-7, no.
this site is one important remnant of the Atlantic Rainforest in eastern 302718/03-6, no. 462003/00-0 and no. 307653/03-0, respectively).
Brazil (Rocha et al. 2003) and because most of the species found in FAPERJ and CNPq partially supported the project (FAPERJ grants:
the leaf litter are endemic (both to the Atlantic Rainforest and to Rio no. E-26/172.383/ 2000 and CNPq: no. 477981/03-8). This study is
de Janeiro State, Rocha et al. , 2004a) it is important to preserve the also part of the BIOTA/FAPESP Project and was partially supported
site as a contribution to the conservation of tropical frog biodiversity with a grant from this Agency (process no. 99/08291-5).
as a whole.

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