Article ZOOTAXA Copyright © 2012 · Magnolia Press ISSN 1175-5334 (Online Edition)

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A new species of the Dendropsophus parviceps group from the western Amazon Basin (Amphibia: Anura: Hylidae)

ANA P. MOTTA1,2, SANTIAGO CASTROVIEJO-FISHER2, PABLO J. VENEGAS3, VICTOR G. D. ORRICO4 & JOSÉ M. PADIAL2 1Departamento de Biologia Animal, Museu de Zoologia João Moojen, Vila Gianetti 32, Universidade Federal de Viçosa, 36570-000, Viçosa, MG, Brazil. E-mail: [email protected] 2Department of Herpetology, American Museum of Natural History, Central Park West at 79th Street, New York, NY, 10024-5192, U.S.A. E-mails: SCF, [email protected]; JMP, [email protected] 3División de Herpetología-Centro de Ornitología y Biodiversidad (CORBIDI), Santa Rita N˚105 Of. 202, Urb. Huertos de San Anto- nio, Surco, Lima-Perú. E-mail: [email protected] 4Departamento de Zoologia, I.B, UNESP - Universidade Estadual Paulista Júlio de Mesquita Filho; Av. 24-A, num 1515, Bela Vista; Cx. P. 199 CEP: 13506-900 - Rio Claro, SP – Brazil. E-mail: [email protected]

Abstract

We describe Dendropsophus frosti sp. nov. from lowland terra firme rainforests of the headwaters of the Amazon River Basin in Colombia and Peru. The new species is known from only two localities, the type locality near Leticia (Departa- mento Amazonas, Colombia, 04° 06' 24.2" S, 69° 56' 57.4" W; 103 m.a.s.l.), and the paratopotypic locality, Piedras in the Putumayo basin (Departamento Loreto, Peru, 02.79278° S, 72.91750° W; 90–170 m.a.s.l.). Maximum likelihood and parsimony analyses of 2436 aligned base pairs of the 12S and 16S rRNA genes recovered the new species as a member of D. parviceps group and sister to D. brevifrons. The new species is most closely related to D. parviceps, D. brevifrons, and D. koechlini, and it can be readily distinguished from these and all other members of the D. parviceps group by, among other characters, its plain dorsal light brown coloration, copper iris, plain immaculate pale yellow to white venter coloration, lack of flash marks on groin and axillae, and absence of white spots on lips.

Key words: Amazon, Colombia, Peru, Dendropsophus frosti, new species

Introduction

The western Amazon Basin harbors the higher level of amphibian species diversity per area in the world (Bass et al. 2010), and fieldwork in the region is still revealing many additional new species (e.g. Köhler & Lötters 2001; Faivovich et al. 2006; Moravec et al. 2006, 2008; Brown & Twomey 2009; Padial & De la Riva 2009; Heinicke et al. 2009; Jungfer et al. 2010). However, in comparison to other groups of frogs for which new species discoveries are frequent in the area or elsewhere, few new Dendropsophus have been recently described. Indeed, only eleven species of Dendropsophus, five from the Amazonian lowlands, have been described since the beginning of the XXI century (Frost 2011). The last species described assigned to the D. parviceps group was D. gaucheri (Lescure and Marty 2000), but Fouquet et al. (2011) transferred it to the D. microcephalus group. The Dendropsophus parviceps group, as redefined by Faivovich et al. (2005), contains 14 species (Frost 2011; Fouquet et al. 2011), distributed in the Amazon from Bolivia to northern Venezuela, through Peru, Ecuador, Colombia, Brazil, Guyana, and French Guiana, and in Brazilian Atlantic Forest, from southern Bahia to northern Rio Grande do Sul. However, Faivovich et al. (2005) and Fouquet et al. (2011) highlighted the need of a more rig- orous test of the monophyly of the group. Subsequently, this group has been recovered as paraphyletic (Wiens et al. 2006, 2010; Moen & Wiens 2009; Pyron & Wiens 2011; Fouquet et al. 2011). Ten of these species occur in the Amazon [D. allenorum (Duellman and Trueb, 1989), D. bokermanni (Goin, 1960), D. brevifrons (Duellman and Crump, 1974), D. grandisonae (Goin, 1966), D. koechlini (Duellman and Trueb, 1989), D. luteoocellatus (Roux,

18 Accepted by M. Vences: 7 Feb. 2012; published: 28 Mar. 2012 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

1927), D. parviceps (Boulenger, 1882), D. pauiniensis (Heyer, 1977), D. schubarti (Bokermann, 1963), and D. timbeba (Martins and Cardoso, 1987)]. Here we describe a new species phenotypically similar to the species of the Dendropsophus parviceps group from terra firme lowland forests of the Upper Amazon plain in Peru and Colombia. Because no phenotypic diag- nostic character is known for the D. parviceps group (Faivovich et al. 2005), we performed maximum parsimony and likelihood phylogenetic analyses of DNA sequences of the 12S rRNA gene and a fragment of the 16S rRNA gene to assess group membership.

Material and methods

Collected specimens were fixed in 75 % ethanol (Colombian) or formalin 10 % (Peruvian) and stored in 70 % ethanol. Prior to fixation, tissue samples from all specimens were collected and deposited in 96% ethanol. Measure- ments were taken to the nearest 0.1 mm using digital calipers. Description of coloration in life is based on notes and digital color photographs taken in the field. Webbing formula follows the standards of Myers and Duellman (1982), whereas all other terminology is that of Duellman (1970). Measurement abbreviations used throughout the text are: SVL, snout-vent length; HL, head length as the straight line distance from the posterior edge of the jaw articulation to the tip of the snout; HW, greatest head width at midlevel of eyes; ED, horizontal eye diameter; ELW, upper eyelid width; ES, eye-tip of snout distance; TYD, horizontal tympanum diameter; DF3, width of disc on third finger; TL, tibia length; THL, thigh length and FL, foot length as the distance from proximal margin of outer metatarsal tubercle to tip of toe IV. For comparisons we used species descriptions, pictures of living specimens (Fig. 1), and specimens from collections. Specimens examined are listed in Appendix I. Museum acronyms are listed in Frost (2011), except ANDES (Museo de la Universidad de los Andes, Bogota) and CORBIDI (Centro de Ornitologia y Biodiversidad, Lima). We sequenced a fragment of the 16S rRNA gene and the complete 12S rRNA gene in five concatenate frag- ments for the holotype of the new species totalizing 2,476 bp (GenBank accession number: JQ088283). Molecular procedures (including primers) are the same as in Faivovich et al. (2005) but sequencing was performed at Macro- gen Inc. (Korea). Data from complementary strands were compared to generate a consensus sequence for each DNA fragment using Sequencher 4.1 (Gene Code Corp, Ann Arbor, USA). In order to assess the relationships of the new species within Dendropsophus, we downloaded homologous sequences of all species of the genus available in GenBank (Appendix II). These included six species of the D. parviceps group (D. allenorum, D. brevifrons, D. giesleri (Mertens, 1950), D. koechlini, D. parviceps, D. schubarti). We used Xenohyla truncata (Izecksohn, 1959) to root trees and Sphaenorhynchus dorisae (Goin, 1957), S. lacteus (Daudin, 1800) and S. orophilus (Lutz and Lutz, 1938) as outgroups following Faivovich et al. (2005). A static alignment of each independent gene was obtained using the web version of the software Mafft version 6.5 (Katoh et al. 2005) under the Q-INS-I strategy, after which the data was concatenated into a single matrix of up to 2436 bp for 42 terminals. Maximum parsimony analyses were performed in TNT (Goloboff et al. 2008) considering gaps as fifth character, and using New Technology searches at level 100, including sectorial searches, ratchet, drift and tree fusing. Bootstrap support (Felsenstein 1985) was evaluated through 500 replicates using New Technology. Maximum likelihood (ML) analyses were performed in Garli2.0 (Zwickl 2006) under default parameters. We implemented the model of nucleotide substitution GTR+I+G selected by ModelTest version 3.7 (Posada & Cran- dall 1998) following the Akaike Information Criterion (AIC). Hundred independent replicates where performed to assess consistency of results during the best tree search. Bootstrap support (Felsenstein 1985) was evaluated through 500 replicates.

Results

Maximum parsimony and maximum likelihood trees support the placement of the new species as sister to D. brevifrons (Fig. 2) with D. parviceps as the most closely related taxon to that pair of species. Although MP analysis recovered the monophyly of a group including D. giesleri, D. allenorum, D. koechlini, D. parviceps, D. brevifrons and the new species, the D. parviceps group, as currently defined, is paraphyletic because D. schubarti is not

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FIGURE 1. Living specimens of 11 species of the Dendropsophus parviceps group sensu Faivovich et al. (2005): A) D. allenorum from Madre de Dios, Peru (photo: W. E. Duellman); B) D. bokermanni from Sucumbios, Ecuador (photo: W. E. Duellman); C) D. luteoocellatus from Venezuela (photo: A. Acevedo); D, E) D. brevifrons from Leticia, Colombia (photos: J. M. Padial); F) D. microps from Brazil (photo: M. Sacramento); G, H) D. giesleri from Brazil (photos: L. Drummond); I) D. ruschii from Espirito Santo, Brazil (photo: P. Peloso); J, K) D. koechlini from Cruzeiro do Sul, Acre, Brazil (photos: T. Grant); L) D. schubarti from Cuzco Amazónico, Madre de Dios, Peru (photo: W. E. Duellman); M, N) D. parviceps from Leticia, Colombia (photos: J. M. Padial); O) D. subocularis from Santander, Colombia (photo: C. Hernandez-Jaimes). Photographs A, B and L are courtesy of the Biodiversity Institute, University of Kansas. Photographs are not at the same scale (i.e. they have no morphometric value) and are presented for coloration, shape, and skin structure comparisons.

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FIGURE 2. A) Maximum parsimony and B) Maximum likelihood best trees for 42 terminals, including 38 samples of 36 species of Dendropsophus, inferred from the 16S and 12S rRNA genes (see methods). Numbers above nodes are bootstrap values. In red are members of the D. parviceps group, and Dendropsophus frosti sp. nov. is highlighted in bold red.

NEW AMAZONIAN DENDROPSOPHUS Zootaxa 3249 © 2012 Magnolia Press · 21 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. closely related to that clade (Fig. 2A). ML analysis however, recovered only D. koechlini, D. parviceps, D. brevifrons and the new species in a clade, while D. allenorum, D. giesleri and D. schubarti are placed elsewhere (Fig. 2B).

Dendropsophus frosti sp. nov. (Figs. 3–4)

Dendropsophus koechlini, Fig. 6H—Von May and Venegas (2010).

Holotype. ANDES-A1025 (field number JMP 1986), an adult male from Km 11 on the road from Leticia to Tara- pacá (04° 06' 24.2" S, 69° 56' 57.4" W; 103 m.a.s.l.), Departamento Amazonas, Colombia (Fig. 3), collected on December 28, 2009 by Santiago Castroviejo-Fisher, José M. Padial, Björn Rogel, and Linn Fenna Groeneveld . Paratypes. ANDES-A1024 (field number JMP 2014) an adult male, same data as the holotype; ANDES- A1026 and ANDES-A1027 (field number AJC 3586–5 respectively) adult female and male with same data as the holotype but collected on December 9, 2011 by Justin Touchon. CORBIDI 05882–05883 (males), CORBIDI 05884 (female), from Peru: Loreto, Provincia de Maynas, Piedras (between the community of Nueva Vida and San Pablo de Totolla) (02.79278° S, 72.91750° W; 150 m) collected on October 25, 2009 by Pablo J. Venegas. Diagnosis. We assigned the new species to the genus Dendropsophus on the basis of our phylogenetic results (Fig. 2) and the overall similarity with other species of the genus (Fig. 1). We could not evaluate the two putative synapomorphies of the genus suggested by Faivovich et al. (2005). Dendropsophus frosti sp. nov. is a medium- sized member of the Dendropsophus parviceps group (sensu our phylogenetic results), (SVL 21.1–23.0 mm in adult males, 25.9–28.8 mm in a single female), diagnosed by the following combination of traits: slender body, head wider than body; snout truncate in dorsal and lateral views; nostrils slightly protuberant; large prominent eyes (EL/HW=0.4); palpebral membrane bearing brownish pigmentation on its border; small tympanum (TYD/ DF3=1.1); moderately developed axillary membrane; bifid distal subarticular tubercles on finger IV; prepollex con- cealed by skin, attached to finger I; nuptial excrescences not visible under a magnifying stereoscope in adult males; hands webbing formula I 2¯ —2¯ II 2½—2¯ III 2+—2 IV, feet webbing formula I 2½—2 II 1+—2+ III 1—2+ IV 2+— 1¯ V; no inner tarsal fold, tarsal tubercles absent; heel and calcar tubercles absent; cloacal opening covered by a cloacal sheath on its dorsal third; in life, dorsal surfaces presenting plain light brown coloration, ventral surfaces pale yellow, thighs and internal region of foot (through fingers III and IV) dark brown; lateral surfaces of body and head dark brown, darker in groin region; fingers I and II, and the tip of finger III pale, iris copper. In alcohol, dorsal surfaces pale brown; venter creamy white; iris gray. Comparison to other species. Dendropsophus frosti sp. nov. differs from all other members of Dendropso- phus by at least the combination of the following traits: plain light brown dorsal coloration, contrasting with dark brown flanks; fingers III and IV dark brown; fingers I and II and tip of finger II pale, and copper iris in life (Fig. 3). It specifically differs from species of the D. parviceps group (sensu Faivovich et al. 2005) as follows: from D. allenorum, D. giesleri, D. koechlini, D. microps Peters 1872, D. parviceps, D. pauiniensis, D. ruschii Weygoldt and Peixoto 1987, and D. timbeba by having a plain ventral coloration; from D. grandisonae by having a pale venter instead of gray. D. grandisonae also differs from the new species by presenting dorsal surfaces of the arms white and a more developed patagium. Dendropsohus frosti sp. nov. differs from D. bokermanni, D. pauiniensis, and D. subocularis Dunn 1934 by having ventral surfaces granular; from D. allenorum, D. giesleri, D. microps, D. pauiniensis, and D. ruschii by lacking any kind of tubercles, spiculae or warts on dorsum. The absence of blotches, spots or bars on surfaces of thighs and groin differentiates D. frosti sp. nov. from D. bokermanni, D. brevifrons, D. luteoocellatus, D. microps, D. pauiniensis, D. timbeba, and D. subocularis. The new species also differs from all the species in the D. parviceps group, except D. giesleri, D. grandisonae, and D. timbeba, by lacking suborbital bars. Females of D. brevifrons present a broad dorsolateral light stripe, which is absent in females of D. frosti sp. nov. Description of holotype. Adult male, SVL 22.8 mm; body slender; head wider than body, slightly wider than long, HW/HL 1.1, widest below eyes; snout truncate in dorsal and lateral views; canthus rostralis present, curved; loreal region flat; lips thin; nostrils slightly protuberant, directed anterolaterally. Interorbital area flat; eyes large (EL/HW=0.4) and protuberant; palpebral membrane translucent, with brown pigmentation on its border. Tympa- num small (TYD/DF3=1.1), distinct, directed laterally. Tympanic annulus and membrane evident, supratympanic

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FIGURE 3. Adult male holotype of Dendropsophus frosti sp. nov. ANDES-A1025, SVL = 22.8 mm. Photos of the live specimen illustrate coloration during daylight. Photos: preserved specimen S. Castroviejo-Fisher, live specimen J. M. Padial.

FIGURE 4. Peruvian paratypes of Dendropsophus frosti sp. nov. presenting the rostral stripe. Photos illustrate coloration during daylight. Photos: P. J. Venegas.

NEW AMAZONIAN DENDROPSOPHUS Zootaxa 3249 © 2012 Magnolia Press · 23 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. fold absent. Forearm slender, not hypertrophied, bearing a slight axillary membrane; fingers short, bearing round discs; relative length of fingers I < II < IV < III; subarticular tubercles small, round, bifid on finger IV, most prominent on finger I; supernumerary tubercles absent; inner metacarpal tubercle flat, elongate; outer metacarpal tubercle flat small, round; nuptial excrescences absent; webbing basal between fingers I and II; webbing formula I 2¯ — 2¯ II 2½— 2¯ III 2+—2 IV. Hind limb long and slender; TL/SVL = 0.5; no tarsal fold; calcar and heel tubercles absent; toes bearing round discs, smaller than those on fingers; relative lengths of toes I < II < III < V < IV; subarticular tubercles, round, elevated; supernumerary tubercles absent; inner metatarsal tubercle flat, elongate; outer metatarsal tubercle flat, round; webbing formula I 2½—2 II 1+—2+ III 1—2+ IV 2+—1¯ V. Skin on dorsum, head, dorsal surfaces of forearms and thighs, flanks and groin smooth; skin on belly and ventral surfaces of thighs granular. Cloacal opening directed posteriorly at midlevel of thighs, covered by cloacal sheath dorsally; cloacal tubercles absent. Tongue cordiform, barely free behind; dentigerous process of vomers evident, in two transverse series, positioned obliquely to choanae, each having two vomerine teeth; choanae large, rounded; vocal slits moderately long, extending from midlateral base of tongue, almost reaching to angle of jaws; vocal sac single, median, subgu- lar. In life, dorsal surfaces of body, head, arms and tibia, and the external region of foot (including toe V) light brown; thighs and internal region of foot (including toes III and IV) dark brown; lateral surfaces of body and head dark brown, darker than dorsal surfaces, with darkest region on groin; fingers I and II, and the tip of Finger III pale; venter pale yellow; iris copper. In alcohol, dorsal surfaces pale brown; venter creamy white; iris gray. The holotype is missing the right foot because it was cut at the level of heel as a tissue sample for molecular studies. Measurements of holotype (in mm): SVL 22.8, HL 8.1, HW 8.8, ED 3.8, ELW 1.8, ES 2.9, TYD 1.4, DF3 1.4, TL 12.3, THL 11.9, FL 9.9. Variation of type series. Type series is morphologically concurrent with the holotype, although the two females are larger than the males. Measurements of the type series are summarized in Table 1. The female COR- BIDI 05884 has four vomerine teeth per dentigerous process of vomer. At night the dorsal coloration of the Peru- vian specimens is yellow in males and pale brown in females without a contrasting coloration in the flanks or thighs. By day, the dorsal coloration of males and females are brown, ventral surfaces pale yellow, thighs and internal region of foot (including toes IV and V) and shanks dark brown (nearly black); toes I-III pale; lateral surface of body and head dark brown, with darkest region on groin; fingers I and II, and the tip of fingers III pale, iris copper. In life, all Peruvian paratypes presented a pale vertical stripe in the rostrum extending posteriorly as a pale narrow stripe from the rostrum to the tympanum through the canthus rostralis and eyelids, fading from the posterior margin of the eyelid over the dorsal margin of the tympanum (Fig. 4). Coloration in preservative resembles that of coloration in life during daylight. The pale vertical stripe in the rostrum of the Peruvian specimens is also apparent in preservative.

TABLE 1. Measurements (mm) and proportions of adult specimens of Dendropsophus frosti sp. nov. (mean and ± standard deviation in parentheses). males (n=4*) females (n=1*) SVL 21.1–25.2 (22.1±0.9) 25.9–28.8 HL 7.1–8.1 (7.5±0.4) 8.6 HW 7.3–8.8 (8.0±0.7) 8.2 ED 3.4–3.8 (3.6±0.2) 3.2 TYD 0.9–1.4 (1.2±0.2) 1.5 DF3 1.0–1.4 (1.2±0.2) 1.0 TL 11.8–12.3 (12.0±0.2) 12.9 TH 11.5–12.0 (11.8±0.3) 12.3 FL 9.5–9.9 (9.7±0.2) 10.3 TL/SVL 0.5–0.6 (0.6±0.0) 0.5 FL/SVL 0.4–0.5 (0.5±0.0) 0.4 HL/SVL 0.3–0.4 (0.3±0.0) 0.3 HW/SVL 0.3–0.4 (0.4±0.0) 0.3 HW/HL 1.0–1.1 (1.1±0.1) 1.0 EL/HW 0.4–0.5 (0.5±0.0) 0.4

*We only included SVL for paratypes ANDES-A1026–7.

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Distribution and ecology. The two localities are in terra firme Amazonian lowland forests close to large river channels (Fig. 5). Justin Touchon (pers. comm.) placed the amplectant pair ANDES-A1026–7 in a semi-natural enclosure consisting of a 1m diameter pool, floating vegetation and emergent vegetation, and surrounded by a nylon mesh cage supported by PVC poles ~2m tall. A thin, black cloth to increase shade covered the cage. The pair was left in the cage overnight and laid a single clutch of 70 eggs, 80 cm above the surface of the water, attached to the PVC supports. Although only a single observation, this indicates that they can and will lay terrestrial eggs attached to a rigid surface (such as a tree trunk in nature). Of course, they may also lay clutches on leaves or other arboreal substrate under different conditions. Justin Touchon carefully removed 5 eggs from the jelly, photo- graphed them on 5mm grid paper and measured their diameters in ImageJ. Average egg diameter was 1.9 mm (0.17 SD). The Peruvian locality is in an extensive complex of high terraces forest, at elevations of 90–170 m.a.s.l., close to the Algodoncillo River, which drains in the Algodón River, of the Putumayo basin. Specimens were found at night perching on low vegetation around ponds in primary forest. Several males were calling and two amplectant pairs were observed in the Peruvian locality, but calls were not recorded. Other Dendrosophus species occurring in the same area at both localities were D. rhodopeplus, including in the same ponds where the new species was collected. Etymology. The name is a patronym for Darrel Frost—a well-known North American herpetologist— in rec- ognition of his contribution to amphibian systematics, his encouragement, and for sharing his ample knowledge with us.

FIGURE 5. Map of western Amazon Basin showing the type (dot) and paratype (square) localities of Dendropsophus frosti sp. nov.

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FIGURE 6. Accumulation curve of named species of Dendropsophus according to Frost (2011).

Discussion

Although the monophyly of the D. parviceps group was doubted from the very beginning of its redefinition (Faivovich et al. 2005; Wiens et al. 2006, 2010; Moen & Wiens, 2009; Fouquet et al. 2011), none of these authors proposed a new group placement for species not recovered in a clade with D. parviceps (with the exception of D. gaucheri, proposed by Fouquet et al. 2011 to be transferred to the D. microcephalus group). In principle, at least, systematists should avoid paraphyletic classifications, but given that the relationships among species of Dendrop- sophus have low support and are contradictory among optimization criteria, authors as we herein, have probably kept recognizing the D. parviceps group to avoid spurious taxonomic actions. Nonetheless, D. parviceps is an Amazonian species, and several morphologically similar species from the Amazon belonging to the group cluster with it. Thus, even after an eventual splitting of the group to accommodate a monophyletic taxonomy, the nominal species group would remain as a small assemblage of morphologically similar Amazonian species including D. frosti, D. brevifrons, and D. koechlini, among other species that still need to be evaluated in future studies. The accumulation curve of described Dendropsophus species seems to be far from stabilization (Fig. 6). None- theless, few species of Dendropsophus have been recently described from the Amazonian lowlands (Frost 2011). Two facts suggest that the number of lowland Dendropsophus in the Amazon is far from known. On one hand, molecular (Fouquet et al. 2007) and integrative evidence (Jansen et al. 2011) have revealed multiple Dendropso- phus lineages that potentially represent new species. On the other hand, the finding of a new and conspicuous Den- dropsophus species in Leticia, a thoroughly sampled and highly diverse Amazonian locality harboring more than 100 species (Lynch, 2005; our own unpublished data), indicate that Amazonian forests can still harbor many new striking species, surely not only of Dendropsophus.

Acknowledgements

To the editor and two anonymous reviewers for improving the manuscript with their comments. To Goran and Adriana from Tanimboca tourism resort (Leticia, Colombia) for providing working facilities in the field. To our

26 · Zootaxa 3249 © 2012 Magnolia Press MOTTA ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited. guide in Leticia, Rodolfo Mesa, for helping during fieldwork and sharing his knowledge on the rainforest. To Björn Rogell and Linn Fenna Groeneveld for their friendship and help in the field. PJV is indebted with A. Del Campo by logistic support in the field. To Andrew J. Crawford, Adolfo Amézquita, Santiago Madriñan, Mauricio Bernal, Alvaro Velázquez, and Lucas Barrientos for helping with our work at the Universidad de los Andes. To Justin Tou- chon and Daniel Moen for sharing their unpublished data on Leticia specimens. To William E. Duellman and the Biodiversity Institute, University of Kansas, and to Aldemar Acevedo, Carlos Hernandez-Jaimes, Jean-Pierre Vacher, Leandro Drummond, Mário Sacramento, Moisés Barbosa de Souza and Pedro Peloso for allowing us to use their pictures. The work of SCF was financed by a Universidad de los Andes (2009–2010) and a Fulbright/Ministry of Education (2010–2012) post-doctoral research contracts. JMP’s research is founded by a Gerstner Postdoctoral Fellowship at the AMNH. VGDO thanks FAPESP for his PhD. scholarship (#2007/57067-9). Peruvian specimens were collected under permits 416-2009-AG-DGFFS-DGEFFS. Colombian specimens were collected under "Reso- lución 0082" of CorpoAmazonia. Fieldwork in northern Loreto (PJV) was part of a Rapid Biological Inventory led by the Field Museum, Chicago in collaboration with the Gobierno Regional de Loreto (GOREL).

References

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APPENDIX I. Additional specimens examined.

Dendropsophus allenorum: PERU: Madre de Dios: Reserva Cuzco Amazonico, 15 km ENE of Puerto Maldonado, 200m, KU 215189–90 (from pictures). Dendropsophus bokermanni: ECUADOR: Sucumbíos: Santa Cecília, KU 123089, 126552 (from pictures). Dendropsophus brevifrons: COLOMBIA: Putumayo, Santa Rosa de Sucumbio, Rio San Miguel, AMNH 88069–79; ECUA- DOR: Napo: Santa Cecília, 340m, KU 126370 (holotype) (from picture), AMNH 93187; PERU: Loreto, Rio Ampiyacu, Estiron, AMNH 114967, 114971–72, 114974. Dendropsophus giesleri: BRAZIL: Rio de Janeiro: Duque de Caxias, Barro Branco, Colonia de Imbarié, SMF 41217 (holotype) (from picture). Dendropsophus koechlini: BOLIVIA: La Paz: Heath River Wildlife Centre, at the Bolivian shore of Heath river, MNCN 44683–89; PERU: Departamento Madre de Dios: Reserva Cuzco Amazónico, on the Río Madre de Dios, approximately 15 km E of Puerto Maldonado, 200m (12° 33′ S, 69° 03′ W), ZUEC 14840 (ex-KU 205703) (from picture). Dendropsophus microps: BRAZIL: Rio de Janeiro: Nova Friburgo, ZMB 7472 (holotype) (from picture); Santa Catarina, AMNH 15573–82. Dendropsophus parviceps BRAZIL: Acre: km 23 on Rio Branco-Porto Velho road, AMNH 139315–17; Amazonas: Fazenda Sao Francisco, 0–1 km N km 49 on Manaus-Manacapuru road, AMNH 139314; BOLIVIA: Cusco: San Miguel, Valle de Marcapata, MNCN 43712; Loreto: Arboretum of Universidad Nacional de la Amazonia, MNCN 44903–11, 44875; Heath River Wildlife Centre, at the Bolivian shore of Heath river, MNCN 44701; Pando: Florida, MNCN 42946; Santa Cruz: Lomas de Arena, MNCN 42624; COLOMBIA: Putumayo: Santa Rosa de Sucumbio, Rio San Miguel, AMNH 88081–82; ECUADOR: Provincia Pastaza: Sarayacú, BMNH 1947.2.13.5 (holotype); Napo: Santa Cecilia, 340m, AMNH 93198–99; VENEZUELA: Amazonas: Solano, Casaquiare Canal, 75m, AMNH 118730–34, 118736–38. Dendropsophus pauiniensis: BRAZIL: Amazonas: Boca do Pauini, MZUSP 49892 (holotype). Dendropsophus schubarti: BRAZIL: Rondonia: Territorio de Rondonia, MNRJ 3669 (ex-WACB 7848) (holotype), MZUSP 73652 (ex-WCAB 7847) (paratype). Dendropsophus subocularis: PANAMA: Darien, AMNH 41117 (holotype), 51777, 79913. Dendropsophus timbeba: BRAZIL: Acre: Xapuri, 160m MZUSP 60560 (ex-ZUEC 5722) (holotype), ZUEC 5728 (paratype) (from picture).

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APPENDIX II. GenBank accession numbers for hylid frog sequences (12S and 16S rRNA) used for this study. 12S 16S Dendropsophus allenorum DQ380348 ― Dendropsophus anceps AY843597 AY843597 Dendropsophus aperomeus AY819450 AY819549 Dendropsophus berthalutzae ― AY843607 Dendropsophus bifurcus AY362975 AY362975 Dendropsophus bipunctatus AY843608 AY843608 Dendropsophus brevifrons AY843611 AY843611 Dendropsophus carnifex AY843616 AY843616 Dendropsophus ebraccatus AY843624 AY843624 Dendropsophus elegans DQ380355 AF308103 Dendropsophus giesleri AY843629 AY843629 Dendropsophus koechlini AY819369 ― Dendropsophus labialis AY843635 AY843635 Dendropsophus leali AY819451 ― Dendropsophus leucophyllatus DQ380360 AF308096 Dendropsophus marmoratus AY843640 AY843640 Dendropsophus microcephalus AY843643 AY843643 Dendropsophus minusculus DQ380362 ― Dendropsophus minutus ― AF308113 Dendropsophus miyatai AY843647 AY843647 Dendropsophus nanus AY819373 ― Dendropsophus parviceps AY843652 AY843652 Dendropsophus pelidna AY819434 ― Dendropsophus rhodopeplus AY843658 AY843658 Dendropsophus riveroi DQ380372 ― Dendropsophus robertmertensi AY819452 ― Dendropsophus rubicundulus AY843661 AY843661 Dendropsophus salli AY362976 AY362976 Dendropsophus sanborni AY843663 AY843663 Dendropsophus sarayacuensis AY843664 AY843664 Dendropsophus sartori AY819453 ― Dendropsophus schubarti DQ380374 ― Dendropsophus seniculus AY843666 AY843666 Dendropsophus triangulum AY843680 AY843680 Dendropsophus walfordi AY843683 AY843683 Sphaenorhynchus dorisae AY843766 AY843766 Sphaenorhynchus lacteus AY819394 ― Sphaenorhynchus orophilus DQ380388 ― Xenohyla truncata AY843775 AY843775