Amphibia, Anura, Hylidae

Bijdragen
tot de
calls
Advertisement
of Bolivian
64
Dierkunde,
SPB Academie
75-85
(2)
The
Publishing bv,
species of Scinax (Amphibia,
(1994)
Hague
Anura,
Hylidae)
de la
Ignacio
Rafael
Riva
de
Departamento
Museo Nacional de Ciencias Naturales,
Biología Evolutiva,
Anura, Hylidae,
Keywords:
& Jaime Bosch
Márquez
Scinax,
advertisement
Bolivia,
Abstract
Myers, 1991;
advertisement
described
species.
A characteristic
for
the
each
and
phenograms
based
constructed,
the first
correlation, a
single
new
the
calling behaviour
as
well
temporal
and
one
the
using
a
of
traditional
second
technique that
one
are
of each
oscillogram
numerical
as
features
the characteristics of the
on
and
nique (UPGMA)
on
of Scinax
species
audiospectrogram
species,
spectral
Bolivian
eight
including information
presented
about
calls of
are
Duellman & Salas,
countries,
contains
of the least known
its
areas, and
anuran
been the
discoveries
Riva,
of several
subject
1990a,
recent
1992a, 1992b,
1990b,
the
1991). Among
Bolivia
some
fauna has
(De
la
1993; Reynolds
information
the
calls.
mating
Two
calls
multivariate
& Foster,
comparisons
1992).
In this
are
tech-
using audiospectrogram
allows holistic
Madrid, Spain
calls, ecology, phylogeny
South American
The
28006
study
Bolivian
we
contribute
herpetofauna
by
to
the
the
presenting
teristics of the advertisement calls of
of
knowledge
charac-
eight species
in
of
the
genus Scinax,
hylid
known
previously
Ololy-
as
vocalizations.
gon
(see
ment
Duellman &
calls of
Wiens,
of these
two
1992).
species
The advertise-
are
described for
the first time.
Resumen
The
Ololygon
as
Se describen
Scinax
de
miento
las llamadas
Un
fonador.
presentados
para
cas
Se presentan
de las
fenogramas
un
ocho
la
y
el
en
known
was
1843.
Fitzinger,
son
numérica
de las vocalizalas característi-
rected from the synonymy of
Delahoussage
Although
by
some
cal
the
validity
authors,
features
based
(1977),
de los
una
novedosa
archivos de
imagen
the
all
by
Wiens,
a
only
to
being
America
are
studies that
the richness
a
most
diverse
anu-
great number of spe-
described each
year. Central and South
the
subject
provide
in
an
anuran
of
a
ever
questioned
morphologi-
species
in
this
nomenclature for
1992). Among
be
Scinax
other
membrane between
number of
more
diversity
exploratory
complex
(see
view of
Donnelly
&
which
toe.
are
& Wiens,
The
and
genus Scinax is
(1985)
Ololygon)
and
II
or
lack-
having
on
have
truncate
disks,
the hands (Duellman
1992).
conspicuous
Frost
long,
fea-
morphological
I
the
Duellman
the inner margin of the
Additionally, they
wider than
(see
Hyla mainly by
toes
vestigial fringe along
a
second
contain by far the
fauna in the world, and
the
&
characters.
sperm
characteristic
correct
considered
was
genus
&
Introduction
ran
years
resur-
Hyla by Fouquette
of this feature was
some
shared
are
Recently
genus.
ing
cies is
was
metodologíamultivari-
audioespectrogramas.
Neotropics
many
Ololygon
on
tures, this genus differs from
The
for
de
comporta-
oscilograma
segundo usando
correlación
especies
su
información
basados
de ellos usando
(UPGMA),
en
de
sobre
temporales y espectrales
uno
metodología basada
información
especie junto con
dos
llamadas,
ante tradicional
de los
apareamiento
audiospectrograma y
cada
sobre las características
ciones.
de
Bolivia, aportando
genus Scinax
groups
reports
and
one
of
54
several
of
the
frogs
species
more
most
in the
in
have
abundant
Neotropics.
the
been
genus
(as
described
/.
76
since
then.
in
present
De
la
Bolivia, although
The
1993).
has
genus
groups. The
present study belong
different
been
bly
S.
S.
chiquitana,
include S.
is the S.
and
is the S.
groups
the S.
only
well
to
Scinax from Bolivia
published
America,
point
to
Finally,
generate phenetic
multivariate
potential
group
an
to
within
species
Scinax.
from the
pared
taxonomie
South
tax-
research.
used
are
to
The
The
method is
first
a
method is
the second
based
technique
of
relationships
some
and
technique,
innovative
of
areas
methodologies
discuss
audiospectrograms.
previously
of affinities among vocalisa-
trees
tions which allow
with
inconsistencies in
of
areas
of
eight species
from other
different
two
the
parkeri
Bolivian Scinax
of
compared
possible
to
future
or
onomy,
and S. nasica. The
group
Calls
are
descriptions
group, that would
which S. nasica has been
to
the correlationof
on
relationships
resulting
study
of the advertisement calls
with the
currently accepted species
are
com-
groups.
authors. Of these four
some
rostrata
have
to
group appears
consistency
phylogenetic
proba-
as
nebulosa. The third
group
assigned according
solid
(possibly four)
as
The fourth
S. nasica.
x-signata
several
group which includes S.
staufferi
possibly
rubra,
rostrata
and S.
garbei
into
first group is the S.
fuscovaria,
second group is the S.
1990b,
considered in the
least three
rubra group that includes S.
species
-
al.
et
In the present paper, the calls of
species
Riva,
divided
groups. The
species
la
(De
eight species
to at
eight
additional
two
have been described since then
phenetic
lists
(1990)
Riva
De la Riva
Duellman &
(see
Material and
methods
Wiens, 1992, for further details on the taxonomy of
the
cies
These four groups include all
group).
considered in the present
which has
castroviejoi,
group (De la Riva,
Since the
study except
been
not
Recordings
eight
assigned
spe-
for
to
S.
any
Puerto
of sound
mating
analysis techniques,
calls have been considered
to
Almacén,
a
(Tosi
Sony
heiser
anuran
isolating
to
their role
mechanisms
pre-zygotic
species
1955, 1958,
1959; Jameson, 1955; Johnson,
as
(Blair,
1959).
recording
seconds)
from
anuran
studies
ecological
of
tropical
the
(Hödl,
or
for each
analysed
species.
characteristics
temporal
for
analysis
tive individual
1965,
Drewry
&
either
Rand,
from taxonomie stand-
"normal"
also based
individual
to the
the
on
and
in
human
general
calls of
genus
terms,
anurans
but
level,
may have
do
it
is
not
accepted
show
that
the
homologies
mating
above the
similarities among calls that
some
phylogenetic
significance
at
generic
and
a
analysis
the
level may be
vocalisations in
phylogenetic
limited and depends
sidered.
According
on
to
natural
phology
are
of
as
the
the group of
to
a
a
is,
species
partitioning
in
con-
which is
used
Tools
to obtain
mor-
convergences
of different fami-
allopatric
and
(Duellman
&
seems
Pyles,
to
a
on
the
each
For
single representa-
and
a
points).
The
tisement
terminology used
calls follows
A total
and
16 bit
resolution
and
to
Frequency
transform
rate
(pulses
frequency,
mum
call,
number
number
note
number
of
al.
generate
per
second),
information
of notes per
pulses
per
repetition
different
call,
with
types
call
(notes
of
were
were:
of the
1024
adver-
notes,
minute
and
a
fundamenmini-
of notes per
rate
within
ratio
in
duration,
energy,
number
repetition
per
compared
note
frequency,
substantial
maximum
note,
rate
was
width
(FFT,
description
dominant
frequency
was
audio-
(1990).
The variables considered
other
of
minute),
for the
et
with
software
Signalyze
characteristics
of 12 different call
phenetic analysis.
pulse
Heyer
digital sig-
editing were completed at
information
Fourier
be
were
of the recorded
Macintosh-based
and
kHz
appeared to
for the choices
recording.
software.
oscillograms.
fast
which
(calls
the
pulse
per
call),
dura-
anuran
have similar characteristics
1983).
with
of 44.1
numerical
through
of the
Digitalisation
hardware
spectrograms
tal
of the
system.
tion/interpulse interval.
communities
of
of
however,
anurans
product
However,
species."
acoustic
validity
certain extent,
the voice is
often found between
lies and
The
studies
Schiotz (1973:313) "the voice
follows the taxonomy
quite
expected.
information
of the identification
quality
sampling frequency
Sound
obtained
infrageneric
the calls
The criteria
ear.
certainty
1966).
nal
In
audio-
2.5 seconds
a
of the sounds.
which emitted vocalisations
Recordings were processed
points (Barrio,
for
either
Senn-
a
longer recording (20-60
A
generate numerical
to
and
representative
presented
were
subtropical
included
recorder
A
microphone.
in
Department
in wet
locality
Ml 120 tape
Sanyo
a
oscillogram
selected
we
of Santa Cruz
part
from
recorded
were
fauna allows for
vocalisations,
1977;
1983; Zimmerman, 1983)
or
segment
and
first author
the
by
specified, calls
Amazonian
an
directional
and
was
spectral
species,
The richness of the
comparative
80
Me
Bolivia
1975). Recording equipment
WM D6C
be im-
in
otherwise
northwestern
al.,
et
spectrogram
portant taxonomie determinantsdue
obtained
were
1990. Unless
(15°46'S 62°15'W),
forest
1993).
inception
1987 to
the
original
different
matrix
variables.
in
Variables
order
to
were
linearly transformed
standardise
The standardised matrix
the
was
scales
of
in
the
used to calcu-
Bijdragen
late the
The
the
pair-group-method arithmetic
calculated
ously
The
1992).
to those
species
for the
from
Hyla
1994
-
of
application
statistical
(NTSYS
Bolivia
by Márquez
the
package,
al.
et
notes
Both
addition
analysis,
UPGMA
to the
Rohlf,
of 18
calls
number of
(891
Hz), and
(2400
(1993).
second
a
animal
on the
audiospectrograms
the
comparison
advertisement
allows
(1987)
of the
This
call.
for
al
acoustical
process
was
Ithaca,
gy,
features.
available
and
To the
of the vocalisations
ment call
was
was
used
composed
for
note which
in
of
more
was
never
the
Museo
de
de la
than
the
the
Lab.
of
a
it is the
the
or
note
the loudest
individuals
were
"Noel
as
the
note).
depositand/or
calls of this
the
the advertisement calls
1
Figs.
oscillograms
of
shown in Table I. The
the
sound
2.5
a
species
and 2. A summary of the
merical information from
Scinax
UPGMA
32.0
correlation
and
shown in
are
trees
by
S. rubra group
This
castroviejoi (De
described
recently
(maximum
Riva,
length
snout-vent
perate valleys of
the
from
northern
Bolivia
recorded in
Santa
Cruz,
migrated
on
a
to
to
small
probably
rain
a
during
and males
lagoon
Bolivia,
the shore
small bushes
or
was
were
two
at
(SVL):
slopes
dusk,
explosive
an
days
and from
locality
1990a)
la
SVL:
males
mm) probably belongs
Riva, 1990b)
and
the
to
occurs
in the
breeder,
Puerto Almacén it
At
1991).
forming large
the
rainy
calls
bushes
or
near
of
males
in
occurs
of
41.9
tem-
the Andes
Calls
were
of
Males
63°38'W).
and called
Scinax
pond
calling.
perched
castroviejoi
it
did
was
heard
only
were
aggrega-
However,
season.
The
single type
(243 ms), pulsed,
intervals
(Fig.
lb).
of 2172 Hz.
was
The
No
on
aquatic
temporal ponds
advertisement call
of
were
repeated
at
dominant
mean
or
was
of intermediate
note
and
previous
species
perched
from
emerging
vicinity.
a
sporadically.
called while
usually
duration
regular
frequency
records of the vocali-
available.
fuscovaria (Lutz,
is
not
visited
out
of
1925)
large species (maximum
1985).
Scinax
of
breeder because
active and
the
(1993) described
same
la Riva,
during
females 53.2 mm)
Bermejo (Dept.
when the
(De
tions of males
by
1993)
mm)
18°97'S
of its
is
analyses
species
near
because
note
species (maximum
Salas,
nu-
produced
Argentina.
rushes.
prolonged
the
eastern
&
(Duellman
audiospectrogram
large
female known 48.7
single
mm,
la
in the sequence
rain forests of northern Bolivia and southern Peru
It
the
occurs
&
Brazil,
breeder
fuscovaria
was
directly
short
on
second)
succession
provided
trunks
and
(Fig. lc).
at
regular
basic
Bolivia.
At
explosive
an
temporal ponds
small trees,
or
or
a
low
The call
one
pulse
was
intervals. De
parameters
usually
thick branches. The
and consisted of
frequency (860 Hz)
per
forests
rains.
Males called from bushes
call
and
was
choruses in
forming large
following heavy
through-
subtropical
Argentina, Paraguay,
Almacén S.
perching
1977; Frost,
Delahoussaye,
and the
mm,
in the S. rubra
in open and forest habitats
Chaco, Cerrado,
Puerto
SVL: males 50.2
probably belongs
(Fouquette
group
Scinax
la Riva
from the
females 35.3
sations of this
3 and 4.
Figs.
primary
chiquitana (De
mm,
Scinax
phenetic
analyses
De
species
This medium-sized
This
the
note
recordings.
same
in their
second section of the calls of the different
shown in
the
as
greater intensity.
composed
are
frequency
quantitative compari-
with other calls the second
mating
and
a
Kempff Mercado",
Results and discussion
audiospectrograms
For the
Hz).
considered
plants
The
dominant
relatively high
a
and 2850
Males
Description of
with
milliseconds),
primary note
primary
the
advertise-
Bolivia.
Sierra,
28
low fundamental frequency
pulses,
correlation
Tropicales, Sevilla, Spain,
Natural
al.
quantitativecompari-
omitted from the call
Historia
et
of Ornitholo-
knowledge,
onenote,
specified, collected
the Centro de Estudios
Santa Cruz
for
our
defined
of
notes
Clark
of different taxa. When the
comparisons (we
Unless otherwise
ed in
best of
in
similar
two
based
representation
used
applied to
single
and
(26
of
rapid succession (Fig. la).
rather than of individu-
"Canary" (Cornell
first time that this method is
son
full
a
frequency
software
The
in
York).
New
of
comparison
a
in time
sound's structure
of
described
method,
used,
was
in
for
method
was
vocalisations
quantitatively comparing
consisted
always
together
short
were
high
sons
In
call
mating
emitted
previ-
similar
were
of advertisement
comparison
phenetic
unweighted
on
used
The
species pair
an
(UPGMA)
average
77
unrooted
resulting
and the parameters
methodology
used
of
matrix
(2)
distance" between each
Sokal, 1973).
generated through
was
tree
taxonomie
"average
Sneath &
(after
de Dierkunde, 64
tot
of the
note
rate
with low
(50 pulses
emitted in fast
la
Riva
calls
(1993)
of S.
fus-
/.
78
Fig.
1. From bottom to top:
oscillogram
showing
and
audiospectrogram
the temporal
details
of the
oscillogram
of
notes; a, Scinax
a
2.5 second
De la Riva
al.
Calls
-
Scinax
chiquitana,
of
Scinax
c,
Bolivian Scinax
mating call. Top: expanded
of a characteristic
recording
castroviejoi; b,
et
fuscovaria; d,
Scinax
garbei.
covaria from Puerto Almacén which
with
our
are
coincident
description.
3715 Hz and
(Fig. Id).
of variable
Scinax
This
garbei (Miranda-Ribeiro,
is
a
medium-sized
males 39.8 mm, females 42.1
group
(Duellman,
upper Amazon
Apparently,
tween
cén S.
rainy
mm)
is
of the S.
there
was
in
the
Bolivia.
marked differences in size be-
are
a
rostrata
widespread
to
SVL:
1977).
prolonged
At Puerto Alma-
breeder
the
during
leaves
called
near
ponds.
Ecuador
the
perched
the shore
of
head-down
temporal
The call consisted of
ms), highly pulsed,
with
a
on
or
one
or
semi-perma-
long
dominant
trunks
note
(950
frequency
of
Scinax
to
be
call
as
being
of
whereas
for the
it
to
of 14
our
the
Ecuadorian
markedly
much
rate
intervals
ms.
described
from Santa
1972, Duellman considered
as
Hyla).
shorter
agreement
notes
1978,
He described the
is
nearly
duration of the
frogs (range
than the
The
either;
note,
per minute. The
recordings
295—1469 ms).
the
Cecilia,
single, moderately long pulsed
at a rate
repetition
(range
a
of 1695 Hz
in the genus Garbeana; in
Duellman referred
repeated
1300
1978)
garbei
1970a and
(in
species
1972,
(1970a,
call of
mating
figure,
season.
Males
nent
1972).
It
Basin from Colombia
populations (Heyer,
garbei
(maximum
species
frequency
separated by long
were
duration, averaging
Duellman
1926)
fundamental
a
The calls
note
calls
reported
160-260
duration
pulse
rates
from
ms)
is
by
us
found
do
note
twice that
not
show
Ecuador
had
Bijdragen
Fig.
2. From bottom to
top:
oscillogram showing
pulse
from 53
that
cy:
to
shows
57
3250
ranging
study.
Hz
In
some
one
The
ranging
rates
only parameter
be-
Duellman
to
3958 Hz
Fig.
3
in
the
pulse
present
rate
and
provided (Fig.
in Duellman,
our
wide, flat, frequency bands,
in
frequency
at
the
1
1972)
Fig.
beginning
on
Pyles
the calls of S.
Santa Cecilia
(1983)
provided
previously
in
the
rate
call
(Ecuador). Curiously enough,
site
from 80
to
240
from
from 54.5
the
be
to
from 1564
to
most
either as
to
650
1978; pulse
1978;
rate,
1978;
in 1983
ranging
dominant
frequency,
1926 Hz in 1983 and estimated
garbei
our
do
of the numerical parameters
calls of Ecuadorian
reported by
ms
in 1983 and from 195
1978. In any event,
described later
repeti-
note
14 notes/minute in
in
in
obtained
discrepan-
The
72.3 notes/minute in
from 370
calls from Bolivian S.
mating
as
ms
to
pulses/second
be 3250 Hz in
recordings
1978).
following parameters:
pulses/second
mating
or
100
Scinax rubra.
within the ranges of the
data for
(Duellman,
reported
to
mating call. Top: expanded
parkeri; d,
not even
and from 160-260
to
several
c, Scinax
duration, ranging
the
of
characteristic
are
ranging
1983 and
a
published
same
ranging
numerical
garbei (then Ololygon)
of
nebulosa,
of the variables
with
Duellman &
recording
Scinax
Id,
the call.
data
2.5 second
nasica; b,
tion
and
(1978)
a
cies include the
is the dominant frequen-
of the differences in
two
of
Scinax
pulses/second,
pulse
audiospectrograms
raising
oscillogram
dramatic differences
to
1970 and
and
79
of the notes; a,
240
to
1994
-
structural similarities with
emphasising
upper
less
described,
spite
in Duellman,
195
pulses/second.
according
the
details
showed
from 3453 Hz
duration,
show
from
slightly
(2)
audiospectrogram
temporal
recordings
the calls
tween
the
ranging
rates
but Bolivian
64
Dierkunde,
de
tot
frogs
Duellman &
on
the
coincide
reported
of this
Duellman (1970a,
by
data
not
for
species,
1972,
1978)
Pyles (1983).
/. De la Riva
80
Further
research
variable
taxonomie
highly
information about
provide
could
species
actual
the
and
widespread
this
on
note
of 58
at
of different popula-
status
The
rate
had
This
is
37.0
(Cope,
1862)
medium-sized
a
to
of
mm)
which has
species
(maximum
phylogenetic
uncertain
been
considered
relationships
member
a
of
the
S.
rubra, x-signata, and staufferi groups (Duellman &
1992; De la Riva,
Wiens,
out the
tem
same
range
explosive
is
S.
as
and
It
1993).
mating
Its
fuscovaria.
breeding
through-
occurs
sys-
in temporary
occurs
Bolivian
ported
obtained
were
W)
Santa Cruz
at
pond,
or
the
call consisted of
with low
(970 Hz)
rapid
one
this
recordings
mating
frequencies
long call
minute)
Riva
species
His
as
fundamental
la
a
note, of short duration,
calls per
De
locality.
the shore. The
emitted in
(337
vocalisations of
same
pulsed
was
(Fig. 2a).
time and
near
dominant and
and it
succession
tervals
ground
la
in the middle of
heavy rains from short grasses,
on
de
where males called after
at
shows
groups
in-
regular
recorded
is
at
the
based
same
on
the
call
a
showing
general
This is
a
We did
a
mm,
species
females 36.0
(Duellman,
1972).
of the S. rostrata
It inhabits open
the lower Amazon Basin from the Guiana
Mato Grosso
Recordings
were
obtained
river banks
were
emitted
or near
by
males
rainy
season
perched
neither
the
on
and
Bolivian
data
at
distance from
long
note
(240 ms) emitted sporadically,
ber of
and
pulses
high
dominant
with
a
low
one
num-
frequency (2870
on
the
Duellman
(1972)
egleri (a junior
Gruber,
described
synonym
1983).
4-7 notes,
with
He
repeated
the
according
described
at a rate
a
calls
to
call
of
Hyla
Hoogmoed
composed
195, fig. 1)
similar
very
with
5000
to
Hz,
to
our
Amazon, Brazil). He
of
types of
two
notes:
note.
secondary
notes
from
temporal
nor
(1977)
Pyles
calls of S.
call of S.
Bolivia.
the
Furthermore,
values
spectral
agree with
re-
recordings
our
of
S.
an
rate
values for
The
nebulosa do
only
the
by
one
rate
note
pulse
to
from 3840
are
not
close
per
de-
call,
of 62.3 notes/
an
average
and dominant fre-
pulses/second,
rate
senior
that
to
average durationof 130 ms,
of 60
data
agree with the
According
repetition
note
descriptive
not
description
1972).
nebulosa has
average
(then Ololygon egleri)
(Brazil).
published
numerical
provided
(1983)
nebulosa
(Duellman,
an
from
egleri
detect the existence of
a
the other values do
4147 Hz.
to
While the
those found
by
show similarities with
us,
our
results.
In summary,
the
descriptions
of the calls of S.
nebulosa appear to be rather conflictive.
the
of
uniformity
this
species
of the
ences
in call
morphological
throughout
unpublished data)
of
Hz) (Fig. 2b).
re-
frogs.
scription
plants
each other. The advertisement call consisted of
1972:
the calls of H.
(central
from Belém de Pará
minute,
Calls
values
secondary
Duellman &
pulse
probably
riverine
in
much shorter
Hödl
ported by
in
Almacén
semi-permanent ponds.
spaced
and
quencies ranging
Puerto
at
the
the
from 2500 Hz
composed
recordings
to
prolonged breeding system,
it. Males called
beyond
on
a
the whole
lasting during
areas
region
and Bolivia.
(Brazil)
where the species had
SVL:
(maximum
mm)
call
note
not
our
author
1824)
small
relatively
males 29.5
group
(Spix,
the
Hz
duration and
in
structure
of Manaus
area
primary
in
particularly
However, the audio-
ms
(1977) described
described
a
the
2b.
Fig.
the
a
previously
Scinax nebulosa
200
over
(mean 55.6),
far from
(1972).
frequencies
for
ours.
Duellman
us
(2374-2867
are
pulse
pulse rate reported
and
ms),
frequencies
emphasised
described the
(1993)
description
in
240
mean
by
a
harmonics
emphasised
Whereas the
recordings),
by
Hödl
Recordings
and
spectrogram provided (Duellman,
ponds.
Sierra (14°48'S 63° 10'
(our
Bolivian Scinax
of
duration of 160 ms,
the values found
emphasised
SVL:
Calls
pulses/second
durations
Scinax nasica
a mean
720 and 1050 Hz.
is similar
tions.
-
al.
et
leads
its
us to
range
(De
la
Riva,
believe that the differ-
descriptions reported
differences in
However,
characteristics
methods of
may
analysis
actual taxonomie differences between
be the result
rather than
populations.
&
of
of 39.5 notes/minute.
Scinax
This
parke ri (Gaige,
extremely
small
1929)
species (maximum
SVL: 23.3
Bijdragen
tot
de Dierkunde, 64
rubra Scinax rubra Scinax
2nd
1st
note
note*
(2)
-
81
1994
fus
parkeri Scinax nebulosa Scinax nasica Scinax garbei Scinax Scinax
covaria
Scinax
Scinax
chiqutan castroviejo castroviejo
are
Scinax
1st
2nd
note
note*
29.5
(24-34.3) (3.7)
1865.76
11
162.9
7
185.7
61
5
240.6
57.5
25
19
947
39
12
243
162.2
38
26.1
(142.7-19.5) (16.2) (167.-218.4) (17.6) (201.-305) (43.6) (42.5-98.7) (10.3) (295-1469) (137.-187.3) (185.3- 8.) (48.6) (21.7-32.5) (24.6-32.)
(275)
2096
2307.7
(34.1)
(88.6)
2374.6
(2.6)
(13)
1865.76
2096
27 7.9
2867.3
2172.4
891.1
905.6
(56.2)
(93.9)
(24.5)
(59.9)
(17.4)
(53.1)
3715.4
2172.4
864.1
2848.7
(176.9-205.6) (1 1.1) (2019.-240.) (2705.8-2 6.9) (2685.-3129.8) (167.7) (84.1- 0.6) (3452.9-357.) (136.9) (827.9-28.) (210-261.5) (276.3-298.5) (2301.9-2584.6)
(34.1)
0-2
10
(9-11)
(0.7)
1
11.4
(56.2)
(50.6)
1
18.6
1
1
13.4
(10-14) (1.3) (17-21) (1.4) (1 -17) (2.6)
5.1
(4-6)
(1.6)
864.1
(0.9)
(24.5)
1
1
53.1
(16-83) (15.6)
(59.9)
(7-9)
8.12
(0.6)
(47.1)
1
23-42
30
(6)
1
2406
(43.1)
1
Number
Note
(ms)
1695.1
972.9
calls
27.8
972.9
(176.9-205.6) (1 1.1) 2019.-240.) (201-42.9) (2 1.-2564.) (124.9) 84.1- 0.6) (154.-176.9) (827.9-28.) (210-261.5) (84.1-928.) (86.3-107.2)
frequency
(Hz)
duration
6
(0.4)
I.
next.
Sum ary
(6-6) (0)
numerical
than
one
type
is
parmetrs
of
the
Fundamental vocaliston.
the
note For
frequency Domina t
fol wed
each
of
the
by
an
asterisk par met rs,
(Hz)
/call
more
of
consider d,
Notes
6.1
(5-7)
shown
When
of
5
Table
/note Pulses
is
the
used
mean
for
the
values
are
342.1
69.7
100.2
55.6
50.12
122.1
(1.8)
(3.7)
234.3
216.1
(291.5-375) (33.7) (67.4-7 .4) (1.983) (96.2-104.9) (54.2-57. ) (74.-13.4) (53.8-57.8) (46. -56. ) (14.2-18.4) (157.-26.5) (25.1) (185.-243.9) (11.8)
(2.9)
91.1
(1.3)
(15)
56
(1)
/second Pulses
construcion fol wed
of
the
185.4
136.7
151.1
22.5
(164.7-206) (29.2) (103.6-156) (28.8) (83-206.7) (54.6) (2.4- 7.5) (22.8)
33.6
33.6
151.1
22.5
(23.8-457.8)
337.1
(56.3)
337.1
29.3
100.2
64.4
(1 .2-49. ) (62.9-12 .9) (15.7) (48.2-84.2) (12.8)
(8.9)
29.3
100.2
64.4
0.678
0.596
1504.2
(1348.-165.4) (1348.-165.4)
1504.2
(79.6)
(79.6)
80.6
80.6
(16. -46.8) (15.5) (16. -46.8) (15.5) (83-206.7) (54.6) (2.4- 7.5) (22.8) 2 3.8-457.8) (56.3) (1 .2-49. ) (62.9-12 .9) (15.7) (48.2-84.2) (12.8) (6.9-1 5.6) (26.6) (6.9-1 5.6) (26.6)
(8.9)
0.708
0.612
0.641
0.828
0.663
0.842
(0.64-0.843) (0.065) (0.45-0.81) (0.137) (0.567-0. 7) (0.045) (0.724-0.912) (0.063) (0.543-0.84) (0.084) (0.5-0.76) (0.074) (0.524-0.85) (0.075) (0.76-0.913) (0.047)
/minute
Notes
interval /inter-pulse
the
phenetic standard
trees.
/minute
by
deviatons
Calls
betwe n
Pulse
parenthse.
duration
Ranges
82
mm)
is
presumably
a
group
formerly
considered
nym of S.
followed
member of
authors
several
considered S. parker i
least in
(1973)
valid
a
northern Bolivia and
breeder
Recordings
differ
De la
rubra.
Wiens
(1992)
a
It
occurs
Brazil.
and is
at
obtained
were
different. Both the
merical
syno-
at
It is
probably
rainy
a
season.
locality,
the type
perched
on
call consisted of
one
activity
edge,
sporadic.
have been
To
species
females 43.6
from
Neotropics
southern
northern South America
taxonomy of both
of
complex
under this
rubra
is
species
of
SVL:
includes both
1985).
forests
and
obtained
at
to
likely
were
where S. rubra is
an
temporal
Males
perching
ponds.
explosive species
emitted
Males formed
and consisted of
longer
their
breeding
calls
in
while
both leaves and branches of bushes and
on
trees.
large
persisted until dawn. The call
The
Puerto Almacén,
note
choruses that often
emitted
was
pulsed
two
(primary note)
notes
isolation and is longer in duration and higher in frequency than the second
sus
29
second
pulses
ms
and
note
(1969)
Hyla rubra)
ms
be
(342 pulses
Rivero
200
can
1865
Hz
(163
in
repeated
per
and 2100 Hz,
ver-
second
The
with
a
high
note).
number of
second).
from Estado
He
Bolívar,
(then
Venezuela
as
a
call with dominant frequency of
compared
this call with that of
of 63
described the
Hyla)
(pulse frequency
mating
Panama
66-72
tion
of
pulse
rate
somewhat
are
teristics of the
of
a
longer
142-197 ms,
range
his
Whereas
ms.
pulse
description
described
rate
not
not
2 of
(fig.
frogs
all in the
Campbell,
depicted
as
those found in
similar
those
(Ecuador)
Panamanian
Campbell
the
1600
at
of
At any rate,
the
calls
described
shorter
Peru
note
recordings
though
this
figure
do resemble
indi-
for
recordings
this
in
species
a
of
description
(then Hy la)
the
from Santa
which coincided with that of the
(1971).
types of calls,
well,
that
audiospectrogram
1971).
provided
(1978)
mating
our
our
(1970b).
Cecilia
a
fact
located
was
reported
Duellman
by
in
call of Scinax rubra
ent
pulses/
with
agree
analyses. Campbell (1971)
our
to
Duellman
(1979)
by
call characteristics of his
cated that the
were
at
(duration
us
characteristics of the call of the Panama-
spectral
nian
reflected
dura-
report the existence
not
dominant frequency of the call
Hz is
a
the charac-
to
However, the
note.
one
duration and
range 67-74
does
because it does
secondary
the
comparable
note
call of
as
pulses/second)
note
140-150
1581
at
pulses/second.
dominant frequency of 1600 Hz and
Hyla) from
described the call of S. rubra
long pulsed
about 1250 Hz.
ms
the
centered
frequency
from
(Ecu-
(mean duration
note
a
(Fig. 2d).
is often emitted in
call of S.
mating
with
Ecuador
sporadi-
(then
since the
complex.
very
rate
a
not
x-signata
S.
versa)
from Santa Cecilia
pulse
later
It would
pulsed
recordings
disturbed
vice
x-
1 in
but it shows
frequency.
short
single
average
rubra
the call
Recordings
cally
Scinax
that
areas.
small
an
second),
habitat of S.
open,
Hz and
cen-
included
being
The
the
throughout
of the Andes
a
Campbell (1971)
males
in
being
Venezuela (fig.
described the
130 ms) with dominant
(ap-
knowl-
our
widespread
currently
(Frost,
name
Duellman (1970b)
Venezuela
in
of Bolivian S.
Fig. 2d,
is
be
nu-
of the call of H.
(and
species
(then Hyla rubra)
and southeastern Brazil. It is
Bolivia,
S. rubra
was
as
Panama
east
actually
to
and the
considered
Rivero
ador)
(maximum
mm)
that
unlikely
with
of the calls of Scinax
is
our
(186 ms)
1768)
(Laurenti,
resembles
lower fundamental
rubra
frequencies
the best
Miranda,
The
published.
This medium-sized
mm,
note
rubra
recordings
our
area.
Hz) (Fig. 2c). Calling
2800
previous descriptions
Scinax rubra
tral
large
pulsed
flooded
fundamental
2300 and
was
no
parkeri
36.1
short
dominant and
high
proximately
a
from
S.
from Estado
slightly
what
for
structure
1969)
Bolivian Scinax
of
and found them
1824)
However, the
Rivero,
be
Calls
audiospectrogram
reported
markedly
signata
Buenavista (17°27'S 63°40' W), where males called
low grasses in
data
-
al.
et
Hyla x-signata (Spix,
position
adjacent
the whole
during
S.
was
1985;
species.
characteristic of open habitats,
prolonged
junior
(Frost,
Duellman &
as
It
1992).
a
1925)
fuscomarginata (Lutz,
by
the dubious
Wiens,
Lutz
by
1990a) although
Riva,
&
(Duellman
staufferi
a
De la Riva
/.
of
this
On
the
the
as
a
(50
call
described
species
other
of
Scinax
rubra
being composed of
two
long (200 ms) pulsed
ms
approximately).
from Bolivia show
(then
differ-
note
and
Similarly,
two note
with shorter durations
by
Schlüter
hand,
(long
types
as
142-167
Bijdragen
Fig.
3.
tot
de Dierkunde, 64
Phenetic tree
Bolivia.
scale represents
recordings
notes
83
of
analyses
12
of the calls of eight species of Scinax from
short 24—34 ms).
ms,
our
The
1994
-
from the UPGMA
resulting
numerical parameters
(2)
to
be
shown
by
frequencies
of
lower for both
slightly
Schlüter
(1979),
how-
whose
&
pulse
of the
rate
longer
note
is similar
frogs (estimated
to
that found
by
Figs.
shown for the
pulses/second)
discrepancy
67-74
(range
us
in
mating
calls
is
phenetic
troviejoi
ador
the
on
one
the
other
hand,
suggests
that
3
is
the
belong
to
two
may have
developed
tree, but does
is
of S. rubra in these locations
status
with
show
not
of the
phenogram
the
UPGMA
concordance with
genus.
Again,
but
to
markedly separated,
not
S.
the
cas-
same
in
as
Fig.
3. This
is
species
with S.
grouped
and S. nebulosa and S.
parkeri (the pair
These four
species probably
dif-
closely clustered).
dif-
distinctly
ferent dialects. A careful re-evaluation of the
nomie
correlation
consistent
individuals
most
or
(Duellman
and 4.
rather
chiquitana,
recorded in these locations may
ferent types
of
hand, and from Peru and Bolivia
degree
on
eight species
correlations.
The advertisement call of S. par-
of the calls from Panama and Ecu-
descriptions
audiospectrographic
calls of
demonstrated
not
the tentative phylogeny
the
the
The scale represents
audiospectrographic
4)
pulses/
between
1992).
from
notes of the
keri
is similar to that of S. nebulosa, according to
(Fig.
The
second).
be 60
to
primary
monophyly
Wiens,
The
Peruvian
resulting
tree
of the
from Bolivia.
both
the
ever,
Phenetic
correlations
Scinax
distances.
The dominant
appear
than those
standard
4.
Fig.
four different
belong
to
in both
Fig.
species
Although
groups.
taxo-
may
3 and
Fig. 4,
S. rubra and S.
garbei,
and
be
S.
and S.
fuscovaria
nasica,
clustered
are
together,
fruitful.
the relative
The
lar
Quantitative comparisons of theadvertisement
calls
if
The UPGMA
netic
cates
a
hand,
solid
the
rently
level
tree
has
are
between the
not
clear
are
the
to
(both belong
are not
to
grouped
is
clearly
the
rostrata
in the
tree.
group)
within
1992)
few
the
by
closely
but
is
rest
size, although
estab-
method
related
to
of
to
the
However,
tween
The
as
currently
group,
does
rate
neither
phenograms
mainly
forest
ence
include
in
notes
there is
in
species
not
and S.
a
(five
nor
out
UPGMA).
correlation bethe
calls.
ecological
species
garbei
variables
call
reflect
clearly
striking
correla-
considered
chiquitana
and S.
rubra
are
found both in the open and in
formations. These last
associated in the
in habitat of S.
either of the
resulting
any
the
no
a
utterances
characteristics of
either. Most of the
forest,
simi-
more
open habitat dwellers. Scinax
euryoecious
tantly
even
included
see,
and
do
same.
in
trees
of emission of the
we can
phylogeny
relationships
is
the
species
audiospectrographic
parameters
lives in the
S.
affinity
not
are
between the
the number of different
twelve
pairs
cluster the
methodologies
consider that the
are
hand,
staufferi
tion
two
pheno-
of the speit
we
The
related
their calls
On the other
is included with S. nasica in the S.
cur-
among the spe-
are
different from the
because of its small
the other
Wiens,
well reflected
garbei
indi-
and the
taxonomists. The
not
co-phe-
relationships
relationships
gram. S. nebulosa and S.
parkeri
tree
a
which
On
consistency.
Scinax (see Duellman &
lished affinities
cies,
(Fig. 3)
phylogenetic
high, although
cies
of
consistency
accepted
the genus
not
phenetic
correlation coefficient of 0.897,
way.
from the
of the
positions
phenograms
two
species
two
phenograms,
chiquitana is
phenograms.
are
dis-
and the differ-
not
reflected in
De la Riva
/.
84
although according
In summary,
to some
authors
closely related species may show similarities in their
calls
(Schiotz,
of the genus Scinax
study
the
1973),
rubra
in
group
Panama
conform
not
this
to
row
results of Duellman (1970b) who did
not
coherence between the classification
of the
of
Hyla
and the similarities among
find great
species
calls
mating
(but
see
Márquez
the
1990 Venezuelan
lack of
al., 1993). This
et
result of the fact that the
tablished. On
tween
the
holistic
the other
approach
being clearly
the
by
G.E.
graphic
correlation
of
become
eventually
is
powerful
a
sounds.
This
es-
a more
and
of numerical variables for multivariate
agile
frog
Garbeana
alter-
rostrata
Kempff Mercado",
collaboration.
of Santa Cruz
Fieldwork
grant
from the
Riva.
Edwin
Chacón
We are
who
the
trogram
de la
de
trees
CYCIT PB 89-0045C
Ciencia, Spain.
methodological
(PI:
P.
advice
and R.
for the
the
to
in
a
I. De la
his
calls
the
for the
were
Alberch)
C.W. Clark
calls
from
help
analyses
Sound
Reig.
Doñana
of
analyses
generated the phenetic
S.
for its
ranch,
grateful to A. Machordom
particularly
phenetic
"Noel
supported by
was
record
to
us
Natural
Sierra, Bolivia,
de
Amigos
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provided by
y
Historia
South America
allowed
Puerto Almacén.
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in
Asociación
performed
de
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through
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funded
Ministerio
Rican
of the
1970:
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