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Altitudinal distribution and advertisement call of Colostethus latinasus (Amphibia: Dendrobatidae), endemic species from eastern Panama and type species of Colostethus , with a molecular assessment of similar sympatric species


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We conducted a molecular assessment of Colostethus-like frogs along an elevational gradient in the Serranía de Pirre, above Santa Cruz de Cana, eastern Panama, aiming to establish their species identity and to determine the altitudinal distribution of C. latinasus. Our findings confirm the view of C. latinasus as an endemic species restricted to the highlands of this mountain range, i.e., 1350–1475 m.a.s.l., considered to be type locality of this species. We described the advertisement call of C. latinasus that consists of a series of 4–18 single, short and relatively loud “peep”-like notes given in rapid succession, and its spectral and temporal features were compared with calls of congeneric species. For the first time, DNA sequences from C. latinasus were obtained, since previously reported sequences were based on misidentified specimens. This is particularly important because C. latinasus is the type species of Colostethus, a genus considered paraphyletic according to recent phylogenetic analyses based on molecular data.
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Accepted by E. Twomey: 2 Feb. 2017; published: 12 Apr. 2017
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ISSN 1175-5326 (print edition)
(online edition)
Copyright © 2017 Magnolia Press
Zootaxa 4254 (1): 091
Altitudinal distribution and advertisement call of Colostethus latinasus
(Amphibia: Dendrobatidae), endemic species from eastern Panama and type
species of Colostethus , with a molecular assessment of similar sympatric species
Smithsonian Tropical Research Institute, Apdo. 0843-03092, Balboa, Ancón, Panamá, Republic of Panama. E-mail:
Círculo Herpetológico de Panamá, Apdo. 0824-00122, Panamá, Republic of Panama.
Departamento de Zoología, Universidad de Panamá, Panamá, Republic of Panama.
G&S Nature and Environmental Consultants S.C., Edgardo Griffith, P.O. BOX 527948, Miami, FL 33152-7948, USA. E-mail:
Department of Biology, University of Maryland, College Park, MD 20742, USA. E-mail:
Departamento de Ciencias Biológicas, Universidad de los Andes, AA 4976, Bogotá, Colombia. E-mail:
Corresponding author.
We conducted a molecular assessment of Colostethus-like frogs along an elevational gradient in the Serranía de Pirre,
above Santa Cruz de Cana, eastern Panama, aiming to establish their species identity and to determine the altitudinal dis-
tribution of C. latinasus. Our findings confirm the view of C. latinasus as an endemic species restricted to the highlands
of this mountain range, i.e., 1350–1475 m.a.s.l., considered to be type locality of this species. We described the advertise-
ment call of C. latinasus that consists of a series of 4–18 single, short and relatively loud “peep”-like notes given in rapid
succession, and its spectral and temporal features were compared with calls of congeneric species. For the first time, DNA
sequences from C. latinasus were obtained, since previously reported sequences were based on misidentified specimens.
This is particularly important because C. latinasus is the type species of Colostethus, a genus considered paraphyletic ac-
cording to recent phylogenetic analyses based on molecular data.
Key words: Neotropical, Anura, systematics, taxonomy, DNA barcoding, Silverstoneia
Realizamos una evaluación molecular de ranas parecidas a Colostethus, a lo largo de un gradiente de elevación en la Ser-
ranía de Pirre, arriba de Santa Cruz de Cana, en el este de Panamá, con el objetivo de establecer la identidad de las especies
y determinar la distribución altitudinal de C. latinasus. Nuestros hallazgos confirman la opinión de C. latinasus como una
especie endémica restringida a las tierras altas de esta serranía, i.e., 1350–1475 m.a.s.l., considerada la localidad tipo de
esta especie. Describimos el llamado de anuncio de C. latinasus que consta de una serie de 4–18 notas sencillas, cortas y
relativamente ruidosas, semejantes a un "piip", dadas en una sucesión rápida; además, comparamos sus características es-
pectrales y temporales con los llamados de especies congéneres. Por primera vez, se obtuvieron secuencias de ADN de C.
latinasus, ya que las secuencias previamente reportadas provienen de especímenes identificados erróneamente. Esto es
particularmente importante debido a que C. latinasus es la especie tipo de Colostethus, un género considerado parafilético
de acuerdo con análisis filogenéticos recientes, basados en datos moleculares.
Colostethus latinasus was originally described by Cope (1863) as a Phyllobates from the Truandó region of
northern Chocó, Colombia. Since its description, the identity of this species has been elusive, being frequently
confused with other Central and South American dendrobatoid species (Savage 1968; Grant 2004), a confusion
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aggravated by the loss of the type specimen (Barbour & Noble 1920; Savage 1968). Aiming to stabilize the
nomenclature of this species, Savage (1968) designated as the neotype a female specimen (National Museum of
Natural History specimen USNM 50198) collected from Cana (= Santa Cruz de Cana), Darién Province, Republic
of Panama. Despite doubts about the precise collecting locality of the neotype, Grant (2004) examined museum
specimens from the Serranía de Pirre that agreed in all aspects with the neotype and concluded that it was probably
collected from the highlands above Cana.
In the past two decades, the systematics of the anuran family Dendrobatidae has been comprehensively revised
(e.g., Grant et al. 2006). Even though several new species of this family have been described (e.g., Grant & Myers
2013), many distinctive frogs remain undescribed (e.g., Grant et al. 2006). Therefore, it is important for future
systematic and taxonomic studies to clearly establish the identity of C. latinasus, the type species of the genus
Colostethus Cope, 1866 (Savage 1968, Grant 2004, Grant et al. 2006, Frost 2016). Moreover, Colostethus is
currently considered to be paraphyletic with respect to the genus Ameerega (Santos et al. 2009, Pyron & Wiens
2011), a condition still waiting to be resolved (Frost 2016).
The geographic distribution of C. latinasus is restricted to the mountain range of Serranía de Pirre. Other
specimens previously assigned to this species (e.g., Savage 1968) from nearby Cerro Malí, a peak in the Altos de
Puna region of the Serranía del Darién (Myers & Lynch 1997), are considered to represent a similar yet
undescribed species (Grant 2004). Given the well-established role of advertisement calls in species recognition and
reproductive isolation (Wells 2007), and the relevance of call characters in anuran phylogenetic and systematic
studies (Padial et al. 2008, Goicoechea et al. 2010), we describe here the advertisement calls emitted by males of
C. latinasus from the Serranía de Pirre, above Cana, currently considered to be the type locality of this species
(Grant 2004). In addition, we conducted a molecular assessment of individuals of similar sympatric species along
an elevational gradient in this region, aiming to establish their identity and to determine the altitudinal distribution
of C. latinasus.
Materials and methods
Survey area and specimen collection. To complement our observations on the altitudinal distribution of C.
latinasus and establish the identity of similar sympatric species present in this geographic area, we collected tissue
samples from individuals of Colostethus-like frogs and tadpoles, along a 449–1606 m.a.s.l. elevational gradient,
from the surroundings (i.e., < 5 km) of Cana field station (7.755800ºN 77.684583ºW) to the ridge of the adjoining
Serranía de Pirre (7.774500ºN 77.733200ºW). We followed the American Society of Ichthyologists and
Herpetologists guidelines for use of live amphibians in field and laboratory research, and our protocol was
approved by the Smithsonian Tropical Research Institute’s IACUC. Field-caught specimens were euthanized with
Benzocaine. Their tissues were stored in a NaCl-saturated buffer containing 0.25 M EDTA and 20% dimethyl
sulfoxide (DMSO) (Seutin et al. 1991). All specimens were vouchered in museum collections. We used the
following collection abbreviations: AJC = Andrew J. Crawford field series, CH = Círculo Herpetológico de
Panamá, and MVUP = Museo de Vertebrados de la Universidad de Panamá. Snout-vent length (SVL) of voucher
specimens were measured using a Mitutoyo Absolute digital caliper (model CD-6”CX) to the nearest 0.01 mm.
DNA barcoding and analysis. DNA extraction, PCR and sequencing protocols followed those of Crawford et
al. (2010) for obtaining mitochondrial DNA sequence data from the COI ‘Barcode of Life’ fragment (Hebert et al.
2003) and the 16S ribosomal gene (Kessing et al. 2004). We obtained 657 aligned base pairs (bp) for the COI gene
fragment, and 556 bp from the 16S gene after excluding sites with gaps.
Candidate evolutionary models were evaluated using jModeltest version 0.1.1 applied to each gene separately
based on the corrected Akaike Information Criterion (AICc) to identify the model that simultaneously minimized
bias and variance in parameter estimates (Anderson 2008, Posada 2008). As a model of intermediate complexity,
the 2-rate HKY+G model, was recommended for the COI gene, and therefore was not partitioned further. The 6-
rate GTR+G model was recommended for the 16S data subset. A 2-gene partitioned likelihood phylogenetic
analysis (Felsenstein 1981) was conducted with GARLI version 2.0 using 20 replicate searches and default settings
(Zwickl 2006). Parameters and rates were independent between the two data partitions. A bootstrap likelihood
analysis was conducted in GARLI, with two replicate searches conducted per each of 300 bootstrapped datasets. To
accelerate search times, parameter values for the best-fit unpartitioned evolutionary model (according to
jModeltest) were set to their ML values.
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Statistical support for clades was also evaluated by Bayesian Markov chain Monte Carlo (MCMC)
phylogenetic inference (Yang & Rannala 1997) with MrBayes version 3.1.2 (Ronquist & Huelsenbeck 2003) on the
CIPRES portal (Miller et al. 2010), assuming default parameters except the prior distribution on the rate
heterogeneity shape parameter, alpha, was given a maximum value of 5.0, and three Metropolis-coupled chains
were run with the temperature parameter lowered to 0.05 to increase swap rate among chains, based on trial runs.
Two independent MCMC analyses were run for 5 million generations each with posterior probabilities estimated
from samples taken every 1,000 generations following a burn-in of 3 million generations, at which point average
standard deviation of split frequencies between runs had dropped below 0.015 and the chains had converged to
similar likelihood scores.
Acoustic recordings and analysis. We used a SONY IC Recorder ICD-P320 with its built-in microphone to
record the calls of the male preserved as voucher specimen CH 6400 (MVUP 2342, see below). The compressed
format DVF files from this recorder were converted to 44.1 kHz and 16-bit WAV files, using the Digital Voice
Editor 2 software. The calls of additional, unvouchered males were recorded using a Roland’s Edirol 24-bit wave/
mp3 recorder R-09 with a Realistic directional microphone (Cat. No. 33-1062). These recordings were made at a
sampling frequency of 44.1 kHz and a wav-16-bit mode. The air temperature was registered during recordings.
All recordings were analyzed with the sound analysis and synthesis laboratory software Avisoft-SASLab Pro
version 5.1.23. We used the waveform display with a resolution of 0.2–0.3 milliseconds (ms) to measure the
temporal variables of the call, and used the spectrogram display with a Hamming window, 1024 points FFT size, 56
Hz bandwidth and 43 Hz resolution for measuring frequency variables, and used the power spectrum to determine
frequency peaks. For the spectral analysis of notes and creating graphics, only the best quality, uncompressed audio
recordings were used. With the spectrogram display we determined the maximum and minimum frequencies of
each note to calculate its bandwidth. Data extracted from calls of different males were pooled for descriptive
statistical analyses. The potential effects of temperature on the temporal features of these calls could not be
adequately assessed due to the small sample size. However, differences in air temperature of these call recordings
were relatively small, i.e., maximum ΔT = 4°C. Moreover, any intraspecific temporal variation in calls that could
be attributed to temperature appeared to be minimal, especially for interspecific comparisons.
Altitudinal distribution. We collected 46 individuals, comprising 5 to 24 specimens of four species (Table 1). Out
of these 46 individuals, only 5 of these could be grouped as C. latinasus and were found at a 1350–1475 m.a.s.l.
elevation range (Fig. 1). The other individuals could be grouped as C. aff. panamansis (7 individuals with a 494–
1246 m.a.s.l. range), C. aff. pratti (24 individuals with a 449–1246 m.a.s.l. range), and Silverstoneia aff. nubicola
(10 individuals with a 462–1597 m.a.s.l. range). These three species had a more extensive distribution along the
elevational gradient compared to C. latinasus. Colostethus aff. panamansis and C. aff. pratti showed a similar
altitudinal distribution pattern, while Silverstoneia aff. nubicola had the most extensive vertical distribution along
the elevational gradient, overlapping the high-altitude distribution of C. latinasus. At high-altitude, individuals of
C. latinasus can be distinguished from these three species by their relatively larger size (SVL of adult males =
29.40–32.39 mm, n = 3; adult female SVL = 29.83–30.16 mm, n = 2) and more terrestrial habits. In addition, adult
individuals of C. latinasus have a marbled throat and chest, being more darkly marbled in males (Figs. 2, 3).
Vocalization. Males of C. latinasus were often observed in shaded and very humid areas with abundant leaf
litter and fallen branches and logs, away from streams, calling on the forest floor or on fallen vegetation during
1130–1245 h, above Cana, near the ridge of Serranía de Pirre (7.76551ºN 77.72418ºW to 7.76893ºN 77.72758ºW,
datum WGS84, 1350–1475 m.a.s.l.), Darién Province, Republic of Panama, on August 8, 2007.
A male (29.40 mm in SVL, preserved as a voucher specimen CH 6400 in the Círculo Herpetológico de Panamá
collection (later deposited in the Museo de Vertebrados de la Universidad de Panamá as MVUP 2342, Fig. 3) and
other calling males were recorded at an air temperature of approximately 23°C. The digital audio file of
advertisement calls of male MVUP 2342 can be heard online at
metas/view/48934. Additional males were recorded at an air temperature of 19°C.
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TABLE 1. Samples of dendrobatid (Colostethus-like) frogs and tadpoles used in molecular phylogenetic analyses. All samples were collected in and around the Cana field station, Darién
National Park, Darién Province, Republic of Panama. AJC = Andrew J. Crawford field series, CH = Círculo Herpetológico de Panamá, and MVUP = Museo de Vertebrados de la Universidad de
Panamá. Asterisk (*) indicates a tadpole specimen.
Genus Species Field series number Collection specimen
GenBank numbers
Colostethus latinasus CH 6381 CH 6381 1475 m KF806998 KF807037
Colostethus latinasus CH 6389 CH 6389 1400 m KF806996 KF807035
Colostethus latinasus CH 6400 MVUP 2342 1350 m KF806999 KF807038
Colostethus latinasus CH 6402 CH 6402 1475 m KF806997 KF807036
Colostethus latinasus CH 6405 CH 6405 1475 m KF806995 KF807034
Colostethus aff. panamansis AJC 1816 CH 9628 550 m KC129188 KC129295
Colostethus aff. panamansis AJC 1856* CH 9654 1246 m KC129215 KC129323
Colostethus aff. panamansis AJC 1857* CH 9655 1246 m KC129214 KC129322
Colostethus aff. panamansis AJC 1859 CH 9634 1246 m KC129213 KC129321
Colostethus aff. panamansis CH 5546
a CH 5546 1246 m KC129211 KC129319
Colostethus aff. panamansis CH 6356 CH 6356 494 m KC129212 KC129320
Colostethus aff. panamansis CH 6448 CH 6448 1246 m KC129189 KC129296
Colostethus aff. pratti AJC 1697 CH 10087 525 m KF807020 KF807059
Colostethus aff. pratti AJC 1698 CH 10088 525 m KF807023 KF807062
Colostethus aff. pratti AJC 1699 CH 10089 525 m KF807021 KF807060
Colostethus aff. pratti AJC 1700 CH 10090 525 m KF807000 KF807039
Colostethus aff. pratti AJC 1801 CH 10091 525 m KF807002 KF807041
Colostethus aff. pratti AJC 1802 CH 10092 525 m KF807004 KF807043
Colostethus aff. pratti AJC 1803 CH 10093 525 m KF807005 KF807044
Colostethus aff. pratti AJC 1804 CH 10094 525 m KF807006 KF807045
Colostethus aff. pratti AJC 1805 CH 10095 525 m KF807007 KF807046
Colostethus aff. pratti AJC 1806 CH 10096 525 m KF807008 KF807047
Colostethus aff. pratti AJC 1807 CH 10097 525 m KF807009 KF807048
Colostethus aff. pratti AJC 1808 CH 10098 525 m KF807010 KF807049
Colostethus aff. pratti AJC 1810* CH 10106 526 m KF807011 KF807050
Colostethus aff. pratti AJC 1811* CH 10107 525 m KF807012 KF807051
……continued on the next page
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TABLE 1. (Continued)
Genus Species Field series number Collection specimen
GenBank numbers
Colostethus aff. pratti AJC 1812 CH 10099 550 m KF807013 KF807052
Colostethus aff. pratti AJC 1815 CH 10100 550 m KF807022 KF807061
Colostethus aff. pratti AJC 1877 CH 10101 1246 m KF807015 KF807054
Colostethus aff. pratti CH 5524 b CH 5524 550 m KF807003 KF807042
Colostethus aff. pratti CH 5598 c CH 5598 535 m KF807017 KF807056
Colostethus aff. pratti CH 5601 c CH 5601 525 m KF807019 KF807058
Colostethus aff. pratti CH 5602 d CH 5602 550 m KF807018 KF807057
Colostethus aff. pratti CH 6330 CH 6330 525 m KF807016 KF807055
Colostethus aff. pratti CH 6363 CH 6363 449 m KF807001 KF807040
Colostethus aff. pratti CH 6446 CH 6446 1246 m KF807014 KF807053
Silverstoneia aff. nubicola AJC 1870 CH 10102 1246 m KF807030 KF807069
Silverstoneia aff. nubicola AJC 1880 CH 10103 1246 m KF807031 KF807070
Silverstoneia aff. nubicola AJC 1881 CH 10104 1246 m KF807026 KF807065
Silverstoneia aff. nubicola AJC 1889 CH 10105 1246 m KF807027 KF807066
Silverstoneia aff. nubicola CH 5558 CH 5558 1130 m KF807024 KF807063
Silverstoneia aff. nubicola CH 5599
e CH 5599 525 m KF807028 KF807067
Silverstoneia aff. nubicola CH 5600 CH 5600 525 m KF807029 KF807068
Silverstoneia aff. nubicola CH 6369 CH 6369 462 m KF807033 KF807072
Silverstoneia aff. nubicola CH 6370 CH 6370 462 m KF807032 KF807071
Silverstoneia aff. nubicola CH 6404 CH 6404 1597 m KF807025 KF807064
a COI was sequenced for the same individual by Grant et al. (2006), referred to as C. panamensis. The mitochondrial genome region 12S-16S was also sequenced for this same individual by
Santos et al. (2009) and referred to as C. panamensis or C. inguinalis, and referred to as C. panamansis in Pyron & Wiens (2011). This individual was collected in the headwaters of Río Cana,
Serranía de Pirre.
b COI was sequenced for the same individual by Grant et al. (2006), referred to as C. pratti-like”. This individual is from Cana, not from “Jungurudó”, Darién, as reported previously, i.e.,
sample 1144 in Grant et al. (2006). This specimen was referred to as C. pratti by Pyron & Wiens (2011).
c Referred to as C. pratti-like” by Grant et al. (2006). 12S-16S was sequenced for the same individual and referred to as C. latinasus by Santos et al. (2009); subsequently, included in Pyron &
Wiens (2011).
d Referred as C. pratti-like” by Grant et al. (2006).
e 12S-16S was sequenced for the same individual by Santos et al. (2009), referred as S. nubicola or C. nubicola. This individual was collected in Cana on the trail to Boca de Cupe.
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FIGURE 1. Likelihood-based phylogeny inferred from a partitioned analysis of partial DNA sequences of the COI and 16S
mitochondrial genes. Scale bar indicates patristic distances estimated from partitioned likelihood analysis. Clade support values
indicate Bayesian marginal posterior probabilities followed by likelihood bootstrap support values as percentage. Unscaled
photographs of Silverstoneia aff. nubicola (CH 5558, SVL = 20.48 mm), Colostethus aff. panamansis (CH 5546, SVL = 25.40
mm), C. latinasus (CH 6389, SVL = 30.16 mm) and C. aff. pratti (CH 1087, SVL = 18.48 mm).
The advertisement call of C. latinasus consisted of a series of single and short, relatively loud “peep”-like
notes given in rapid succession (Fig. 4). Males called at a rate of 4.0 ± 1.4 calls/min (mean ± standard deviation, n
= 5 males), apparently synchronizing their calls to those of neighboring males but often partially overlapping them.
The advertisement calls had 4–18 notes (10 ± 5 notes, mode = 7 notes, n = 15 calls of 8 males), 0.3–1.6 second (s)
duration (975 ± 468 ms) and a note repetition rate of 11.3 ± 0.9 notes/s (n = 11 calls of 4 males). The dominant
frequency of calls had one major peak at 2658 ± 79 Hz (n = 14 calls of 7 males). The notes had a duration of 41 ± 6
ms (n = 79 notes of 8 calls from 2 males), and inter-note intervals of 57 ± 9 ms (n = 71 inter-note intervals of 8 calls
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from 2 males). These notes were slightly upwardly frequency modulated. The lower frequency of the notes was
2427 ± 26 Hz, their higher frequency is 2878 ± 69 Hz, their bandwidth was 451 ± 66 Hz and the dominant
frequency of notes peaked at 2673 ± 40 Hz (n = 56 notes of 5 calls from 1 male). The first note of calls often had a
slightly lower frequency peak, less frequency modulation and lasted a few ms longer than the other notes.
FIGURE 2. Color in life of Colostethus latinasus individuals. Unscaled photographs. A–C) adult female (CH 6381), SVL =
29.83 mm; D–E) adult female (CH 6389), SVL = 30.16 mm; F) adult male (CH 6402), SVL = 31.72 mm.
As commented in Grant (2004), the presence of C. latinasus at Cana (500–550 m.a.s.l.) is unlikely. We have not
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seen individuals nor heard any calls of this species at this site, despite of approximately 18 days of surveys
collectively accumulated during years 2002–2010. However, we have routinely heard males calling from the slopes
high above Cana, along a trail to the top of the Serranía de Pirre ridge, i.e., approx. 1350–1500 m.a.s.l. Our
observations support the view that C. latinasus is a species restricted to the highlands. Therefore, specimens that
were previously assigned to C. latinasus from lower elevations around Cana (e.g., Santos et al. 2009, Pyron &
Wiens 2011), as well as their associated tissues and DNA sequences (see Table 1), do not belong to this species. In
the present study, DNA sequences of C. latinasus are published for the first time.
FIGURE 3. Voucher specimen of Colostethus latinasus (CH 6400, MVUP 2342), adult male. A) ventral view; B) dorsal view.
As with other dendrobatoid species (Ibáñez & Smith 1995), the description of the advertisement call of C.
latinasus may be helpful for species identification in studies of the diversity and systematics of anurans from
Darién and adjacent Colombia. We analyzed two advertisement calls recorded by William E. Duellman from a
male specimen (University of Kansas Natural History Museum KU 76830) on Cerro Malí, at a temperature of
20°C, and deposited as recording No. 7227 in the Fonoteca Zoológica, Museo Nacional de Ciencias Naturales de
Madrid ( We assumed no errors in the available recording, and this specimen was previously
considered to represent C. latinasus (Savage 1968). Nonetheless, these calls differ from those of C. latinasus,
described here, mainly in having a longer duration (mean 8.4 vs. a maximum of 1.6 s in C. latinasus), slower note
repetition rate (mean 6.6 vs. 11.3 notes/s in C. latinasus) and notes lacking evident frequency modulation (present
in C. latinasus). Therefore, morphological data (Grant 2004) and call characteristics reported here support
specimens from Cerro Malí as being another species.
The advertisement call of C. latinasus is also distinct from other species of the genus Colostethus. Detailed
descriptions of the vocalizations, based on spectral and temporal characteristics, from C. panamansis (as C.
inguinalis) have been published (Wells 1980). No voucher specimens are associated with any of these calls, and the
reported air temperature was 25°C (Wells 1980). The advertisement call from a population of C. panamansis at
Cerro Campana, in central Panama, was described as a whistled trill composed of 2–5 frequency modulated notes,
each note with an approximate duration of 60 ms (Wells 1980). It differs from C. latinasus by having fewer notes
(mode of 3 vs. mode of 7 notes in C. latinasus) with a longer duration (60 vs. 41 ± 6 ms in C. latinasus) and a
higher frequency (3200–4547 vs. 2427 ± 26 to 2878 ± 69 Hz in C. latinasus) (Wells 1980). In addition, males of C.
panamansis are known to emit advertisement calls at a higher rate (46–58 vs. 4.0 ± 1.4 calls/min in C. latinasus)
(Wells 1980).
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FIGURE 4. Waveform (upper), power spectrum (lower left) and spectrogram (lower right) of the advertisement call produced
by a male Colostethus latinasus, recorded at 19°C air temperature. This is one call in the series of five calls that can be heard
online at
We thank W.E. Duellman for permission to use his recording of a male from Cerro Malí and R. Márquez and L.
González for facilitating access to this recording, D. Medina, S.V. Flechas, R. Brenes, and the guides from Ancon
Expeditions for their assistance in the field. We thank C.A. Jaramillo for photographic images of some specimens,
M. Dimas for curatorial assistance, N.A. Ibáñez and J. Alemán for their help with figures, and A. Driskell at the
Smithsonian’s Laboratory of Analytical Biology for obtaining the DNA sequence data. Throughout the years, we
were accompanied by many other people on our field expeditions to Cana, most notably M. Evans, M. Hughey,
D.G. Mulcahy, K.E. Nicholson, and K. Zamudio. Our fieldwork was primarily funded by the grant 8133-06 from
the Committee for Research and Exploration of the National Geographic Society to AJC, RID, and KRL, and was
conducted under ANAM permits No. SE/A-07902 and SE/A-37-07 to RID. RID was supported by the Panama
Amphibian Rescue and Conservation Project, and the Sistema Nacional de Investigación of Panama. Suggestions
from E. Twomey and T. Grant helped to improve greatly the final version of the manuscript.
Anderson, D.R. (2008) Model Based Inference in the Life Sciences: A Primer on Evidence. Springer, New York, 184 pp.
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... Santos et al. (2009) did not challenge or even mention Grant et al.'s (2006) conclusion and did not analyze or discuss the sequences generated by Grant et al. (2006), despite their similarity (e.g., as assessed by BLAST) to the sequences they reported as C. latinasus. Recently, DNA sequences of C. latinasus were published by Ibáñez et al. (2017), which allowed us to test the relationships of that species (see Other ingroup relationships: Genera, above). Santos et al. (2009) took special exception to the recognition of the inclusive clade of "dendrobatid frogs" (sensu Grant et al., 2006) as Dendrobatoidea, comprising two families, each with three subfamilies. ...
... Terminals labelled "Colostethus 'latinasus'" are conspecific with C. sp. pratti-like ofGrant et al. (2006; see alsoIbáñez et al., 2017) but were incorrectly identified as C. latinasus bySantos et al. (2009). SeeGrant et al. (2006: 135) regarding the terminal labeled as "Ameerega sp. ...
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Despite the impressive growth of knowledge on the phylogenetic systematics of dart-poison frogs and their relatives (Dendrobatoidea) over the past decade, many problems remain to be addressed. We analyzed up to 189 phenomic characters (morphology, behavior, defensive chemicals) and 15 mitochondrial and nuclear loci scored for 564 dendrobatoid and outgroup terminals, including 76 newly sequenced terminals and > 20 previously unanalyzed species, using tree-alignment and the parsimony optimality criterion in the program POY v.5.1.1 and additional analyses of the implied alignment using TNT v.1.5. Even though data coverage was highly heterogeneous, the strict consensus of 639 optimal trees is highly resolved and we detected only one instance of wildcard behavior involving a small clade of outgroup species. The monophyly of the median lingual process (MLP) possessing genus Anomaloglossus is decisively refuted, with the cis-Andean species being sister to Rheobates within Aromobatidae and the trans-Andean species nested within Hyloxalinae, implying two independent origins of the structure in Dendrobatoidea. Although this result was unexpected, it is not surprising given that the MLP evolved at least five times in Asian and African ranoids, including Arthroleptidae, Dicroglossidae, Mantellidae, and Rhacophoridae and either once in the most recent common ancestor of the massive clade Victoranura followed by independent losses or multiple times within component lineages. We restrict Anomaloglossus to the cis-Andean MLP-possessing species, describe a new genus for the trans-Andean MLP-possessing species, and resurrect Paruwrobates for its sister group, which includes Dendrobates andinus (formerly Ameerega), D. erythromos (formerly Hyloxalus and, until recently, Ameerega), and Prostherapis whymperi (formerly Hyloxalus). We also transfer Dendrobates maculatus from Ameerega to Epipedobates, making Ameerega an exclusively cis-Andean group. We describe two new species of the trans-Andean MLP-possessing genus-one from Cerro Tacarcuna, near the Colombo-Panamanian border, and the other from 800-900 m elevation on the western versant of the Colombian Cordillera Occidental (Cauca Department)-bringing the total number of species in the genus to seven. The discrete, round, white to yellowish-brown dots found on the venter of the new species from Cerro Tacarcuna and at least one other trans-Andean MLP-possessing species are formed by large, ellipsoid, densely distributed (up to 80 glands/mm²) granular glands. Although specimens of the new species from Cerro Tacarcuna exuded a noxious milky substance when handled, lipophilic alkaloids were not detected. In addition to the unexpected placement of the trans-Andean MLP-possessing species, major findings include the unexpected placement of Colostethus ruthveni and its undescribed sister species (the "C." ruthveni group) within Dendrobatinae as sister of the newly recognized tribe Dendrobatini (all dendrobatines except Phyllobates and the "C." ruthveni group). We describe a new genus for C. argyrogaster and C. fugax to remedy the paraphyly of Colostethus caused by the placement of those species as sister to Ameerega. Our evidence rejects the sister group relationship of Dendrobates + Oophaga in favor of Dendrobates + Adelphobates, which is consistent with their uniquely low diploid chromosome number of 2n = 18 (2n = 20 in Oophaga). With the exception of Anomaloglossus and Colostethus, all other genera are monophyletic. We recognize several monophyletic species groups-including the Atlantic Forest, trans-Andean, and 22-chromosome groups within Allobates, the An. stepheni, An. megacephalus, and An. beebei groups in Anomaloglossus, the C. latinasus (formed by the C. inguinalis and C. latinasus clades) and C. fraterdanieli groups within Colostethus, and the Am. braccata and Am. rubriventris groups within Ameerega-identify unambiguously optimized phenomic synapomorphies, and summarize patterns in the evolution of the diploid chromosome number, swelling of Finger IV in males, relative length of Fingers II and III, length of Finger V, and testicular and intestinal pigmentation. Finally, we address criticisms of the current taxonomy of Neotropical poison frogs and their relatives, concluding that they are either overstated, misguided, or false, and that the current system of names better communicates knowledge of the diversity of these frogs. Our results highlight the importance of increased taxon sampling, and we conclude by identifying key species to include in future phylogenetic analyses.
... Due to the isolation between the mountain ranges, these sites present numerous endemic species (Bermúdez et al. 2000;ANAM 2010). Cerro Pirre has been considered to harbor a high level of endemism, as it includes many endemic plant species (Luteyn 1976;Croat 1986bCroat , 1997Rojas-Alvarado 2002;Zhu and Croat 2004;Kolanowska et al. 2012;Monro 2012;Croat et al. 2017b;TROPICOS 2018) and endemic animals (Wake et al. 1970;Robbins et al. 1985;Stattersfield et al. 1998;Angehr et al. 2003;Batista et al. 2014;Hruska et al. 2016;Ibáñez et al. 2017;Renjifo et al. 2017). ...
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Ninety-four species belonging to 12 genera of Araceae are recorded on Cerro Pirre (Darién Province): Adelonema Schott (two); Anthurium Schott (39), Chlorospatha Engl. (2), Dieffenbachia Schott (3); Heteropsis Kunth (1); Monstera Adans. (6); Philodendron Schott (28), Rhodospatha Poepp. (2); Spathiphyllum Schott (2); Stenospermation Schott (3); Syngonium Schott (4) and Xanthosoma Schott (2). Taxonomic notes, identification key and comments on habitat and ecology of the species are presented.
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Vocalizations are one of the most important communication modalities in amphibian biology, and advertisement call is the type of vocalization mostly emitted by anurans (frogs and toads). In the present study, we carried out a scientometric analysis of the advertisement call in species of anurans in Colombia to determine the state of knowledge of this science in the country. We recorded the number of call descriptions and its trend throughout more than 60 years; in addition, we identified how many species there are with the call described per taxonomic family, geographic units and threat status. According to our review, between 1958 and June 30 of 2021, at least 296 studies have been published that describe the advertisement call of 307 species (of 785 in the country); only 130 descriptions come from recordings to individuals in Colombian populations. Leptodactylidae and Hylidae are the families with the highest percentage of species whose call has been described (84.6% and 68.4%, respectively). On the contrary, Bufonidae (19.5%) and Craugastoridae (12.9%) exhibit a low percentage of described calls. The Central and Eastern cordilleras were the regions with the best knowledge of advertisement calls, while the least known were the Caribbean – interAndean valleys, Orinoquía, and Sierra Nevada de Santa Marta. In terms of the threat status, the species with the least concern (LC) were the ones that presented the highest number of species with the call described. Despite an increase in the most recent two decades, our findings still reveal notable gaps in knowledge of the advertisement calls in Colombian anurans, which constitutes an incentive to develop future research on this subject. Finally, based on this meta-analysis, we highlight some recommendations that we hope that we hope stimulate new studies in ecoacoustics, using anurans in Colombia as a study model.
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For the past several decades, it has been thought that Colostethus inguinalis (Cope, 1868) (type species of Prostherapis Cope, 1868) is distributed in the Chocoregion of western Co- lombia and throughout much of Panama. This study shows that C. inguinalis is a Colombian endemic known only from the lowlands of the Chocoand Magdalena Valley—an unusual distribution pattern among dendrobatids but one shared with a several other anuran species typically known from the Chocoregion.Colostethus cacerensisRivero, 2000 ''1995'' is argued to be a junior synonym of C. inguinalis. The available name for the tetrodotoxin-possessing species found in Panama is C. panamensis (Dunn, 1933), which is redescribed. The first record of C. panamensis in Colombia is also reported. Colostethus inguinalis and C. panamensis differ from each other in ventral coloration of adult males and adult females, flank coloration, head coloration, relative tympanum size, and mean adult female snout-vent length. Colostethus latinasus (Cope 1863) (type species of Colostethus Cope, 1866) is most similar to C. pana- mensis but differs in a variety of characters, including ventral coloration and toe webbing. The exact provenance of the neotype of C. latinasus is unclear, but material that agrees closely with it was collected in Panama in the Serrania de Pirre; specimens previously reported asC. latinasus from Cerro Maliin the Serrania del Darien are not conspecific with that taxon and represent an undescribed species to be named elsewhere. It is doubtful that the Colombian holotype of C. latinasus (lost for over 80 years) was conspecific with the Panamanian neotype, and specimens that agree with the neotype have yet to be discovered in Colombia. Limited
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A problem frequently faced by researchers involved in collecting tissues for DNA isolation is the preservation of samples in the field prior to and during their transportation to the laboratory. Prevention of DNA degradation is usually achieved through freezing. As this is not always practical, we have tested the efficiency of chemical solutions containing high concentrations of salts (e.g., NaCl, EDTA, and diaminocyclohexanetetraacetate) and detergent at preserving DNA in bird tissue and blood samples stored at ambient temperature for extended periods of time. For blood samples, we recommend the use of a buffer that lyses the cells and nuclei and contains 0.01 M Tris, 0.01 M NaCl, 0.01 M EDTA, and 1% n-lauroylsarcosine, pH 7.5. Tissue samples are best preserved as small pieces in a saline solution made of 20% dimethyl sulfoxyde, 0.25 M EDTA, saturated with NaCl, pH 8.0. DNA extracted from samples preserved in these solutions for up to 24 weeks was compared with DNA recovered from tissue samples stored at -70-degrees-C and blood samples at -70 and -20-degrees-C. Yields were similar, averaging 300-mu-g/0.2 g of tissue and 500-mu-g/50-mu-L of blood. Quality of DNA in terms of fragment size, ability to be cut by restriction enzymes, and ability to hybridize to radioactive probes was also similar between cryopreserved and chemically preserved samples. Yields of DNA recovered from tissue samples preserved in 70% ethanol for 6 or 11 weeks was very low and significant degradation was observed. We have also examined how DNA contained in crude avian blood samples withstands freeze-thaw cycles. We found normal yields and no significant degradation of DNA in samples that experienced up to six cycles. We encourage field researchers who refrain from preserving tissue samples because of logistical problems, such as transporting liquid nitrogen containers in the field, to consider using these solutions. Both solutions can also facilitate exchanges of samples between laboratories, and they form an alternative to storage of samples at -70-degree-C for laboratories and museums with limited access to deep-freezing facilities.
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The taxonomic status of a very poorly known group of Andean frogs (the Eleutherodactylus discoidalis group) is assessed through acoustic and statistical analyses of differences in temporal parameters of advertisement calls, such as the number of pulses and the call duration, and also in a spectral parameter, dominant frequency. As these species are usually misidentified or ignored because of their taxonomic complexity in both ecological and biodiversity studies, we provide a bioacoustical diagnosis for each species in order to facilitate identification in the field. Differences in acoustic parameters support the specific status of Eleutherodactylus cruralis, E. discoidalis, Eleutherodactylus ibischi, and Eleutherodactylus madidi. The name E. cruralis is probably applied to three different species: the nominal form from Amazonian forests of the Andean slopes and adjacent lowlands, and two cryptic species restricted to inter-Andean dry valleys and cloud forests from central Bolivia. Moreover, the distribution of energy through the call and the aggregation of pulses seem to be useful meristic characters for detecting interspecific differences. Populations from each macrohabitat can be recognized by distinctive advertisement calls, usually corresponding to a recognized species. For the whole group, pulse rate is significantly correlated to latitude, which could indicate a speciation process along the Andes in relation to habitat changes and isolation. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 152, 353–365.
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1. A population ofColostethus inguinalis, a small diurnally active frog, was studied along a stream in seasonal tropical forest on Cerro Campana, Panama, from January through November 1976. Females laid eggs on land and carried up to 40 tadpoles on their backs to water. Tadpoles remained on their mothers' backs for up to 9 days. Sexual maturity was reached about 4–6 months after metamorphosis. 2. Some females produced two clutches in 11 months. The average interval between clutches was 133 days. Reproduction occurred in every month except April, with a peak in May and June, the beginning of the wet season. Reproduction decreased in the wettest months. 3. C. inguinalis has three vocalizations: an advertisement call, an encounter call used to challenge territorial intruders, and a close-range encounter call given just before an attack. The first two are given only by males, the last by females as well. 4. Agonistic behavior included postural displays, chases, attacks, and wrestling. Encounters between same sex individuals were more likely to end in wrestling than those between males and females. Males were more effective than females in repelling intruders without physical contact. 5. Courtship was prolonged and involved closerange vocal, visual, and tactile interactions between males and females. 6. The behavior and spatial organization of males and females changed seasonally. Both sexes defended small territories near pools in the dry season, although males remained in one place longer than most females. In the wet season, males defended widely dispersed territories ten times the size of dry season territories. Females were more mobile and seldom defended territories for more than a few days at a time. Dry season territories appeared to provide access to moist retreats and feeding sites in short supply. Large male wet season territories were used for feeding, shelter, courtship, and mating. 7. Both males and females defended territories against frogs of other species, but only encounters between maleC. inguinalis and maleColostethus pratti were frequent. Most of these occurred in the dry season, when the two species competed for the same retreats. There is some evidence that the two species can perceive each other's vocalizations.
Conference Paper
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Understanding the evolutionary history of living organisms is a central problem in biology. Until recently the ability to infer evolutionary relationships was limited by the amount of DNA sequence data available, but new DNA sequencing technologies have largely removed this limitation. As a result, DNA sequence data are readily available or obtainable for a wide spectrum of organisms, thus creating an unprecedented opportunity to explore evolutionary relationships broadly and deeply across the Tree of Life. Unfortunately, the algorithms used to infer evolutionary relationships are NP-hard, so the dramatic increase in available DNA sequence data has created a commensurate increase in the need for access to powerful computational resources. Local laptop or desktop machines are no longer viable for analysis of the larger data sets available today, and progress in the field relies upon access to large, scalable high-performance computing resources. This paper describes development of the CIPRES Science Gateway, a web portal designed to provide researchers with transparent access to the fastest available community codes for inference of phylogenetic relationships, and implementation of these codes on scalable computational resources. Meeting the needs of the community has included developing infrastructure to provide access, working with the community to improve existing community codes, developing infrastructure to insure the portal is scalable to the entire systematics community, and adopting strategies that make the project sustainable by the community. The CIPRES Science Gateway has allowed more than 1800 unique users to run jobs that required 2.5 million Service Units since its release in December 2009. (A Service Unit is a CPU-hour at unit priority).
The dendrobatid genus Silverstoneia is a clade of forest-dwelling frogs that share small adult size (<22 mm snout-vent length), basal webbing between toes a solid blackish brown flank divided by a conspicuous pale oblique lateral stripe from the groin to the eye, a pale ventrolateral stripe, finger I>II, male nurse frogs (known in four species), and larvae possessing an umbelliform oral disc with few to many submarginal papillae and medially emarginate, posterior labium (also known hi four species). The clade is distributed from Costa Rica to southern Departamento del Valle del Cauca in western Colombia and includes eight species, five of which are described herein as new species. As in all species of Dendrobatoidea, in Silverstoneia the distal tendon of insertion of the m. semitendinosus inserts dorsad to the distal tendon of the mm. gracilis complex and is strapped to the dorsal edge of the inner surface of the mm. gracilis complex by a unique binding tendon; The species of Silverstoneia may be diagnosed on the basis of adult size, adult ventral coloration, thigh coloration, and degree of expansion of finger III in adult males; additionally, there are clear species differences among the known tadpoles. Taxonomic comments are given for three previously named species: Silverstoneia erasmios (Rivero and Serna), S. flotator (Dunn), and S. nubicola (Dunn). We were unable to distinguish erasmios from nubicola. However, only females of erasmios are known and its validity needs confirmation. The species S. flotator sensu lato and S. nubicola sensu lato occur north through Panama to Costa Rica; they are distinct from one another, but some intraspecific variation suggests the possible presence of unnamed sibling species. The specimen long recognized as the type of S. flotator is not the holotype, which we consider lost; however, ample material is available from the well-known type locality (Barro Colorado Island) and neotype designation is not needed.
Colostethus flotator was described by E. R. Dunn in 1931 from specimens collected in the Canal Area of Panama. In 1933, Dunn synonymized this species with Colostethus nubicola, originally described by him in 1924 from western Panama, upon discovery of some apparently intermediate specimens from an intervening locality. However, two species referable to C. flotator and C. nubicola breed sympatrically in some locations in the Republic of Panama. We herein demonstrate that call characteristics, ventral coloration, and size differences indicate that C. flotator is a species separate from C. nubicola and raise C. flotator to specific status. /// Colostethus flotator fue descrita por E. R. Dunn en 1931 a partir de especimenes colectados en el Area del Canal de Panamá. En 1933 Dunn sinonimizó a esta especie con Colostethus nubicola, originalmente descrita por él en 1924 del oeste de Panamá, al descubrir algunos especímenes aparentemente intermediarios en una localidad intermedia. Sin embargo, las dos especies referidas como C. flotator y C. nubicola se reproducen simpátricamente en algunos lugares de la República de Panamá. Aquí se demuestra que diferencias en características del llamado, coloración ventral y tamaño, indican que C. flotator es una especie distinta de C. nubicola; por consiguente, elevamos a C. flotator al estado específico.
The strictly neotropical frog family Dendrobatidae is represented in Central America by three genera and 11 species. The bright colored, toxic poison arrow frogs belong to the genera Dendrobates and Phyllobates with five (auratus, granuliferus, minutus, pumilio, and speciosus) and one species (lugubris), respectively, in the region. Most of the species previously placed in Phyllobates, Prostherapis, or Hyloxalus are inconspicuous and apparently non-toxic forms for which Colostethus Cope 1867, is the first available name. Five Central American species (inguinalis, latinasus, nubicola, pratti, and talamancae) represent the latter genus in Central America. In all known members of the family eggs are laid on land and the larvae are carried to water on the back of a parent after hatching. Tadpoles of eight Central American species are known, those of C. inguinalis, C. latinasus, and P. lugubris here described for the first time. All Central American dendrobatids are found in wet evergreen forest situations and none ranges north of Nicaragua. The area of greatest species diversity is near the Colombia-Panamá boundary, where seven species (C. inguinalis, C. latinasus, C. nubicola, C. pratti, C. talamancae, D. auratus, and D. minutus) occur, but six of these forms comprise the dendrobatid fauna of central Panamá (only latinasus is lacking). In Costa Rica and western Panamá four species (C. nubicola, C. talamancae, D. auratus, and P. lugubris) are found on both Atlantic and Pacific lowlands. D. pumilio (Atlantic) and D. granuliferus (Pacific) apparently replace one another. C. pratti does not occur in Costa Rica but forms part of the Atlantic western Panamá fauna. Only D. auratus and D. pumilio range into Nicaragua. Seven of the species are restricted to the tropical lowlands; two lowland forms range into the subtropical zone, C. nubicola to its upper limit at 1700-1800 m and D. pumilio to 960 m. C. latinasus (1100-1400 m) and D. speciosus (1300-1400 m) are not known from lowland environments. As many as five species of dendrobatids occur sympatrically at localities on the Atlantic and Pacific lowlands of Costa Rica.
Science is about discovering new things, about better understanding processes and systems, and generally furthering our knowledge. Deep in science philosophy is the notion of hypotheses and mathematical models to represent these hypotheses. It is partially the quantification of hypotheses that provides the illusive concept of rigor in science. Science is partially an adversarial process; hypotheses battle for primacy aided by observations, data, and models. Science is one of the few human endeavors that is truly progressive. Progress in science is defined as approaching an increased understanding of truth – science evolves in a sense.
Goicoechea, N., De La Riva, I. & Padial, J. M. (2010). Recovering phylogenetic signal from frog mating calls. —Zoologica Scripta, 39, 411–154. Few studies have tried to analyse the phylogenetic information contained in frog mating calls. While some of those studies suggest that sexual selection deletes any phylogenetic signal, others indicate that frog calls do retain phylogenetic informative characters. Discordant results can be the outcome of disparate rates of character evolution and evolutionary plasticity of call characters in different groups of frogs, but also the result of applying different coding methods. No study to date has compared the relative performance of different coding methods in detecting phylogenetic signal in calls, hampering thus potential consilience between previous results. In this study, we analyse the strength of phylogenetic signal in 10 mating call characters of 11 related species of frogs belonging to three genera of Andean and Amazonian frogs (Anura: Terrarana: Strabomantidae). We use six quantitative characters (number of notes per call, note length, call length, number of pulses per note, fundamental frequency and dominant frequency) and four qualitative ones (presence/absence of: pseudopulses, frequency modulation in notes, amplitude modulation in notes and amplitude modulation in pulses). We code quantitative characters using four different coding and scaling methods: (i) gap-coding, (ii) fixed-scale, (iii) step-matrix gap-weighting with between-characters scaling, and (iv) step-matrix gap-weighting with between-states scaling. All four coding methods indicate that frog calls contain phylogenetic information. These results suggest that divergent selection on frog mating calls may not always be strong enough to eliminate phylogenetic signal. However, coding methods strongly affect the amount of recoverable information. Step-matrix gap-weighting with between-characters scaling and gap-coding are suggested as the best methods available for coding quantitative characters of frog calls. Also, our results indicate that the arbitrariness in selecting character states and the method for scaling transitions weights, rather than the number of character states, is what potentially biases phylogenetic analyses with quantitative characters.