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Phyllomedusa- 3(1), September 2004
51
Ecological aspects of the casque-headed frog
Aparasphenodon brunoi (Anura, Hylidae) in a
Restinga habitat in southeastern Brazil
Daniel Oliveira Mesquita1, Gabriel Corrêa Costa2 and Mariana G. Zatz3
Universidade de Brasília, 70910-900, Brasília, DF, Brazil.
1Pós-graduação em Biologia Animal, Instituto de Ciências Biológicas. E-mail: danmesq@unb.br.
2Pós-graduação em Ecologia, Instituto de Ciências Biológicas. E-mail: costagc@unb.br.
3Colecão Herpetológica da UnB – CHUNB, Departamento de Zoologia, Instituto de Ciências Biológicas. E-mail:
mariana@unb.br.
Phyllomedusa 3(1):51-59, "
© 2004 Melopsittacus Publicações Científicas
ISSN 1519-1397
Received 6 March 2004.
Accepted 4 August 2004.
Distributed 30 September 2004.
Abstract
Ecological aspects of the casque-headed frog Aparasphenodon brunoi (Anura,
Hylidae) in a Restinga habitat in southeastern Brazil. We describe some aspects
of the ecology of Aparasphenodon brunoi, a species associated with bromeliads. We
comment on the relationships of this species with bromeliad size, microhabitat use,
diet and sexual dimorphism. This study was conducted on a Restinga habitat near
Presidente Kennedy, state of Espírito Santo, southeastern coast of Brazil. When the
animals were found inside the bromeliads, we measure bromeliad and head size of
frogs. We analyzed stomach contents and determined the sex and reproductive
condition. We found 17 individuals (58.6%) in bromeliad leafs, six (20.7%) in
Cactaceae, three (10.3%) in liana and three (10.3%) on trunks. The correlation between
head measurements and bromeliad size were high, indicating that animals apparently
use bromeliads based on their size, which could be related to the minimization of water
loss. The most common prey items were beetles, ants, and insect larvae, suggesting
that the species is relatively generalist in prey consumption. Aparasphenodon brunoi
showed significant sexual size and shape dimorphism with females having larger bodies
than males (size) and females having tibia, eye diameter and SVL larger than males
(shape), but larger sample size and more detailed ecological and life history data are
needed to elucidate the factors that have led to sexual size dimorphism.
Keywords: Anura, Hylidae, Aparasphenodon brunoi, casque-headed frog, diet,
microhabitat use, bromeliads, sexual dimorphism.
Introduction
Many species of invertebrates and vertebra-
tes use bromeliads for foraging, reproduction
and escaping from predators. Among vertebra-
tes, amphibians and reptiles are the most
common inhabitants. For example, the lizard
Mabuya macrorhyncha preys on animals that
live between bromeliad leaves of Neoregelia
(Vrcibradic and Rocha, 1996), and the frog
Phyllodytes luteolus carries out its entire life
cycle inside bromeliads (Teixeira et al. 1997,
Eterovick 1999).
Phyllomedusa- 3(1), September 2004
52
Aparasphenodon are tree frogs
characterized by a strongly ossified skull, which
gives them the common name “casque-headed
frog” (Pombal 1993). The ossified skull appears
to have evolved independently six times in
hylids, apparently as an adaptation to similar
habitats where water is scarce (Trueb 1970).
Several genera of casque-headed frogs occur in
South America, including Osteocephalus,
Phrynohyas, Trachycephalus, Corythomantis,
Aparasphenodon and some species of Scinax
and Hyla. Aparasphenodon may be closely
related to Corythomantis (Trueb 1970). The
genus Aparasphenodon consists of three
species, ranging from southern Brazil to the
Orinoco river basin in northern Venezuela
(Argôlo 2000, Frost 2002). Aparasphenodon
brunoi Miranda-Ribeiro, 1920 occurs in coastal
areas of São Paulo, Rio de Janeiro, Espírito
Santo and Bahia states and continental areas of
the Parque Estadual do Rio Doce, in Minas
Gerais state (Feio et al. 1998, Argôlo 2000).
The species is relatively common in Restinga
habitats, which are white sand dunes partially
covered by herbaceous plants and shrubs. This
vegetation forms dispersed islands of vegetation
(Suguio and Tessler 1984), where the frog is
usually found associated to bromeliads.
Bromeliads are abundant in the Restinga and
occur on many kinds of substrates, including
soils with organic material, sandy soils, and tree
trunks (Cogliatti-Carvalho et al. 2001). The
leaves of bromeliads typically form a rosette,
within which water accumulates. The shape and
size of the bromeliads determine the amount of
water that can accumulate (Leme 1984). This
frog species is highly associated with
bromeliads and reaches 80 mm in snout vent
length (SVL) (Feio et al. 1998), but little is
known about their ecology.
Herein, we describe some aspects of the
ecology of Aparasphenodon brunoi from a
Restinga in the southern Espírito Santo state,
southeastern Brazil. We address the following
questions: (1) What are the patterns of habitat
and microhabitat use? (2) Is there a relationship
between the size of bromeliads and frog size?
(3) Is there a significant sexual dimorphism? (4)
What are the most important prey items?
Material and Methods
The study was performed in a gradient that
varies from a Restinga (21°17’59’’S,
40°57’30’’W) to a forested area within Restinga
(21°17’40’’S, 40°57’35’’W), near Presidente
Kennedy, Espírito Santo state, on the
southeastern coast of Brazil, from 20 to 27
September 2001. Aparasphenodon brunoi
(Figura 1) was collected during the day by the
Restinga shrubs and by searching bromeliads.
The forest site was visited during the day and
night. Individuals were located visually and by
their calls. Microhabitat categories, including
branch, liana, Cactaceae, trunk, bromeliad
leaves, and inside bromeliads were recorded for
29 individuals as the original position that it was
found at first sight. The height (in cm) of each
individual above the ground was recorded. All
frogs were killed in 10% alcohol solution and
preserved in 10% formalin. Specimens were
deposited in the Coleção Herpetológica da
Figure 1 - Aparasphenodon brunoi from Presidente
Kennedy (ES) (sex unknown). Photo: Adrian
A. Garda.
Mesquita et al.
Phyllomedusa- 3(1), September 2004
53
Universidade de Brasília (CHUNB 15935,
15937–15940, 24369, 24781–24800, 24909–
24945).
The following measurements were taken
only on frogs found inside the bromeliads and
of the correspondent plant which they were
associated: snout-vent length (SVL), head length
(from the tip of the snout to the commissure of
the mouth), head width (at its broadest point)
and head height (at its highest point) (N=10)
(using a Mitutoyo® digital caliper, to the nearest
0.01 mm), plant height, width between external
leaves, and diameter of the bromeliad rosette
(using a ruler and a measuring tape). Canonical
correlation analysis was used to investigate the
relationship between frog size and bromeliad
size.
For the analysis of feeding habits we
removed the stomachs (N=63) and identified
prey items to order. We recorded the length and
width (to the nearest 0.01 mm) of intact prey
with Mitutoyo® digital calipers, and estimated
prey volume (V) as an ellipsoid:
,
223
42
π= lw
V
where w is prey width and l is prey length.
We calculated the numeric and volumetric
percentages of each prey category for individual
frogs and for pooled stomachs. To investigate
the relationship between prey size and frog head
measurements, we used a canonical correlation
analysis with two sets of variables: maximum
prey length and width vs. frog head width,
height, and length.
We determined sex and reproductive condi-
tion of each frog using direct observation of
gonads. The females were considered repro-
ductive when their ovaries were extremely
convoluted and enlarged. We considered as
reproductive males, individuals that have
completely evident vocal sac. We recorded for
all individuals, collected inside and outside of
bromeliads and previously deposited in CHUNB
(N=63), the following morphometric variables:
SVL, head width, height, and length; tibia,
forearm and foot length; and tympanum and eye
diameter. We considered the SVL of the smaller
reproductive male and female as the SVL of
sexual maturity; all individuals with SVL equal
or superior to that ones were considered adults.
We log-transformed (base 10) all morpho-
metric variables prior to analyses to meet the
requirements of normality (Zar 1998). To
partition the total morphometric variation
between size and shape variation, we defined
body size as an isometric size variable (Rohlf
and Bookstein 1987) following the procedure
described by Somers (1986). We calculated an
isometric eigenvector defined a priori with
values equal to p-0.5, where p is the number of
variables (Jolicoeur 1963). Next, we obtained
scores from this eigenvector, hereafter called
body size, by post-multiplying the n by p matrix
of log-transformed data, where n is the number
of observations, by the p by 1 isometric
eigenvector. To remove the effects of body size
from the log-transformed variables, we used
residuals of regression between body size and
each shape variable. To test the null hypothesis
of no difference between sexes, we conducted
separate analyses of variance on body size
(ANOVA) and the shape variables (MANOVA)
of adult individuals.
We carried out statistical analyses using
SYSTAT 9.0 for Windows with a significance
level of 5% to reject null hypotheses.
Throughout the text, means appear ± 1 SD. All
measures are in mm.
Results
We collected, during the night, a total of 29
active individuals, being 17 (58.6%) in
bromeliad leaves, six (20.7%) in cactaceae,
three (10.3%) in lianas and three (10.3%) in
trunks (Figure 2). The animals were at mean
height of 65.7 cm (25-210 cm). During the day,
the individuals were found only inside the
bromeliads (N=10). In this period their activity
Ecology of the casque-headed frog Aparasphenodon brunoi (Anura, Hylidae)
Phyllomedusa- 3(1), September 2004
54
was restricted to emitting call from the interior
of bromeliads.
The correlation between the measures of the
body and the measures of the bromeliads were
high. The first and second canonic variables of
the body measures show that the three measures
have equal influence in the composition of the
canonic variable. The first canonic variable of
the bromeliad measurements gave more
emphasis in the bromeliad width and height. The
first canonical correlation was 0.972, having
statistical significance (p=0.022), showing
association between the body measures of
Aparasphenodon brunoi and the bromeliad
measurements (Table 1).
We analyzed 85 stomachs and 26% (22)
were empty. We found 10 prey categories, being
more frequent beetles (56.1%) and ants (16.8%).
Considering the number of items per stomach,
the diet consisted mainly of beetles (44.0%) and
ants (17.1%). By volume, beetles were the most
important prey item (60.5%), followed by insect
larvae (16.5%) and ants (16.1%) (Table 2). The
Figure 2 - Frequency distribution of Aparasphenodon
brunoi according to microhabitat categories.
Sample sizes are indicated at the top of the
bars.
Standardized canonical coefficients for the body measurements
First canonical variable Second canonical variable
Snout-vent length 0.837 0.529
Head height 0.814 0.340
Head length 0.887 0.427
Standardized canonical coefficients for the bromeliad measurements
First canonical variable Second canonical variable
Bromeliad height 0.950 –0.234
Bromeliad width 0.964 –0.232
Cup diameter 0.643 0.734
Canonical variables Canonical correlation F p
I 0.972 3.94 0.022
II 0.684 1.22 0.361
Table 1 - Canonical correlation among bromeliad and body measurements of Aparasphenodon brunoi (N=10).
results based on pooled stomach were similar.
Numerically, beetles were most important
(38.1%), followed by ants (27.4%), and
volumetrically, beetles were dominant (54.0%),
followed by insect larvae (20.0%) (Table 2).
Mesquita et al.
Phyllomedusa- 3(1), September 2004
55
Occurrence Stomach Means Pooled Stomachs
Prey categories f f% N %N Vol. (mm
3
) %Volume N %N Vol. (mm
3
) %Vol.
Aranae 1 2.44 0.02 ± 0.16 2.5 ± 15.81 819.36 ± 5246.49 4.17 ± 20.41 1 1.19 33593.91 7.79
Coleoptera 23 56.10 0.78 ± 0.88 43.96 ± 43.12 5680 ± 8982.79 60.48 ± 44.40 32 38.10 232915.50 54.03
Formicidae 11 16.83 0.561 ± 1.76 17.08 ± 31.01 798.95 ± 2566.64 16.12 ± 31.70 23 27.38 32756.85 7.60
Gastropoda 2 4.88 0.05 ± 0.22 1.88 ± 8.75 1105.20 ± 7076.72 2.75 ± 13.50 2 2.38 45313.10 10.51
Orthoptera 5 12.20 0.15 ± 0.42 7.44 ± 23.53 ––6 7.14 ––
Isoptera 3 7.32 0.195 ± 0.95 4.64 ± 17.16 ––8 9.52 ––
Insect larvae 4 9.76 0.10 ± 0.30 7.50 ± 24.15 2109.00 ± 8998.80 16.48 ± 37.64 4 4.76 86468.96 20.06
Non identified 6 14.63 0.15 ± 0.36 12.50 ± 31.52 ––6 9.52 ––
Plant material 1 2.44 0.02 ± 0.16 1.25 ± 7.91 ––1 1.19 ––
Insect egg 1 2.44 0.02 ± 0.16 1.25 ± 7.91 ––1 1.19 ––
Table 2. Diet composition of Aparasphenodon brunoi (N= 63).
Table 3. Canonical correlation among prey and head measurements of Aparasphenodon brunoi (N=26).
Standardized canonical coefficients for the head measurements
First canonical variable Second canonical variable
Head width 0.003 0.154
Head length 0.099 –0.186
Standardized canonical coefficients for the prey measurements
First canonical variable Second canonical variable
Maximum prey width 0.077 –0.038
Maximum prey length –0.020 0.077
Canonical variables Canonical correlation F p
I 0.330 0.70 0.594
II 0.094 0.21 0.654
The correlation between the head measure-
ments and the prey measurements were low. The
first canonical variable of the head measure-
ments showed that both measures have equal
influence on the composition of the canonical
variable while the second variable showed a
contrast between the head width and height. The
first canonical variable of the measurements of
the prey represents a contrast between the
maximum width and weight of the prey. The
first canonical correlation was 0.330 but it has
no statistical significance, showing no asso-
ciation between head and prey measurements
(Table 3).
Ecology of the casque-headed frog Aparasphenodon brunoi (Anura, Hylidae)
Phyllomedusa- 3(1), September 2004
56
The smallest adult female measured 56.32
mm SVL, whereas the smallest adult male was
48.88 mm. The largest male measured 62.42 mm
and the largest female measured 81.24 mm. We
found significant differences between sexes in
body size (ANOVA F1,25= 9.743; p=0.005). In
addition, we found significant differences
between the sexes in shape variables (MANOVA
Wilk’s Lambda = 0.444; p=0.032). To determine
which of the shape variables contributed most
to sexual dimorphism, we performed a stepwise
discriminant analysis (Tabachnick and Fidell
1996). Three shape variables were selected as
the most powerful discriminators of the two
sexes, correctly classifying 78% of individuals
(Table 4). Tibia length was the first variable
selected, correctly classifying 74% of indivi-
duals, followed by eye and tympanum diameter.
To determine whether important variables were
excluded from the analysis due to inter-
correlation with tibia length, we excluded tibia
length and repeated the analysis. Eye diameter
was then selected first, correctly classifying 70%
of the individuals, followed by SVL and
tympanum diameter. We repeated the analysis
once more with the exclusion of tibia length and
eye diameter and this time only SVL was
selected. These results indicate that besides tibia
length, eye and tympanum diameter, SVL is also
important in stating differences between the
sexes, with females having tibia, eye diameter
and SVL larger than males (Table 5).
Discussion
Aparasphenodon brunoi shows higher
Step Variable entered R2Wilk’s Lambda p < Error-rate
1 Adjusted tibia length 0.13 0.87 0.06 0.26
2 Adjusted eye diameter 0.22 0.68 0.009 0.29
3 Adjusted tympanum 0.16 0.57 0.004 0.22
Table 4 - Stepwise discriminant analysis of shape variables of Aparasphenodon brunoi. Error-rate indicates posterior
probability error-rate estimates based on cross-validation.
activity during the night, being easily found in
the Restinga Forest, outside the bromeliads.
However, in most cases, they were found within
the bromeliad leaves. Like most vertebrates,
anuran diurnal activities are highly affected by
requirements of food, mate, and shelter to avoid
predation and maintain ideal physiological
conditions, because they have a very permeable
skin, being highly susceptible to water loss by
evaporation (Hodgkison and Hero 2001).
Therefore, most anurans are typically nocturnal
(Duellman and Trueb 1994). In the case of A.
brunoi, since bromeliads retains a large amount
of water and the air humidity in the forest is
usually stable and high, these animals can show
diurnal activity, even in the sun (Silva et al.
1988). Although unusual, diurnal activities have
been reported in many anurans species, such as
Litoria nannotis (Hodgkison and Hero 2001),
dendrobatids (Zug et al. 2001), some
leptodactylids (Kwet and Di-Bernardo 1999,
Zug et al. 2001), and others (Duellman and
Trueb 1994). In this study, we observed diurnal
activity in A. brunoi in moisture days, when we
noticed some individuals calling from inside
bromeliads. However, no individual was
observed outside the bromeliads during the day.
We found a significant correlation between
frog head and bromeliad size. It has been
showed that the phragmatic behavior in A.
brunoi effectively reduces evaporative water
loss (Andrade and Abe 1997). Our data suggest
that these animals are selecting bromeliads
based on size, which could be an effort to
minimize water loss. The annual precipitation in
Restinga is high, varying from 1100 to 1500 mm
Mesquita et al.
Phyllomedusa- 3(1), September 2004
57
(Louro and Santiago 1984), but the high
permeability of sandy soils reduces the water
availability. In addition, the shape of bromeliads
promotes the accumulation of water and the
head adjustment in the bromeliad rosette is
important to maintain ideal physiological
conditions.
Some groups of amphibians are considered
dietary specialists, for example ants are the
primary diet item of dendrobatids (Toft 1995).
However, this is not the case for other groups.
For example, Rana nigromaculata (Ranidae) is
generalist and eats a high variety of prey items
dependent more so with availability than
selectivity (Hirai and Matsui 1999). Rana
catesbeiana and R. clamitans, in Michigan, also
show a highly diverse diet, eating mainly
coleopterans, hemipterans and spiders (Werner
et al. 1995). In Argentina, the diet of Pseudis
paradoxa and Lysapsus limellus (Hylidae,
Pseudinae) primarily consists of other
amphibians, beetles, mosquitoes and
Osteichthyes fishes, respectively (Duré and Kehr
2001). In a study in the Restinga of Jurubatiba,
Rio de Janeiro state, southeastern Brazil,
Aparasphenodon brunoi ate mainly beetles (Van
Sluys et al. 2004). In a study in the same area,
based on the diets of 21 individuals, A. brunoi
ate mainly ants, cockroaches and grasshoppers
(Teixeira et al. 2002). Our study shows that A.
brunoi has a very diverse diet, eating mainly
beetles, insect larvae and ants. Based on
proportion of prey use, this species could be
considered a generalist; however data on
arthropod availability in the study area and the
relationship between availability and prey
Ecology of the casque-headed frog Aparasphenodon brunoi (Anura, Hylidae)
Table 5 - Summary statistics of morphometric characters of adult Aparasphenodon brunoi. Values indicate mean ± 1
standard deviation. Size-adjusted values (see text) are in parentheses. Values are in mm.
Males (N=14)
3.535 ± 0.078
55.239 ± 3.794
(-0.010 ± 0.018)
24.388 ± 1.225
(-0.005 ± 0.014)
32.971 ± 1.832
(0.011 ± 0.013)
19.841 ± 1.125
(-0.011 ± 0.019)
18.221 ± 1.168
(-0.000 ± 0.017)
4.135 ± 1.532
(-0.019 ± 0.096)
6.393 ± 0.323
(-0.002 ± 0.025)
24.680 ± 1.428
(-0.002 ± 0.019)
Females (N=14)
3.663 ± 0.125
65.176 ± 8.244
(0.015 ± 0.054)
28.459 ± 2.725
(0.016 ± 0.038)
35.635 ± 9.627
(-0.041 ± 0.232)
24.843 ± 9.382
(-0.014 ± 0.084)
20.938 ± 2.158
(-0.010 ± 0.045)
4.078 ± 0.281
(-0.006 ± 0.035)
6.736 ± 0.490
(-0.019 ± 0.024)
28.446 ± 2.723
(0.009 ± 0.041)
Character
Body size
Snout-vent length
Tibia length
Foot length
Head length
Head width
Tympanum diameter
Eye diameter
Forearm length
Phyllomedusa- 3(1), September 2004
58
selectivity are needed to support this statement.
Still, it will be necessary more samples throughout
the year to avoid the seasonality effects.
Several authors have indicated that
relationships between prey and head
measurements could be related to resource
partitioning between sexes and/or species
(Magnusson and Silva 1993, Van Sluys et al.
2001). Teixeira et al. (2002) did not find any
correlation between prey and body size,
however they did not use any statistical test to
support this evidence. In our study, based on a
canonical correlation, no significant relationship
between prey and head measurements was found
in Aparasphenodon brunoi. These relationships
are likely more related with resource
partitioning among species in an assemblage
rather than due to size variation within the same
species (Magnusson and Silva 1993).
Three main hypotheses have been proposed
to explain the existence of sexual dimorphism
in frogs. Several studies have found that when
females are larger than males there is a
significant relationship between female SVL and
clutch size (Kuramoto 1978, Kaplan 1980,
Duellman and Trueb 1994). We found statistical
differences in body size between sexes in A.
brunoi, with the females being larger than
males; however, we did not collect clutch size
data for A. brunoi, and we are thus unable to
access this hypothesis.
Another hypothesis to explain sexual
dimorphism could be related to male-male
competition for mates, with larger males
benefiting with this kind of dimorphism
(Duellman and Trueb 1994). Alternatively,
sexual size dimorphism can be a mechanism for
reducing intersexual competition for food
resources (Magnusson and Silva 1993, Van
Sluys et al. 2001), where the difference in the
head size between sexes leads to a difference in
prey size consumed by each sex. However the
canonical correlation between head and prey
size was not significant. Furthermore, this
pattern could be due to resource sharing among
species of an assemblage rather than difference
between sexes (Magnusson and Silva 1993). In
our study area, there are four other species that
use bromeliads in their life cycle (Hyla
albomarginata, Scinax altera, S. cuspidatus and
Trachycephalus nigromaculatus) (Teixeira et al.
2002), which could interact with and influence
A. brunoi ecology.
Teixeira et al. (2002) stated that sexual size
dimorphism occurred in A. brunoi with females
being larger than males; however their statistical
tests do not support this affirmation. Based on
univariate and multivariate analyses of variance
on a set of morphometric variables we found
sexual dimorphism in A. brunoi. Nevertheless,
a study including a larger sample size and more
detailed ecological and life history data are
needed to elucidate the factors that have led to
sexual size dimorphism.
Acknowledgements
We thank Alexandre Bamberg Araújo,
Mirella A. Kacinskis and Mariana F. P. de
Araújo for help with the fieldwork; Antônio
Sebben, Reuber A. Brandão, Janalee P.
Caldwell, Adrian A. Garda, Donald B. Shepard
and Alison M. Gainsbury for insightful
comments on a previous version of the
manuscript and for help in English. This work
was supported by a doctorate’s fellowship from
Coordenação de Aperfeiçoamento de Pessoal de
Nível Superior – CAPES to DOM and by a
master’s fellowship from Conselho Nacional de
Desenvolvimento Científico e Tecnológico–
CNPq to GCC.
References
Andrade, D. V. and A. S. Abe. 1997. Evaporative water
loss and oxygen uptake in two Casque-Headed tree
frogs, Aparasphenodon brunoi and Corythomantis
greeningi (Anura, Hylidae). Comparative
Biochemistry and Physiology 118A: 685–689.
Argôlo, A. J. S. 2000. Aparasphenodon brunoi: Geogra-
phic distribution. Herpetological Review 31: 108.
Cogliatti-Carvalho, L., A. F. N. Freitas and C. F. D. Ro-
Mesquita et al.
Phyllomedusa- 3(1), September 2004
59
cha. 2001. Variação na estrutura e na composição de
Bromeliaceae em cinco zonas de restinga no Parque
Nacional da Restinga de Jurubatiba, Macaé, RJ. Re-
vista Brasileira de Botânica 24: 1–9.
Duellman, W. e L. Trueb. 1994. Biology of Amphibians.
Baltimore and London. The Johns Hopkins University
Press. 670 pp.
Duré, M. I. and A. I. Kehr. 2001. Differential exploitation
of trophic resources by two pseudid frogs from
Corrientes, Argentina. Journal of Herpetology 35:
340–343.
Eterovick, P. C. 1999. Use and sharing of calling and
retreat sites by Phyllodytes luteolus in a modified
environment. Journal of Herpetology 33: 17–22.
Feio, R. N., U. M. L. Braga, H. C. Wiederhecker, and P.
S. Santos. 1998. Anfíbios do Parque Estadual do Rio
Doce (Minas Gerais). Viçosa, Universidade Federal de
Viçosa.
Frost, D. R. 2002. Amphibian Species of the World: an
online reference. Vol. V2.21 (15 July 2002), http://
research.amnh.org/herpetology/amphibia/index.html.
Hirai, T. and M. Matsui. 1999. Feeding habits of the pond
frog, Rana nigromaculata, inhabiting rice fields in
Kyoto, Japan. Copeia 1999: 940–947.
Hodgkison, S. and J. M. Hero. 2001. Daily behavior and
microhabitat use of the waterfall frog, Litoria nannotis
in Tully Gorge, Eastern Australia. Journal of
Herpetology 35: 116–120.
Jolicoeur, P. 1963. The multivariate generalization of the
allometry equation. Biometrics 19: 497–499.
Kaplan, R. H. 1980. The implications of ovum size
variability for offspring fitness and clutch size within
several populations of salamanders (Ambystoma).
Evolution 34: 51–64.
Kuramoto, M. 1978. Correlations of quantitative parame-
ters of fecundity in amphibians. Evolution 32: 287–
296.
Kwet, A. and M. Di-Bernardo. 1999. Pró-Mata: Anfibios.
Amphibien. Amphibians. Porto Alegre. EDIPUCRS.
107 pp.
Leme, E. M. C. 1984. Bromélias. Ciência Hoje 3: 66–72.
Louro, R. P. and L. J. M. Santiago. 1984. A região de Barra
de Maricá-RJ e a importância de sua preservação. Atas
da Sociedade Botânica do Brasil 2: 109–118.
Magnusson, W. E. and E. V. Silva. 1993. Relative effects
of size, season and species on the diets of some
amazonian Savanna lizards. Journal of Herpetology
27: 380–385.
Pombal, J. P., Jr. 1993. New species of Aparasphenodon
(Anura: Hylidae) from Southeastern Brazil. Copeia
1993: 1088–1091.
Rohlf, F. J. and F. L. Bookstein. 1987. A comment on
shearing as a method for “size correction”. Systematic
Zoology 36: 356–367.
Silva, H. R., M. C. Britto Pereira, U. Caramaschi, and R.
Cerqueira. 1988. Utilização de Neoregelia cruenta
(Bromeliaceae) como abrigo diurno por anfíbios
anuros na restinga de Maricá, Rio de Janeiro. Pp. 307–
318 in Anais do VI Seminário Regional de Ecologia,
São Carlos, SP.
Somers, K. M. 1986. Multivariate allometry and removal
of size with principal component analysis. Systematic
Zoology 35: 359–368.
Suguio, T. and M. G. Tessler. 1984. Planícies de cordões
litorâneos quaternários do Brasil: origem e nomencla-
tura. Pp. 15–25 in L. D. Lacerda, D. S. D. Araújo, R.
Cerqueira, and B. Turcq (eds.), Restingas – origem,
estrutura e processos. Niterói, CEUFF.
Tabachnick, B. G. and L. S. Fidell. 1996. Using
Multivariate Statistics. New York, HarperCollins
Publishers Inc.
Teixeira, R. L., J. A. P. Schineider and G. I. Almeida. 2002.
The occurrence of amphibians in bromeliads from a
southeasthern Brazilian Restinga habitat, with special
reference to Aparasphenodon brunoi (Anura, Hylidae).
Brazilian Journal of Biology 62: 263–268.
Teixeira, R. L., C. Zamprogno, G. I. Almeida, and J. A. P.
Schineider. 1997. Tópicos ecológicos de Phyllodytes
luteolus (Amphibia, Hylidae) da restinga de Guriri São
Mateus-ES. Revista Brasileira de Biologia 57: 647–
654.
Toft, C. A. 1995. Evolution of diet specialization in
poison-dart frogs (Dendrobatidae). Herpetologica 51:
202–216.
Trueb, L. 1970. Evolutionary relationships of casque-
headed tree frogs with co-ossified skulls (family
Hylidae). Publications of the Museum of Natural
History of the University of Kansas 18: 547–716.
Van Sluys, M., C. F. D. Rocha and M. B. Souza. 2001.
Diet, reproduction, and density of the leptodactylid
litter frog Zachaenus parvulus in an Atlantic rain
forest of southeastern Brazil. Journal of Herpetology
35: 322–325.
Van Sluys, M., C. F. D. Rocha, F. H. Hatano, L.
Boquimpani-Freitas, and R. V. Marra. 2004. Anfíbios
da restinga de Jurubatiba: composição e história na-
tural. Pp. 165–318 in C. F. D. Rocha, F. A. Esteves
and F. R. Scarano (eds.), Pesquisas de Longa Dura-
ção na Restinga de Jurubatiba - ecologia, história
natural e conservação. RiMa, São Carlos, SP.
Vrcibradic, D. and C. F. D. Rocha. 1996. Ecological
differences in tropical sympatric skinks (Mabuya
macrorhyncha and Mabuya agilis) in Southeastern
Brazil. Journal of Herpetology 30: 60–67.
Ecology of the casque-headed frog Aparasphenodon brunoi (Anura, Hylidae)
Phyllomedusa- 3(1), September 2004
60
Werner, E. E., G. A. Wellborn and M. A. McPeek. 1995.
Diet composition in postmetamorphic bullfrogs and
green frogs: implications for interspecific predation
and competition. Journal of Herpetology 29: 600–607.
Zar, J. H. 1998. Biostatistical Analysis. Englewood Cliffs,
Prentice-Hall, Inc.
Zug, G. R., L. J. Vitt and J. P. Caldwell. 2001.
Herpetology – an introductory biology of
amphibians and reptiles. San Diego, Academic
Press.
Mesquita et al.