Corresponding author: Santiago R. Ron (
Academic editor: F. Andreone
We present a new phylogeny, based on DNA sequences of mitochondrial and nuclear genes, for frogs of the genus
Presentamos una nueva filogenia, basada en secuencias de ADN de genes nucleares y mitocondriales, para ranas del género
The Upper Amazon region has the highest alpha diversity of amphibians in the World with several sites exceeding 100 species in less than 10 km2 (
One such group is
Despite recent contributions to the taxonomy of the group (e.g.,
For ease of comparison, we generally follow the format of
Principal Components Analysis (PCA) and Discriminant Function Analysis (DFA) were used to assess the degree of morphometric differentiation between species. Only well preserved specimens (
Advertisement calls recordings were made with a Sennheiser™ ME-67 directional microphone with digital recorder Olympus™ LS10. Calls were analyzed using software Raven 1.2.1 (
Total DNA was extracted from muscle or liver tissue preserved in 95% ethanol or tissue storage buffer using standard phenol–chloroform extraction protocols (
We estimated phylogenetic relations between species of
Preliminary sequence alignment was done with MAFFT 6.814b software with the L-INS-i algorithm (
Because our dataset includes several loci, it is unlikely that it fits a single model of nucleotide substitution. Thus, we partitioned the data to analyze each partition under a separate model. The best model for each partition was chosen with JModelTest version 0.1.1 (
Each Bayesian analysis consisted of two parallel runs of the Metropolis coupled Monte Carlo Markov chain for 5 × 106 generations. Each run had four chains with a temperature of 0.05. The prior for the rate matrix was a uniform dirichlet and all topologies were equally probable a priori. Convergence into a stationary distribution was determined by reaching average standard deviation split frequencies < 0.05 between runs. We also used software Tracer ver. 1.5 (
Because the only nuclear gene analyzed had low variability and few informative sites, it was concatenated to the mitochondrial genes into a single matrix. We recognize the advantages of species-tree methods (e.g.,
Genbank accession numbers for DNA sequences used in the phylogenetic analysis.
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KU 143119 |
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QCAZ 15981 |
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LAC 2216 |
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-- | -- |
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CORBIDI 7458 |
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This study |
CORBIDI 7459 |
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This study |
CORBIDI 7462 |
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This study |
CORBIDI 7516 |
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This study |
LAC 2216 |
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QCAZ 14948 |
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This study |
QCAZ 24446 |
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QCAZ 24447 |
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QCAZ 28277 |
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QCAZ 28395 |
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QCAZ 28427 |
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This study |
QCAZ 36703 |
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This study |
QCAZ 39073 |
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This study |
QCAZ 39074 |
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This study |
QCAZ 39285 |
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This study |
QCAZ 43071 |
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This study |
QCAZ 48093 |
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This study |
QCAZ 48827 |
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This study |
AJC 2566 |
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This study |
AJC 2567 |
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This study |
CORBIDI 120 |
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This study |
CORBIDI 5819 |
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This study |
CORBIDI 5821 |
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This study |
LSUMZ H-13720 |
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QCAZ 27923 |
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This study |
QCAZ 28231 |
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CORBIDI 9368 |
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-- | This study |
CORBIDI 9370 |
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-- | This study |
CORBIDI 9394 |
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-- | This study |
CORBIDI 9507 |
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-- | This study |
QCAZ 25469 |
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QCAZ 31016 |
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This study |
QCAZ 31032 |
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This study |
QCAZ 31033 |
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This study |
QCAZ 32506 |
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QCAZ 32508 |
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QCAZ 37175 |
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QCAZ 39633 |
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This study |
QCAZ 40258 |
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This study |
QCAZ 45909 |
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This study |
QCAZ 46472 |
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This study |
QCAZ 49572 |
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This study |
CBF 6051 |
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NMP6d 28/2009 |
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NMP6V 73810/3 |
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NMP6V 73820 |
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QCAZ 20711 |
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This study |
NMP6V 71262/2 |
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CORBIDI 623 |
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CORBIDI 760 |
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This study |
CORBIDI 10461 |
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CORBIDI 1965 |
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This study |
CORBIDI 9585 |
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CORBIDI 9587 |
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QCAZ 38420 |
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QCAZ 39364 |
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QCAZ 41039 |
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QCAZ 20785 |
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CORBIDI 5505 |
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CORBIDI 8267 |
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This study |
CORBIDI 8284 |
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This study |
141 MC |
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AMNH-A 131254 |
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CORBIDI 4645 |
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This study |
CORBIDI 9369 |
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-- | This study |
KU 221930 |
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QCAZ 25603 |
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QCAZ 25684 |
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This study |
QCAZ 28223 |
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QCAZ 28646 |
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QCAZ 28647 |
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QCAZ 29430 |
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This study |
QCAZ 30925 |
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This study |
QCAZ 30926 |
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QCAZ 40253 |
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QCAZ 41030 |
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This study |
QCAZ 42999 |
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QCAZ 46470 |
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-- | This study |
QCAZ 46471 |
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This study |
14 MC |
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MNHN 2001.0828 |
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KU 221933 |
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NMP6V 71174/1 |
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NMP6V 71264/1 |
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NMP6V 71264/2 |
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QCAZ 20797 |
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214 MC |
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JM 2007/60 |
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KU 221941 |
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QCAZ 18230 |
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KU 205406 |
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CORBIDI 9477 |
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CORBIDI 9525 |
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KU 217751 |
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QCAZ 15942 |
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QCAZ 15991 |
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QCAZ 17285 |
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QCAZ 20544 |
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QCAZ 22201 |
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QCAZ 26304 |
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This study |
QCAZ 32032 |
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QCAZ 41108 |
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QCAZ 45344 |
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This study |
CORBIDI 4773 |
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This study |
QCAZ 14947 |
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QCAZ 27816 |
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QCAZ 27998 |
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NMP6d 41/2009 | -- |
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NMP6V 72173/1 | -- |
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NMP6V 72173/3 | -- |
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NMP6V 73105 | -- |
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QCAZ 35405 |
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This study |
QCAZ 38075 |
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This study |
Throughout this section, genetic distances are uncorrected
The topology (
All species within the
Populations of
The phylogeny recovers a monophyletic
Samples of
Clade D comprises five samples from four populations. For two individuals (from Brazil and French Guyana) only GenBank sequences were available and thus we cannot determine if they belong to
Post burn-in averages for parameters of Bayesian analyses. Abbreviations are: I = proportion of invariant sites, G = shape parameter of the gamma distribution of rate variation.
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GTR+G | – | 0.201 | 0.049 | 0.347 | 0.071 | 0.028 | 0.494 | 0.009 | 0.330 | 0.249 | 0.182 | 0.238 |
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SYM+I+G | 0.592 | 1.229 | 0.079 | 0.209 | 0.123 | 0.022 | 0.547 | 0.019 | – | – | – | – |
K80+I | – | – | – | – | – | – | – | – | – | – | – | ||
F81 | – | – | – | – | – | – | – | – | 0.168 | 0.269 | 0.155 | 0.408 | |
GTR+G | – | 2.972 | 0.024 | 0.651 | 0.027 | 0.027 | 0.236 | 0.035 | 0.272 | 0.324 | 0.093 | 0.310 | |
Control region | HKY+G | – | 0.367 | – | – | – | – | – | – | 0.399 | 0.183 | 0.085 | 0.333 |
HKY+G | – | 0.198 | – | – | – | – | – | – | 0.319 | 0.254 | 0.180 | 0.247 | |
HKY+I+G | 0.716 | 0.043 | – | – | – | – | – | – | 0.177 | 0.290 | 0.126 | 0.407 | |
GTR+G | – | 1.780 | 0.032 | 0.601 | 0.036 | 0.018 | 0.295 | 0.019 | 0.353 | 0.273 | 0.098 | 0.275 | |
POMC, 1st position | F81+G | – | 0.128 | – | – | – | – | – | – | 0.412 | 0.169 | 0.259 | 0.160 |
POMC, 2nd position | F81 | – | – | – | – | – | – | – | – | 0.419 | 0.183 | 0.199 | 0.199 |
POMC, 3rd position | HKY+I+G | 0.485 | 0.761 | – | – | – | – | – | – | 0.318 | 0.319 | 0.164 | 0.199 |
Bayesian consensus phylogram depicting relationships within
Records of
Variation in dorsal coloration of preserved specimens of adult
Variation in ventral coloration of preserved specimens of adult
Boxplots for snout-vent length (SVL; left) and the ratio tympanum diameter/snout-vent length (TD/SVL; right). The line in the middle of the box represents the median, and the lower and upper ends of the box are the 25% and 75% quartiles respectively. Each individual is shown with a symbol; the cross in
(
20 adult males, 1 adult female. QCAZ 33256, adult male, collected by I. G. Tapia, D. Almeida-Reinoso and M. Páez on 30 March 2007; QCAZ 39579, 39586–87, adult males, collected by D. Salazar-Valenzuela and G. Diaz between 12 and 14 December 2008; QCAZ 40909–10, adult males, collected by I. G. Tapia, L. A. Coloma, and S. R. Ron on 31 March 2008; QCAZ 40252, 40258, adult males, collected by D. Salazar-Valenzuela, D. Acosta-López and C. Korfel between 23 February and 1 March 2009; QCAZ 45271–72, 45277, 45281, adult males, collected by D. Acosta-López between 30 July and 2 August 2009; QCAZ 45907, 45909, adult males, collected by P. Peña-Loyola on 16 October 2009; QCAZ 49569–71, adult males, collected by N. Peñafiel between 26 June and 3 July 2010; QCAZ 49021–22, adult males, collected by R. Tarvin, and L. Bustamante on 3 August 2010; QCAZ 49439, adult female, collected by R. Tarvin and P. Aguilar on September 2010; QCAZ 48744 adult male, collected by S. R. Ron, L. Bustamante, I. G. Tapia , P. Peña-Loyola and R. Tarvin on 3 July 2010.
42 adult males, 2 adult females. Ecuador: Provincia Morona Santiago: Bobonaza (
Throughout this section, coloration refers to preserved specimens unless otherwise noted.
Adult male, 52.85 mm SVL, head length 18.61, head width 18.53, eye diameter 5.08, tympanum diameter 3.31, femur length 25.84, tibia length 30.05, foot length 22.73. Head narrower than body, slightly longer than wide; snout truncate in lateral and dorsal views; distance from nostril to eye longer than diameter of eye; canthus rostralis distinct and rounded; loreal region concave; internarial area depressed; nostrils moderately protuberant, directed laterally; interorbital area flat, lateral margins of the frontoparietals inconspicuous through skin; eye large, strongly protuberant; tympanic membrane clearly evident, large, slightly wider than high, about two thirds of eye diameter, separated from eye by ca. 85% of its diameter; tympanic annulus distinct except dorsally where it is covered by supratympanic fold; posterior end of supratympanic fold reaches arm insertion. Arm slender, axillary membrane present, reaching one third of arm length; four small low tubercles present along ventrolateral edge of forearm; relative length of fingers I<II<IV<III; fingers bearing large, oval discs, that of third finger about three fourths of tympanum diameter; subarticular tubercles prominent, round to ovoid, single; supernumerary tubercles present; palmar tubercle small, elongated; prepollical tubercle large, flat, elliptical; prepollex enlarged; large dark keratinous nuptial excrescences covering inner surface of prepollex up to half the distance between subarticular tubercle and proximal border of disk of thumb; webbing formula of fingers I basal II12/3—22/3III2½—2+IV. Medium sized to small tubercles on tibiotarsal articulation; scattered tubercles on tarsus, more abundant on outer edge; small tubercles scattered along ventrolateral edge of foot; toes bearing discs slightly wider than long, smaller than those of fingers; relative length of toes I<II<V<III<IV; outer metatarsal tubercle ill defined, small, round; inner metatarsal tubercle large, ovoid; subarticular tubercles single, round, protuberant; supernumerary tubercles restricted to the soles; webbing formula of toes I1—2II1—2+III1+—2+IV2–—1V. Skin on dorsum, head, and dorsal surfaces of limbs smooth, with scattered tubercles; skin on flanks areolate; skin on venter coarsely granular; skin on ventral surfaces of head and thighs granular, those of shanks smooth. Cloacal opening directed posteriorly at upper level of thighs; short simple cloacal sheath covering cloacal opening; round tubercles below vent; two conspicuous white tubercles ventrolateral to vent. Tongue cordiform, widely attached to floor of mouth; dentigerous processes of the vomer angular, adjacent medially, posteromedial to choanae, bearing 12 and 9 (left/right) vomerine teeth; choanae trapezoidal, oblique; vocal sac barely distinct above the arm and below the ear.
The specific name
Variation in dorsal and ventral coloration of preserved specimens is shown in
Ventral surfaces of preserved specimens (
Head shape is truncate in dorsal and lateral view (e.g., QCAZ 39579). Lateral head coloration varies between dark brown (QCAZ 49569) to cream (QCAZ 32506). There is a cream subocular mark. The tympanic annulus is concealed dorsally and has lighter color than the background. The distal subarticular tubercle on Finger IV is single (e.g., QCAZ 40909) or bifid (e.g., QCAZ 45272).
Morphometric data pertain to adults and are summarized in
Based on digital photograph of adult male QCAZ 48744 (
Based on digital photograph of juvenile QCAZ 40859 (
In life the Peruvian specimens have extensive blue coloration in the groins, concealed surfaces of thighs and tibia, dorsal surfaces of tarsus, armpits and posterior surfaces of arms (e.g., CORBIDI 10534;
Green coloration in life changes to cream in preserved specimens.
The advertisement calls of
Most specimens were collected at Río Pucayacu, a river surrounded by a mixture of primary and secondary forest. Frogs were found next to the river, perching over broad leaves or on tree branches 50 to 230 cm above the ground. At the collection site, the river has an average width of approximately 10 m, fast running water, and a rocky bottom. Males were calling next to the river between June 26 and July 3 2010. Several adults and a juvenile were found on a small stream, tributary of Río Rivadeneira, surrounded by secondary forest, near Río Pucayacu, in March 2008.
Vegetation types (according to the classification of
Specimens from Peru were collected in Cordillera de Kampankis within an elevational range of 300 to 365 m above sea level in tall, closed-canopy forest on low hills with well-drained soils at the base of the mountains. The soils have variable proportions of silt, clay and sand, but there are some small patches of sandy soil and limestone outcrops. The forest canopy is about 30 m tall, with emergent trees reaching 45 m. All individuals were collected in riparian vegetation of low-velocity and low-volume streams with rounded slate rocks lining the stream bed. Some individual were found on leafs of dense populations of rheophytic plants or shrubby
Records of
Dorsolateral and ventral views of
Adult
Adult
Lateral view of the head of the holotype of
Advertisement calls of
Ventral views of the left hand and foot of
(
(
(
Throughout this section, coloration refers to preserved specimens unless otherwise noted.
Adult male, 41.26 mm SVL, head length 12.79, head width 14.23, eye diameter 5.23, tympanum diameter 3.79, femur length 22.3, tibia length 23.1, foot length 17.97. Head narrower than body, slightly wider than long; snout rounded in dorsal view and truncate in lateral view; distance from nostril to eye longer than diameter of eye; canthus rostralis distinct and straight; loreal region concave; internarial area depressed; nostrils moderately protuberant, directed laterally; interorbital area with tiny keratinized conical tubercles, lateral margins of frontoparietals inconspicuous through skin; eye large, strongly protuberant; tympanic membrane clearly evident, large, slightly wider than high, about two thirds of eye diameter, separated from eye by ca. 85% of its diameter; tympanic annulus distinct except dorsally where it is covered by supratympanic fold; posterior end of supratympanic fold reaches arm insertion. Arm slender, axillary membrane present, reaching less than one third of arm length; four small low tubercles present along ventrolateral edge of forearm; relative length of fingers I<II<IV<III; fingers bearing large, oval discs, that of third finger about three fourths of tympanum diameter; subarticular tubercles prominent, round to ovoid except for bifid distal subarticular tubercle of Finger IV; supernumerary tubercles present; palmar tubercle small, elongated; prepollical tubercle large, flat, elliptical; prepollex enlarged; large dark keratinous nuptial excrescences covering inner surface of prepollex up to two thirds the distance between subarticular tubercle and proximal border of disk of thumb; webbing absent between fingers I and II; webbing formula of fingers II2–—3III2½ —3–IV. Small tubercles on tibiotarsal articulation; dorsal surface of tarsus covered by tiny keratinized conical tubercles, more abundant on outer edge; minute tubercles scattered along ventrolateral edge of foot; toes bearing discs slightly wider than long, smaller than those of fingers; relative length of toes I<II<V<III<IV; outer metatarsal tubercle ill defined, small, round; inner metatarsal tubercle low, ovoid; subarticular tubercles single, round, protuberant; supernumerary tubercles restricted to the soles; webbing formula of toes I1—2II1—2III1+—2IV2–—1V. Skin on dorsum, head, and dorsal surfaces of limbs shagreen covered by conical tubercles with keratinized tips, tiny on head and limbs; skin on flanks areolate; skin on venter coarsely granular; skin on ventral surfaces of head and thighs granular, those of shanks smooth. Cloacal opening directed posteriorly at upper level of thighs; short simple cloacal sheath covering cloacal opening; round tubercles below vent; two conspicuous white tubercles ventrolateral to vent at lower level of thighs. Tongue cordiform, widely attached to floor of mouth; dentigerous processes of the vomers angular, adjacent medially, posteromedial to choanae, bearing 5 and 6 (left/right) vomerine teeth; choanae trapezoidal, oblique; vocal slits moderately long, extending diagonally from lateral end of tongue toward to the angle of snout; vocal sac indistinct above the arm and below the ear.
Dorsum brown with irregular dark brown marks; flanks brownish cream with dark brown spots and flecks; dorsal surfaces of thighs, shanks, and forelimbs brown with transversal dark brown bands. Venter whitish cream with brown flecks in throat; ventral surfaces of thighs tan. Iris bronze with thin black reticulations (G. Chávez field notes April 2010).
The new species is dedicated to our colleague German Chávez (CORBIDI), one of the best friends of PJV, for his contributions to Peruvian herpetology and collecting the type series and tissues of this new species.
Variation in dorsal and ventral coloration of preserved specimens is shown in
Ventral surfaces of preserved specimens (
Snout is truncate in lateral view except for a female with rounded snout (CORBIDI 06633). Lateral head coloration varies from dull brown (CORBIDI 06633) to cream with dark brown blotches (CORBIDI 05505). The tympanic annulus is concealed dorsally and has lighter color than the background. The distal subarticular tubercle on Finger IV is bifid in all the specimens.
Adult morphometric data are summarized in
Based on digital photograph of adult male CORBIDI 06660: dorsum brown with irregular dark brown marks and some scattered light green blotches; canthal region greenish brown with greenish cream subocular mark and dark labial bars; tympanum light brown; flanks light green with dark brown reticulation and dark brown blotches posteriorly; dorsal surfaces of thighs, shanks, and forelimbs brown with transversal dark brown bands and scattered light green blotches. Iris bronze with brown horizontal midline and thin black reticulations.
Based on digital photograph of adult female CORBIDI 06633: dorsum brown with few scattered irregular dark brown marks; canthal region dark brown with greenish cream subocular mark speckled by three small dark brown blotches; tympanum light brown; flanks light brown with dark brown blotches; ventrolateral region cream with fine dark reticulation; dorsal surfaces of thighs, shanks, and forelimbs brown with transversal dark brown bands. Anterior half of venter whitish cream with fine brown reticulation in throat and chest; posterior half of venter and ventral surfaces of thighs tan; iris bronze with diffuse brown mid-horizontal line and thin black reticulations.
Based on digital photograph of adult female CORBIDI 05505 (
All specimens were collected next to temporary pools, perching over broad leaves or on tree branches 100 to 200 cm above the ground. Many streams surround the collection sites.
In the phylogeny (
Lateral view of the head of the holotypes of
Adult
Dorsolateral, frontal, and ventral views of
(
Five adult males: Ecuador: Provincia de Orellana: Pompeya-Iro road, km 80, Río Beye, QCAZ 51205, collected by E. Toral, I. G. Tapia, T. Camacho, and S. R. Ron on 31 May 2011; Provincia Pastaza: Nuevo Corrientes, 250 m above sea level, QCAZ 14947, collected by F. Villamarín on August 2000. Peru: Provincia Datem del Marañón: Andoas (
Throughout this section, coloration refers to preserved specimens unless otherwise noted.
A cream to bronze iris with black reticulations distinguishes
Adult male, 51.85 mm SVL, head length 18.9, head width 19.0, eye diameter 6.8, tympanum diameter 4.9, femur length 28.0, tibia length 28.7, foot length 22.1. Head narrower than body, nearly as wide as long; snout truncate in lateral and dorsal views; distance from nostril to eye longer than diameter of eye; canthus rostralis distinct and straight; loreal region concave; internarial area depressed; nostrils moderately protuberant, directed laterally; interorbital area flat, lateral margins of frontoparietals distinct through skin; eye large, strongly protuberant; tympanic membrane clearly evident, slightly wider than high, about two thirds of eye length, separated from eye by ca. 85% of its diameter; tympanic annulus distinct except dorsally where it is covered by supratympanic fold; posterior end of supratympanic fold reaches mid arm insertion. Arm slender, axillary membrane present, reaching one third of arm length; three small low tubercles present along ventrolateral edge of forearm; relative length of fingers I<II<IV<III; fingers bearing large, oval discs, that of third finger about three fourths of tympanum diameter; subarticular tubercles prominent, round to ovoid, bifid in distal subarticular tubercle of Finger IV; supernumerary tubercles present; palmar tubercle small, elongated; prepollical tubercle large, flat, elliptical; prepollex enlarged; large dark keratinous nuptial excrescences covering inner surface of prepollex almost reaching the proximal border of disk of thumb; webbing basal between fingers I and II; webbing formula of fingers I basal II1½—2½III2+—2IV. Medium sized to small tubercles on tibiotarsal articulation; scattered low tubercles on tarsus, more abundant on outer edge; small tubercles scattered along ventrolateral edge of foot; toes bearing discs slightly wider than long, smaller than those of fingers; relative length of toes I<II<V<III<IV; outer metatarsal tubercle ill defined, small, round; inner metatarsal tubercle large, ovoid; subarticular tubercles single, round, protuberant; supernumerary tubercles restricted to the soles; webbing formula of toes I1—2–II1—2III1—2IV2—1–V. Skin on dorsum, head, and dorsal surfaces of limbs shagreen, with scattered tubercles; minute keratinized conical tubercles present on the eyelids and dorsal surface of head; skin on flanks areolate with big flattened warts; skin on venter coarsely granular; skin on ventral surfaces of head and thighs granular, that on shanks smooth. Cloacal opening directed posteriorly at upper level of thighs; short simple cloacal sheath covering cloacal opening; round tubercles below vent; two distinct white tubercles ventrolateral to vent. Tongue cordiform, widely attached to floor of mouth; dentigerous processes of the vomers angular, adjacent medially, posteromedial to choanae, bearing 9 and 6 (left/right) vomerine teeth; choanae trapezoidal, oblique; vocal slits short and curved posteroventral to the angle of snout at the base of tongue; vocal sac barely distinct above the arm and below the ear.
The specific name is a patronym for Vilma Duran, in recognition of her continued work and efforts toward the improvement of the herpetological collection of CORBIDI and also for collecting the holotype and tissue of this new species.
Dorsal and ventral coloration of preserved specimens is shown in
Ventral surfaces of preserved specimens (
Head shape is truncate in dorsal view and truncate in lateral view. Lateral head coloration varies from light brown with dark mottling (CORBIDI 01086) to grayish white with dark brown canthus rostralis and preocular stripes (CORBIDI 05031). All specimens have a white to cream subocular mark. The tympanic annulus is concealed dorsally and has lighter color than the background. The distal subarticular tubercle on Finger IV is bifid in all specimens.
Adult morphometric data are summarized in
Based on a digital photograph of adult male CORBIDI 01086: dorsum light brown with irregular dark brown and light green marks; canthal region greenish brown with white subocular mark and dark brown band along posterior half of upper lip; tympanum pink contrasting with dark brown tympanic annulus; flanks light green with dark brown reticulation anteriorly and few irregular dark brown blotches posteriorly; dorsal surfaces of thighs, shanks and forelimbs brown with dark brown transversal bands; posterior surfaces of thighs light green; venter white speckled with light brown blotches; iris light cream with brown mid-horizontal line and fine black reticulations.
Descriptive statistics for morphometric measurements of species of the
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42.54 ± 2.51(39.66–45.72) | 17.46 ± 0.92(16.15–18.57) | 15.33 ±0.73(14.33–16.29) | 14.86 ± 0.73(14.11–15.66) | 4.11 ± 0.38(3.54–4.56) | 3.4 ± 0.34(3–3.9) | 23.62 ± 1.04(22.37–25.14) | 21.93 ± 1.11(20.09–22.9) | |
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Females ( |
66.55 ±5.44(62.64–72.77) | 28.31 ± 2.69(26.12–31.32) | 21.68 ± 1.25(20.76–23.11) | 18.36 ± 1.59(17.3–20.2) | 5.86 ± 0.43(5.42–6.28) | 3.85 ± 0.16(3.7–4.02) | 37.15 ± 3.25(34.17–40.62) | 34.64 ±2.19(32.89–37.11) |
BobonazaMales ( |
44.79 ± 2.72(42.86–46.72) | 18.55 ± 0.80(17.98–19.12) | 15.34 ± 1.39(14.36–16.33) | 12.85 ± 0.45(12.53–13.17) | 4.94 ± 0.12(4.85–5.03) | 2.91 ± 0.18(2.78–3.04) | 4.82 ± 1.64(23.66–25.98) | 21.34 ± 0.69(20.85–21.83) |
PomonaMales ( |
49.65 ± 5.54(43.67–54.61) | 21.06 ± 2.69(18.5–23.88) | 17.66 ± 1.72(15.27–18.31) | 14.29 ± 1.41(12.66–15.23) | 5.02 ± 0.68(4.44–5.78) | 3.21 ± 0.36(2.83–3.56) | 26.08 ± 2.98(23.52–29.36) | 24.21 ±2.86(21.52–27.22) |
YawiMales ( |
41.86 ± 2.91(38.46–45.46) | 17.11 ± 0.78(15.96–18.1) | 14.65 ± 0.95(13.86–16.24) | 12.53 ± 0.98(11.39–13.67) | 4.61 ± 0.42(4.24–5.34) | 2.92 ± 0.47(2.16–3.48) | 22.41 ± 1.21(20.68–23.77) | 20.22 ± 1.05(18.87–21.69) |
Female ( |
64.25 | 27.5 | 20.76 | 17.58 | 5.42 | 3.7 | 36.66 | 32.89 |
ZanjarajunoMales ( |
50.07 ± 0.96(49.39–50.75) | 21.46 ± 1.34(20.51–22.41) | 17.73 ± 0.07(17.68–17.79) | 14.15 ± 0.16(14.03–14.27) | 5.07 ± 0.09(5.01–5.14) | 3.24 ± 0.16(3.13–3.36) | 28.17 ± 2.24(26.59–29.76) | 24.75 ± 1.36(23.79–25.75) |
Female ( |
72.77 | 31.32 | 23.11 | 20.2 | 6.28 | 4.02 | 40.62 | 37.11 |
41.34 ± 2.41(38.01–45.25) | 16.42 ± 1.07(14.51–18.34) | 14.46 ± 0.74(13.05–15.82) | 12.49 ± 1.26(10.84–15.35) | 4.26 ± 0.30(3.76–4.84) | 3.51 ± 0.19(3.20–3.88) | 22.05 ± 1.21(20.07–24.24) | 20.14 ± 1.20(17.76–22.37) | |
Females ( |
45.68 ± 7.44(40.42–50.95) | 18.06 ± 1.52(16.99–19.14) | 16.08 ± 2(14.67–17.5) | 13.47 ± 1.49(12.42–14.53) | 4.69 ± 0.53(4.32–5.07) | 3.66 ± 0.2(3.52–3.81) | 25.14 ± 3.63(22.57–27.71) | 23.49 ± 2.8(21.51–25.47) |
41.26–41.45 | 17.97–18.17 | 12.79–12.99 | 14.23–14.82 | 4.51–5.23 | 3.79–3.99 | 23.10–23.50 | 22.30–22.70 | |
Females ( |
49.16–50.76 | 21.00–22.10 | 13.67–15.00 | 17.23–17.67 | 5.10–5.35 | 3.80–4.17 | 26.80–27.70 | 25.00–27.00 |
50.74 ± 3.17(48.23–55.77) | 21.06 ± 1.16(19.61–22.11) | 16.78 ± 1.32(14.90–18.09) | 18.03 ± 1.13(16.46–19.22) | 6.092 ± 0.62(5.27–6.80) | 4.43 ± 0.29(4.10–4.90) | 27.90 ± 0.64(27.00–28.70) | 25.93 ± 1.50(24.20–28.00) |
Descriptive statistics for call parameters of
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Combined(n = 5) | Río Piraña(n = 1) | Río Pucayacu(n = 4) | Jatun Sacha(n = 2) | |
Duration of first component note | 0.425 ± 0.053(0.356–0.489) | 0.356 | 0.442 ± 0.042(0.389–0.489) | 0.059 ± 0.004(0.056–0.063) |
Call Rate | 0.3066 ± 0.113(0.208–0.454) | 0.454 | 0.269 ± 0.090(0.208–0.402) | 1.524 ± 0.151(1.417–1.631) |
First component interval | 4.114 ± 1.722(2.142–6.004) | 2.142 | 4.607 ± 1.528(2.568–6.004) | 0.725 ± 0.140(0.625–0.824) |
Dominant Frequency | 1049.54 ± 247.18(771.6–1412.6) | 771.616 | 1119.02 ± 221.99(765.68–1472.26) | 745.66 ± 0.87(745.04–746.28) |
Number of pulses | 12.213 ± 1.585(9.8–14.2) | 12 | 12.266 ± 1.825(9.8–14.2) | 3.328 ± 0.181(3.2–3.457) |
Pulse rate | 28.932 ± 4.095(23.847–34.016) | 33.661 | 27.749 ± 3.610(22.004–33.495) | 55.833 ± 1.565(41.772–69.893) |
Duration of second component | 0.307 ± 0.106(0.216–0.488) | 0.216 | 0.329 ± 0.108(0.25–0.488) | NA |
Duration of second component note | 0.032 ± 0.004(0.027–0.037) | 0.027 | 0.033 ± 0.004(0.027–0.037) | NA |
Number of second component notes | 0.866 ± 0.339(0.445–1.287) | 1 | 0.832 ± 0.381(0.225–1.439) | NA |
Quack rate | 0.140 ± 0.026(0.108–0.177) | 0.108 | 0.148 ± 0.022(0.125–0.177) | NA |
Ventral views of left hand and foot of
Adult male
Three components with eigenvalues > 1.0, accounting for 70.9% of the total variation, were extracted from the PCA for males (
Three components with eigenvalues > 1.0 were extracted from the PCA for females (
In the DFA classification on males, all
Character loadings and eigenvalues for Principal Components (PC) I–III. The analysis was based on seven morphometric variables of adult
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Femur length |
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0.126 | 0.052 |
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0.030 | 0.193 |
Foot length | 0.425 | –0.400 | 0.046 | 0.451 | 0.374 | –0.106 |
Head length | 0.106 | 0.459 |
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–0.004 | –0.326 |
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Head width | 0.433 | –0.053 | –0.480 | 0.161 |
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–0.083 |
Eye diameter | 0.173 |
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–0.332 | –0.018 | 0.349 |
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Tympanum diameter | 0.066 |
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–0.233 | –0.376 | 0.421 | 0.201 |
Tibia length |
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–0.108 | 0.298 |
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–0.164 | 0.080 |
Eigenvalue | 2.451 | 1.432 | 1.083 | 2.311 | 1.768 | 1.222 |
Principal components from analysis of seven size-corrected morphological variables. See Table 5 for character loadings on each component.
Similarly to previous studies on Amazonian amphibians (e.g.,
Genetic evidence is a valuable taxonomic tool but, in most cases, is insufficient to define species boundaries without reference to other sets of characters like advertisement calls or external morphology. Taxonomic reviews of Amazonian amphibians suggest that morphological characters are too conservative to define species boundaries because closely related species share similar morphology (e.g.,
We suspect that the difficulty in defining species boundaries based on morphology arises from the high intraspecific polymorphism in coloration characteristic of most groups of dull-colored Amazonian amphibians like
Most
Examination of the geographic ranges of sister species can provide insights into modes of speciation. Our phylogeny of the
At the intraspecific level, we found low genetic divergence with the only exception of
This study was supported by grants from the Secretaría Nacional de Educación Superior, Ciencia, Tecnología e Innovación de Ecuador SENESCYT (PI-C08-0000470) and Dirección General Académica of Pontificia Universidad Católica del Ecuador. For the loan of specimens and access to collections we are indebted to A. Almendáriz and Barry Clarke. Santiago Castroviejo y J. M. Padial provided tissues of Colombian