D
iet
of
D
enDropsophus
microcephalus
anD
s
carthyla
vigilans
(a
nura
: H
yliDae
)
at
a
locality
in
nortH
-
western
V
enezuela
witH
notes
on
microHabitat
occupation
K
imberlyn
a. f
onseca
-p
érez
¹²³
c
ésar
m
olina
¹
†
z
aiDa
t
árano
²⁴
ABSTRACT
The coexistence of species with similar ecological requirements (food, space, time) has typically
drawn attention of researchers because competition for resources is expected to be high. The diet and
microhabitat occupation of two morphologically and ecologically similar species of Hylidae (Anura),
Dendropsophus microcephalus and Scarthyla vigilans, were analyzed at a locality in north-
western Venezuela, with the aim of addressing the potential for space and food competition between
them. Diet was estimated through the analysis of stomach contents and microhabitat occupation
was estimated through recordings of perch type, height and horizontal distance to water. Thirty-six
prey categories (32 families, 4 orders) of arthropods were identified: 30 categories in D. microceph-
alus and 21 categories in S. vigilans. The most important prey (RII) in D. microcephalus were
Agelenidae (11.1%), Tachinidae (9.32%) and Lepidoptera-larvae (7.96%). Gryllidae (14.13%),
Cicadidae (9.1%), Cicadellidae (8.3%) and Delphacidae (8.02%) were the most important in
S. vigilans. Diet overlap was relatively low (0.32). Both species have relatively generalist diets. Both
species occupied the same type of perches (leaves and stems of Dicotyledons and Monocotyledons) and
heights (average: S. vigilans,
24 ± 16.2 cm; D. microcephalus,
22.7 ± 9.5 cm). The potential for
space competition is high if perches are limited and food competition is expected to be low.
Key-Words: Diet; Generalist; Niche overlap; Resource partitioning; Microhabitat; Mor-
phometry.
INTRODUCTION
Morphological and ecological similarity between
species is believed to hinder their coexistence because
competition for resources is likely. It is thought that
at least one ecological difference in resource use be-
tween them (i.e., food, time or space partitioning) is
necessary to allow coexistence (Pianka, 1994; Gor-
www.mz.usp.br/publicacoes
www.revistas.usp.br/paz
ISSN impresso: 0031-1049
ISSN
1807-0205
on-line:
Museu de Zoologia da Universidade de São Paulo
Papéis Avulsos de Zoologia
http://dx.doi.org/10.11606/0031-1049.2017.57.07
Volume 57(7):93‑104, 2017
1. Laboratorio de Biología y Conservación de Anfibios, Instituto de Zoología y Ecología Tropical, Facultad de Ciencias,
Universidad Central de Venezuela.
2. Laboratorio de Comportamiento Animal, Instituto de Biología Experimental, Facultad de Ciencias, Universidad Central de Venezuela.
3. E-mail: kimbi415@gmail.com.
4. Correspondence autor. E-mail: zaida.tarano@ciens.ucv.ve.
† This is a posthumous publication for César Molina.
don, 2000; Vignoli & Luiselli, 2011). The relative
importance of food, time and space use in structur-
ing animal communities varies from one another and
between habitats. Several authors have suggested that
the space dimension is often more important than
the food dimension, and that the latter is often more
important than the temporal dimension (Schoener,
1974; Giller, 1984); however, this ordering is by no
means universal. For instance, food is the main di-
mension in structuring several anuran communities
(Toft, 1980a,b; Lima, 1998), while microhabitat is in
others (e.g., Crump, 1974; Toft, 1985; Cardoso et al.,
1989).
Amphibians, and specially anurans, are remark-
ably abundant in tropical ecosystems and have been
considered extremely important in food webs and
energy flow (Stebbins & Cohen, 1995). Nonethe-
less, there are relatively few studies of feeding pref-
erences and behavior in this group (Toft, 1980a,b;
Duellman, 1993; Piñero & Durant, 1993; Lima &
Magnusson, 1998; Caldwell & Vitt, 1999; Parmelee,
1999), and most of them have focused on a limited
number of taxa. With regards to habitat occupation,
available literature indicates that in general, there are
substantial differences in microhabitat and activity
periods both within and among species (Schoener,
1974; Drewry & Rand, 1983; Toft, 1985; Muñoz-
Guerrero et al., 2007; Tárano, 2010). Several stud-
ies with hylids have demonstrated that microhabitat
segregation is associated to body size (Bevier, 1997),
which in turn has a strong impact on the diet and
prey-capture behavioral strategies (Toft, 1980a,
1981).
In anurans, diet composition is usually related
to body size, sex, and habitat and microhabitat pref-
erences (Toft, 1980a,b; Christian, 1982; Woolbright
& Stewart, 1987; Piñero & Durant, 1993; Bevier,
1997; Hirai & Matsui, 2000). The diet typically
changes with age (e.g., Labanick, 1976; Christian,
1982; Strussmann et al., 1984; Woolbright & Stew-
art, 1987; Donnelly, 1991; Wiggins, 1992), season
(da Rosa et al., 2002) and the size and behavior of
preys (Freed, 1980; Lima, 1998; Parmelee, 1999).
Since anurans swallow whole prey, mouth width pos-
es an upper limit to the maximum size or volume of
prey. Therefore, as an individual grows, the maximum
size of its preys may increase concomitantly (Lima &
Moreira, 1993; Parmelee, 1999). In general, anurans
that consume relatively small and slow-moving prey
have narrow mandibles and symmetric feeding cycles
(i.e., the time devoted in capturing is similar to that
devoted in retrieving to the mouth). On the other
hand, anurans feeding on relatively large slow-moving
prey have wide mandibles and asymmetric feeding
cycles (Emerson, 1985).
Most anurans analyzed so far feed upon inverte-
brates as adults while a few also prey upon small verte-
brates (Duellman & Trueb, 1994). A great majority of
the anurans analyzed have been labeled as food-gener-
alists based on estimations of diet richness and equi-
tability, despite the fact that prey availability has not
been estimated in most studies (but see Toft, 1980b,
1981; Christian, 1982; Hirai & Matsui, 2000). With
regards to prey specificity, anurans can be arranged
in a continuum ranging from ant specialists through
non-ant specialists to generalists (Toft, 1981).
In the present study we aimed to describe the
diet and microhabitat occupation of two hylid frogs
of similar morphology which occur syntopically over
a wide range in northern Venezuela, Scarthyla vigilans
and Dendropsophus microcephalus. Scarthyla vigilans is
an arboreal anuran traditionally thought to be restrict-
ed to the Maracaibo Lake basin in northwestern Ven-
ezuela (Barrio-Amorós, 1998). It is currently known
for inhabiting the northern Caribbean lowlands, the
Magdalena River basin in Colombia and the llanos
of Colombia and Venezuela (Barrio-Amorós et al.,
2006; Lotzkat, 2007; Rojas-Runjaic et al., 2008). The
species is currently expanding into the Orinoco River
Delta (Rojas-Runjaic et al., 2008) and Trinidad and
Tobago (Smith, J.M. et al., 2011). Dendropsophus mi-
crocephalus has been regarded as widely distributed in
Venezuelan lowlands (Barrio-Amorós, 2009). There-
fore, both species coexist in vast areas of their distri-
bution providing opportunity to assess the potential
for food and space segregation. Previous studies in
Colombia have documented similar microhabitat
preferences (Lomolino et al., 2006, Muñoz-Guerrero
et al., 2007; Armesto et al., 2009), overlapping diets
(Muñoz-Guerrero et al., 2007) and partially disjoint
activity patterns throughout the rainy season (Mu-
ñoz-Guerrero et al., 2007). Nonetheless, so far there is
scarce information on the habits of both species. With
this study we aimed to add to the comprehension of
the coexistence of D. microcephalus and S. vigilans and
to address potential regional differences.
METHODS
Study Site and Subjects
We performed the study at Hacienda La Guá-
quira (10°20’4”N, 68°39’17”W), in the mountain
complex Macizo de Nirgua, at the western-most edge
of the Coastal Mountain Chain (Cordillera de la Cos-
Fonseca-Pérez, K.A.
et
al
.: Diet and microhabitat Hylinae
94