ZOOLOGIA 26 (1): 97102, March, 2009

The leaf litter of most tropical forests is the habitat of a

rich herpetofauna, mainly composed of small lizards and frogs
(e.g., HEATWOLE & SEXTON 1966, LLOYD et al. 1968, SCOTT 1976,
1982, INGER 1980, HEINEN 1992, VONESH 2001, ALMEIDA-GOMES et
al. 2008). Leaf litter frogs tend to be more abundant in Neotro-
pical forests when compared to the Old World tropics (SCOTT
1976, 1982, ALLMON 1991, VONESH 2001), although this is not a
rule (HOFER & BERSIER 2001, WATANABE et al. 2005). The reasons
for this difference are not clear, but may be in part due to the
greater availability of nutrients in many Neotropical areas, as-
sociated with their more recent geological age (ALLMON 1991).
Environmental and structural parameters at each local-
ity partially explain the differences in species richness and den-
sity among communities (e.g. SCOTT 1976, WATANABE et al. 2005,
VAN SLUYS et al. 2007). Some studies about litter frogs in tropical
forests have included density estimates, which allow quantita-
tive comparisons among anuran faunas (ALLMON 1991 and in-
cluded references, GIARETTA et al. 1997, 1999, ROCHA et al. 2001,
2007, HUANG & HOU 2004, WATANABE et al. 2005, ALMEIDA-GOMES
et al. 2008). In South America, studies of tropical forest leaf
litter frog assemblages providing data on frog densities are still
relatively rare, particularly with regard to the Brazilian Atlan-
tic Rainforest (GIARETTA et al. 1997, 1999, ROCHA et al. 2000, 2001,
2007, VAN SLUYS et al. 2007, ALMEIDA-GOMES et al. 2008). The At-
lantic Rainforest extends along the eastern coast of Brazil and
is considered as one of the worlds biodiversity hotspots be-
cause of its high biological diversity and rates of endemism,
and severe deforestation rate (MYERS et al. 2000). This biome
may harbor the worlds greatest diversity of anuran species
(DUELLMAN 1999, YOUNG et al. 2004).
Despite intense deforestation, the state of Rio de Janeiro
still has proportionately large forested areas (TANIZAKI-FONSECA
& MOULTON 2000, ROCHA et al. 2003) and has relatively high
anuran endemism rates (ROCHA et al. 2004). Herein, data are
presented on species composition, richness, relative abundance,
and density for the leaf litter anuran assemblage of an area
within the Serra dos rgos mountain range, which represents
one of the largest continuous areas of Atlantic Rainforest in
the state of Rio de Janeiro (ROCHA et al. 2003). The results are
compared with those reported for other studied rainforest ar-
eas worldwide.
MATERIAL AND METHODS
The Parque Estadual dos Trs Picos (hereafter PETP) is a
recently created conservation unit that encompasses much of
the Serra dos rgos mountain range, and extends along five
municipalities in the state of Rio de Janeiro, in southeastern
Brazil. With an area of more than 46,000 ha, it comprises one
of the most extensive areas of Atlantic Rainforest in the state
(ROCHA et al. 2003). The study was carried out within a private
property (Fazenda Santa Brbara; 2225S and 4235W) located
inside the PETP, in the municipality of Cachoeiras de Macacu.
The annual rainfall in the area is approximately 2,500 mm and
the mean annual temperature varies between 16 and 18C.
Density and richness of leaf litter frogs (Amphibia: Anura) of an Atlantic
Rainforest area in the Serra dos rgos, Rio de Janeiro State, Brazil
Carla C. Siqueira
1, 2
; Davor Vrcibradic
1
; Mauricio Almeida-Gomes
1
; Vitor N. T. Borges-Junior
1
;
Patrcia Almeida-Santos
1
; Marlon Almeida-Santos
1
; Cristina V. Ariani
1
; Diego M. Guedes
1
;
Pablo Goyannes-Arajo
1
; Thiago A. Dorigo
1
; Monique Van Sluys
1
& Carlos F. D. Rocha
1
1
Departamento de Ecologia, Universidade do Estado do Rio de Janeiro. Rua So Francisco Xavier 524, Rio de Janeiro,
20550-011 Rio de Janeiro, Brasil.
2
Corresponding author. E-mail: carlacsiqueira@yahoo.com.br
ABSTRACT. Data on species composition, richness, and density are presented for the leaf litter frog assemblage of an
area of Atlantic Rainforest at the Serra dos rgos mountain range, in the state of Rio de Janeiro, southeastern Brazil.
Three sampling methods were used: plot sampling, visual encounter surveys, and pitfall traps. The local assemblage of
leaf litter frogs was composed of 16 species, with the direct-developing species, Euparkerella brasiliensis (Parker, 1926),
being the most abundant. The estimated density of the local leaf litter frog assemblage based on plot sampling was 17.1
ind/100 m
2
and the estimated overall leaf litter frog mass was 684.2 g/ha. The estimated density of leaf litter frogs at the
present study is the highest currently reported for Atlantic Rainforest areas, which reinforces the idea of higher densities
of leaf litter frogs in the Neotropical Region compared to the Old World tropics.
KEY WORDS. Amphibian survey; relative abundance; tropical forest.
98 C. C. Siqueira et al.
ZOOLOGIA 26 (1): 97102, March, 2009
Surveys were conducted during late October and early
November 2006 at altitudes between 500 and 800 m, using
three sampling methods: plots or quadrats (JAEGER & INGER 1994),
visual encounter surveys (CRUMP & SCOTT 1994), and pitfall traps
with drift fences (CORN 1994).
For the plot method, 25 quadrats of 5 x 5 m were estab-
lished on the forest floor during the afternoon, totaling 625
m
2
of sampled area. The corners of each plot were marked with
wooden stakes and the area inside was enclosed with a 50 cm
high soft plastic fence, whose base was buried or attached to
the ground. After sunset, each plot was carefully searched for
about 30 minutes by five people using headlamps, moving on
hands and knees, side-by-side. During the search, leaves,
branches, and stones were overturned with hand rakes; rock
crevices and fissures among tree roots were also checked.
For visual encounter surveys, 150 transects lasting 30 min
each were carried out by 10 people, totaling 75 hours of sam-
pling effort (7.5 h/person). Equal numbers of transects (50) were
surveyed during each period of the day (diurnal, crepuscular,
and nocturnal). During each transect sampling, the observer
moved at a slow walking pace, carefully searching all types of
potential microhabitats for frogs.
Three pitfall trap systems were used during a total of 18
days. Each system consisted of 10 30-liter buckets buried on
the ground and set ca. 5 m apart, with soft plastic drift fences
about 50 cm high extended among them. Six buckets were set
in line and the other four were placed at opposite ends of the
fence, perpendicularly to the main axis. Pitfalls were checked
once a day, always in the morning.
All frogs found using the three sampling methods were
collected and identified. Besides, all individuals found during
casual encounters were also recorded. For an estimate of spe-
cies composition and richness of the leaf litter frog assemblage,
the species recorded by all three sampling methods plus those
collected during casual encounters were considered. For esti-
mates of density only the data obtained by plot sampling were
considered, as this method has been frequently used in tropi-
cal forests worldwide (e.g. ALLMON 1991, WATANABE et al. 2005)
and allows comparisons with other forested areas for which
data is available. The total mass of leaf litter frogs per hectare
(g/ha) was also estimated. Voucher specimens of all frog spe-
cies recorded during the study were deposited at the Museu
Nacional, Rio de Janeiro.
RESULTS
Sixteen frog species belonging to eight families were re-
corded in the leaf litter during the study (Tab. I). The local
anuran assemblage was dominated by species with direct de-
velopment (Terrarana sensu HEDGES et al. 2008), with Euparkerella
brasiliensis (Parker, 1926) being the most abundant species (com-
prising 51% of all individuals collected), followed by
Brachycephalus didactylus (Izecksohn, 1971) (16%). The species
with the highest mean body mass were Rhinella icterica (Spix,
1824) (91.2 88.0 g), followed by Proceratophrys appendiculata
(Gnther, 1873) (36.0 g), R. ornata (Spix, 1824) (35.2 21.8 g),
and P. boiei (Wied-Neuwied, 1824) (32.0 14.4 g), whereas B.
didactylus (0.07 0.02 g) was the species with lowest mean
body mass (Tab. I).
A total of 107 individuals belonging to seven frog spe-
cies were found in the plots (Tab. I). Three frogs escaped (one
E. brasiliensis, one B. didactylus and one individual that could
not be identified). The estimated overall frog density of the
local leaf litter frog assemblage was 17.1 ind/100 m
2
, and the
estimated overall frog mass was 684.2 g/ha (Tab. I). The frogs
that occurred with the highest densities were E. brasiliensis (9.6
ind/100 m
2
) and B. didactylus (4.0 ind/100 m
2
), which together
comprised about 80% of all individuals found in plots. Hylodes
pipilans Canedo & Pombal, 2007 was the species with the low-
est estimated density (0.2 ind/100 m
2
) on the forest floor. The
species with the highest estimated mass per hectare were E.
brasiliensis (380.3 g/ha) and Haddadus binotatus (Spix, 1824)
(150.1 g/ha), whereas H. pipilans (8.1 g/ha) and Ischnocnema
parva (Girard, 1853) (9.4 g/ha) had the lowest estimated mass
per hectare (Tab. I).
Twelve frog species were recorded during transect sam-
plings, with E. brasiliensis (N = 23, or 56% of all individuals
found) being the most abundant (Tab. I). Almost all frog spe-
cies sampled in transects were found during the crepuscular
period (N = 10 species, or 83% of the species in the assem-
blage), whereas only three (B. didactylus, Zachaenus parvulus
(Girard, 1853), and E. brasiliensis) were found during diurnal
samplings (25%). Most individuals were found during crepus-
cular (N = 24, or 58% of all individuals sampled) and noctur-
nal (N = 14, or 34%) transects, whereas the proportion of frogs
found during diurnal transects (N = 3, or 7.3%) was compara-
tively lower.
Twelve frogs belonging to seven species were captured
by pitfall traps (Tab. I). The most frequently captured species
in the pitfalls were R. icterica (N = 4, or 33% of all specimens
captured) and E. brasiliensis (N = 3, or 25%).
DISCUSSION
Our data indicate that frog density at the studied area is
relatively high, as the estimated total density was twice as high
as that of another Atlantic Forest area (Reserva Ecolgica de
Guapiau) located only ca.15 km from the study area (Tab. II).
Other large-plot studies have yielded even lower density es-
timates for different Atlantic Forest areas of southeast Brazil
(1.4-5.9 ind/100 m
2
; Tab. II). In fact, estimated density of leaf
litter frogs at the Fazenda Santa Brbara is the highest yet re-
ported for Atlantic rainforest areas. Considering other tropical
rainforest areas worldwide for which litter frog densities were
estimated (Tab. II), our data for the PETP is also on the high
side. The estimated leaf litter frog density in our study was
higher than that of a survey conducted in the Brazilian Ama-
zon, but close to that reported for a Peruvian site during the
99 Density and richness of leaf litter frogs of an Atlantic Rainforest area
ZOOLOGIA 26 (1): 97102, March, 2009
wet season (Tab. II). In Central America, litter frog densities are
generally similar or higher than that of the present survey (Tab.
II). In most studied areas of tropical Africa and Asia the esti-
mated densities were lower than that of our study, except for
one site at Iriomote Island (Ryuku Archipelago, Japan) for which
a much higher value (41.8 ind/100m
2
) was obtained (Tab. II).
Our data thus reinforces the idea that higher densities of leaf
litter frogs tend to occur in the Neotropical region compared
to the Old World tropics.
Individuals of species with direct development numeri-
cally dominated the anuran assemblage at PEPT, as it has been
observed in most studied Neotropical litter frog assemblages
(e.g. SCOTT 1976, LIEBERMAN 1986, FAUTH et al. 1989, GIARETTA et al.
1997, 1999, ROCHA et al. 2001, 2007). Euparkerella brasiliensis, a
small direct-developing frog, was the most abundant frog spe-
cies at the PETP and had the highest estimated value of mass
per hectare of all frogs sampled in plots, despite its small body
size. This is a poorly known species, probably due to its small
body size and relatively limited geographical range (previously
restricted to the municipalities of Rio de Janeiro and
Guapimirim; IZECKSOHN 1988). Another small direct-developing
frog, B. didactylus, the smallest frog sampled in the present study,
was the second most abundant species in the study area.
Brachycephalus didactylus (adult SVL up to 10-11 mm) is argu-
ably the worlds smallest tetrapod species (ESTRADA & HEDGES
1996, HEDGES et al. 2008) and it was previously known from
only three other localities in the state of Rio de Janeiro (VAN
SLUYS et al. 2007). The present data extend the distribution of
E. brasiliensis and B. didactylus some 20 km to the east.
The estimated total mass of frogs per hectare was lower
than that reported for another Atlantic forest area (Ilha Grande)
by ROCHA et al. (2001) based on data from 8 x 8 m plots (1,150 g/
Table I. Number of individuals recorded for each sampling method and mean body mass ( one standard deviation) of each species, and
estimated density and mass per area of frogs recorded in the leaf litter of an Atlantic Rainforest area within the Parque Estadual dos Trs
Picos, in southeastern Brazil.
Species Plots Transects Pitfalls Extra Total
Body mass
(g)
Density
(frogs/100m
2
)
Mass (g/ha)
Brachycephalidae
Brachycephalus didactylus (Izecksohn, 1971)* 25 2 1 28 0.07 0.02 4.0 26.1
Ischnocnema guentheri (Steindachner, 1864) 5 2 4 11 1.00 0.50 0.8 56.0
I. parva (Girard, 1853) 3 2 5 0.30 0.10 0.5 9.4
I. octavioi (Bokermann, 1965)* 1 1 0.20
Craugastoridae
Haddadus binotatus (Spix, 1824) 3 3 3.30 2.40 0.5 150.1
Cycloramphidae
Proceratophrys appendiculata (Gnther, 1873) 1 1 36.00
P. boiei (Wied-Neuwied, 1824) 3 4 7 32.00 14.40
Thoropa miliaris (Spix, 1824) 1 1 26.20
Zachaenus parvulus (Girard, 1853) 9 2 1 4 16 1.10 0.80 1.4 54.2
Hylodidae
Crossodactylus aeneus Mller, 1924 2 2 1.10 0.40
Hylodes pipilans Canedo & Pombal, 2007* 1 1 2 0.70 0.20 0.2 8.1
Leptodactylidae
Leptodactylus marmoratus Steindachner, 1867 1 1 1.20
Bufonidae
Rhinella icterica (Spix, 1824) 1 4 5 91.20 88.00
R. ornata (Spix, 1824) 1 1 2 35.20 21.80
Microhylidae
Myersiella microps (Dumril & Bibron, 1841) 1 1 2 0.70 0.20
Strabomantidae
Euparkerella brasiliensis (Parker, 1926)* 60 23 3 3 89 0.40 0.20 9.6 380.3
Total (**) 107 41 12 19 176 17.1 684.2
* Species endemic to Rio de Janeiro state. ** Four individuals were seen but escaped in the course of the study: one unidentified frog,
one B. didactylus and one E. brasiliensis during plot sampling, and one E. brasiliensis during transect sampling.
100 C. C. Siqueira et al.
ZOOLOGIA 26 (1): 97102, March, 2009
ha). As the estimated frog density was lower at Ilha Grande, this
difference in mass may be due to the assemblage composition
in each locality. At the PETP, the most abundant frogs found in
plots were small (E. brasiliensis) to very small (B. didactylus),
whereas at Ilha Grande the assemblage was dominated by spe-
cies about equal in size (I. parva) or larger (Z. parvulus) than E.
brasiliensis (ROCHA et al. 2001). In another Atlantic rainforest area
(GIARETTA et al. 1999), the estimated mass of frogs per hectare
(476.6 g/ha) was slightly lower than that of the present study,
whereas in rainforests studied in other parts of the world they
were higher, being 1,088 g/ha in Peru (TOFT 1980a) and 11,460
g/ha in Iriomote island, Japan (WATANABE et al. 2005).
Although most species of frogs at the PETP were sampled
in transects (75% of all sampled species) and three species
Table II. Summary of data from studies of leaf litter frog assemblages using large-plot (5 x 5 m or larger) sampling in tropical rainforests
worldwide. Acronyms for Brazilian states are: (AM) Amazonas, (RJ) Rio de Janeiro, and (SP) So Paulo.
Locality Altitude (m) Season Density (ind/100 m
2
) Reference
South America
Brazil, P. E. dos Trs Picos (RJ) 500-800 wet 17.1 This study
Brazil, Reserva Ecolgica de Guapiau (RJ) 40-400 wet 8.4 ROCHA et al. (2007)
Brazil, Ilha Grande (RJ) 220-230 wet 5.9 ROCHA et al. (2001)
Brazil, Morro So Joo (RJ) 10-320 dry 4.5 ALMEIDA-GOMES et al. (2008)
Brazil, Serra do Japi (SP) 850-1000 dry 1.4 GIARETTA et al. (1997)
Brazil, Parque Florestal de Itapetinga (SP) * 900-1250 wet 5.1 GIARETTA et al. (1999)
dry 4.2
Brazil, INPA-WWF reserve (AM) 150 wet 6.0 ALLMON (1991)
dry 3.0
Peru, Panguana 210 wet 15.5 TOFT (1980a)
dry 4.4
Central America
Costa Rica, Osa lowland wet 18.9 SCOTT (1976)
dry 14.0
Costa Rica, La Selva 100 wet 21.4
dry 13.4
Costa Rica, San Vito 1200 wet 62.3
dry 55.1
Costa Rica, La Selva * 60 both 15.7 LIEBERMAN (1986)
Costa Rica, La Selva * lowland dry 11.5 HEINEN (1992)
Panama, Pipeline road 30 wet 7.5 TOFT (1980b)
dry 19.4
Panama, Carti road 300 wet 11.8
dry 30.9
Panama, Silugand lowland wet 30.2 HEATWOLE & SEXTON (1966)
Asia
Thailand, Sakaerat ** lowland both 0.5-2.6 INGER & COLWELL (1977)
Borneo, nanga Tekalit lowland both 1.2 LLOYD et al. (1968)
Taiwan, Nanjen-shan Nature Reserve** 10-460 both 3.5-10.2 HUANG & HOU (2004)
Japan, Iriomote Island 25 both 41.8 WATANABE et al. (2005)
Africa
Uganda, Kibale * 1530 wet 2.2 VONESH (2001)
dry 1.5
Cameroon, Douala-Eda Game Reserve lowland dry 9.4 SCOTT (1982)
* Only density estimates from unburned/undisturbed sites were considered. ** Two different forest environments were sampled.
101 Density and richness of leaf litter frogs of an Atlantic Rainforest area
ZOOLOGIA 26 (1): 97102, March, 2009
Ischnocnema octavioi (Bokermann, 1965), Crossodactylus aeneus
Mller, 1924 and Leptodactylus marmoratus Steindachner, 1867
were recorded only by this method, three other species were
sampled only using other methods. One species, H. binotatus,
was sampled only in plots and the other two, P. appendiculata
and Thoropa miliaris (Spix, 1824) (one individual each), were
captured only by pitfall traps. Thus, the data indicates that,
although a particular method may be more efficient to capture
more individuals (plot sampling) or more species (transects),
the three sampling methods should be used to better estimate
the richness and abundance of frogs in leaf litter assemblages,
at least in Atlantic Rainforest areas. Moreover, during the
present study, frogs were captured mainly during the crepus-
cular and nocturnal transects (92.7% of all individuals). This
has also occurred in previous studies carried out in other At-
lantic rainforest areas (ROCHA et al. 2000, 2007, ALMEIDA-GOMES
et al. 2008). Thus, the data obtained at the PETP reinforce the
idea that most of the leaf litter frog activity in the Atlantic
forest is crepuscular to nocturnal.
The PETP, created as recently as 2002, currently repre-
sents the largest protected area of Atlantic Rainforest in the
state of Rio de Janeiro (ROCHA et al. 2003). Although the present
study represents a short-term survey carried out at a compara-
tively small area within this reserve, the results indicate a rela-
tively high species richness and density of leaf litter frogs. This
suggests that the whole area encompassed by the park (over
46,000 ha) may harbor a considerable anuran richness and
numerous sites where the forest floor is densely populated by
frogs. The presence of at least four species endemic to the state
of Rio de Janeiro attest to the importance of the PETP as a con-
servation unit for anuran species, which are currently facing a
worldwide decline due to factors such as global climate change,
pollution, habitat loss, and the spread of diseases such as
chytridiomycosis (e.g. YOUNG et al. 2004, TOLEDO et al. 2006, NAVAS
& OTANI 2007).
ACKNOWLEDGEMENTS
This study was sponsored by Critical Ecosystem Partner-
ship Fund, Conservao Internacional and Aliana para a
Conservao da Mata Atlntica. CFDR (Processes 307653/2003-
0 and 477715/2006-0) and MVS (Process 301401/2004-7) re-
ceived research grants from the CNPq. CFDR received a research
grant from Fundao de Amparo Pesquisa do Estado do Rio
de Janeiro (FAPERJ) through the Programa Cientistas do Nosso
Estado (Process E-26/100.471.2007). Graduate fellowships were
granted by the FAPERJ to CCS, by the CAPES to PAS, and by
CNPq to VNTBJ. We are especially thankful to Alexander J. Davis
for local support and for making many facilities available. We
also thank CAG Cruz, J.P. Pombal Jr, C. Canedo, B.V.S. Pimenta,
and E. Izecksohn for their help with the identification of frogs.
Currently CCS is associated to the Programa de Ps-Graduao
em Ecologia from Universidade Federal do Rio de Janeiro and
receives a PhD grant from CNPq.
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Submitted: 23.VI.2008; Accepted: 11.III.2009.
Editorial responsibility: Glauco Machado