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Anura: Hemiphractidae: Gastrotheca) from the Andes of Southern Peru

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Anura: Hemiphractidae: Gastrotheca) from the Andes of Southern Peru 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. Zootaxa 3095: 1–14 (2011) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2011 · Magnolia Press ISSN 1175-5334 (online edition) A new species of marsupial frog (Anura: Hemiphractidae: Gastrotheca) from the Andes of southern Peru WILLIAM E. DUELLMAN1,3, ALESSANDRO CATENAZZI2 & DAVID C. BLACKBURN1 1Biodiversity Institute, University of Kansas, Lawrence, KS 66045-7561 USA. E-mail: [email protected]; [email protected] 2Department of Biology, Gonzaga University, 502 E. Boone Ave., Spokane, WA 99201 USA. E-mail: [email protected] 3Corresponding author Abstract Gastrotheca nebulanastes sp. nov. from cloud forests in the upper Kosñipata Valley, Manu National Park, in the Andes of southern Peru is similar to G. excubitor, which inhabits grasslands in higher elevations than the cloud forests. The two spe- cies differ in relative lengths of the fingers, skin texture, coloration, and advertisement call. Although the new species has an elevational range of 2000–3300 m, it is most abundant at 2400–2800 m. A phylogenetic analysis of a previously defined clade of Gastrotheca based on a fragment of 16S mitochondrial gene provides strong support that the sister taxon to the new species is G. atympana, a species from farther north in the Cordillera Oriental in Peru. Key words: Anura, distribution, ecology, Gastrotheca, Hemiphractidae, new species, phylogenetic relationships Resumen Gastrotheca nebulanastes sp. nov. de los bosques nublados de la parte alta del valle de Kosñipata, Parque Nacional del Manu en los Andes del sur de Perú se asemeja a G. excubitor, una especie que vive en la puna a elevaciones por encima del límite superior del bosque nublado. Las dos especies se diferencian por la longitud relativa de sus dígitos, textura de la piel, coloración y canto nupcial. A pesar de tener un rango de distribución altitudinal entre los 2000 y 3300 m, la nueva especie es más abundante a elevaciones entre 2400–2800 m. El análisis filogenético de un fragmento del gen mitocondrial 16S de un clado de Gastrotheca previamente definido apoya la hipótesis de que el taxon hermano de la nueva especie es G. atympana, una especie distribuida más al norte en la Cordillera Oriental en Perú. Palabras claves: Anura, distribución, ecología, Gastrotheca, Hemiphractidae, nueva especie, relaciones filogenéticas Introduction In 1971 the senior author and Thomas H. Fritts discovered a distinctive, terrestrial species of Gastrotheca at the crest of Abra Acanacu (now Abra Acjanaco), a pass in the Cadena del Paucartambo, the northwestern part of the Cordillera de Carabaya, which is a part of the extensive Cordillera Oriental of the Andes of Peru and now protected as part of Manu National Park. They found the new species, G. excubitor (Duellman & Fritts, 1972) in wet puna also inhabited by the widespread G. marsupiata (Duméril & Bibron, 1841). Subsequently, both species were col- lected again at Abra Acanacu in 1975 and 1977, but during the latter trip specimens referred to G. excubitor were also found in humid montane forests at lower elevations northwest of Abra Acjanaco, as well as at other abras at elevations above 3500 m in Departamento (= Región) Cusco in southern Peru. More recent fieldwork (2007 and 2009) by one of us (AC) resulted in the collection of more individuals at localities in the humid montane forest, as well as the discovery that Hyla antoniiochoai De la Riva and Chaparro (2005) is an arboreal species of Gastrotheca in the upper Kosñipata (formerly Cosñipata) Valley (Catenazzi & Lehr 2009). Accepted by S. Castroviejo: 12 Oct. 2011; published: 10 Nov. 2011 1 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. Material and methods The 16 morphological measurements (Table 1), 25 external descriptive characters, and numbered diagnostic char- acters are those used by Duellman and Pyles (1980), Duellman and Hillis (1987), Duellman and Trueb (1988), and Duellman et al. (2001, 2004, 2006). All measurements are in millimeters; snout–vent length is abbreviated SVL. If sex could not be determined by the presence of a brood pouch or vocal sac, determination was made by examina- tion of gonads. Fourteen morphological measurements were retained for Principal Component Analysis (Table 2) to maximize sample size of n =18 for G. nebulanastes and n = 23 for G. excubitor. Morphological data (non-trans- formed, after checking for normality) were analyzed with the princomp function using eigen on the correlation matrix in the stats package in R 2.10.1 (The R Foundation for Statistical Computing; http://www.R-project. org). Principal components 1 and 2 (representing 60.2% of variation) were used to produce a scatter plot. Photographs of frogs taken by Catenazzi and Duellman were used in the description of color in life; these photos are available at the Calphoto online database (http://calphotos.berkeley.edu). Specimens of other species examined and/or used in the phylogenetic analysis are listed in Appendix I. We recorded advertisement calls of male MUSM 28060 (SVL 42.0 mm) at the type locality between 1900 and 2030 on 4 April 2011. The air temperature during that time was 10.7–12.3ºC. We used a digital recorder (Song Meter SM1, Wildlife Acoustics, Inc., Concord, Massachusetts) and recorded advertisement calls in uncompressed .wav format. We recorded the advertisement call of several males (exact number unknown; males were not cap- tured) of G. excubitor at Abra Acjanaco on 23–31 January 2009 (recorded between 2030 and 2230 and at tempera- tures of 7.5–9.0°C). We used Cool Edit version 96 (Syntrillium Software Corporation) and Raven Lite, version 1.0 (Cornell Laboratory of Ornithology) to resample recordings at a sampling frequency of 44 KHz, FFT width 512 points, and 16-bit resolution and analyze call variables. The Hamming window function for the spectrogram was set at 256 bands. Averages are reported ± SD. We analyzed a total of 20 calls for G. nebulanastes and 30 calls for G. excubitor. The following institutional abbreviations are used: AMNH = American Museum of Natural History, New York; FML = Fundación Miguel Lillo, Tucumán, Argentina; KU = Museum of Natural History, University of Kan- sas, Lawrence; LM = Linda Maxson tissue collection, Pennsylvania State University; MNCN = Museo Nacional de Ciencias Naturales, Madrid, Spain, MTD = Museum für Tierkunde Dresden; MHNC = Museo de Historia Natural, Universidad Nacional San Antonio Abad, Cusco, Peru; MUSM = Museo de Historia Natural, Universidad Nacio- nal Mayor de San Marcos, Lima, Peru; MVZ = Museum of Vertebrate Zoology, University of California, Berkeley; TNHC = Texas Natural History Collection, University of Texas, Austin; and USNM = National Museum of Natural History, Washington. Spelling of Acjanaco and Kosñipata follows terms currently used in Peru and in recent publi- cations; however, spellings in Carta Nacional “Calca,” Hoja 27-s (Instituto Geográfico Nacional, Lima) are Acanacu and Cosñipata. We determined the phylogenetic relationships of the new species through analysis of DNA sequence data for the mitochondrial 16S ribosomal RNA gene. We obtained data from three specimens of the new species of Gas- trotheca, as well as topotypic specimens of G. excubitor (MUSM 26280–81), G. antoniiochoai (MUSM 27944), and the recently described G. pachachacae (MUSM 28492). We amplified the 16S gene using polymerase chain reaction (PCR) and the primers (16Sc and 16Sd) of Darst and Cannatella (2004), resulting in 810–880 bp of sequence per specimen. Genomic DNA extraction, as well as purification of PCR products and sequencing follows the methods of Esselstyn et al. (2008). Newly collected sequence data are accessioned in GenBank. Details on the taxa and the corresponding GenBank sequences used are provided in Appendix II. We generated a multiple alignment of 16S data using MAFFT v6.850 (Katoh et al., 2005) with minor adjust- ments made by eye; the alignment used for analysis is deposited in Dryad (doi:10.5061/dryad.bt760). We analyzed the 16S alignment as a single partition. Using the Akaike information criterion (AIC) and jModeltest v.0.1.1 (Pos- ada, 2008), we selected the GTR + I +G model (lnL = –2648.47, AIC = 5388.93) as the best-fit model of sequence evolution (vs. next best, GTR + G: lnL = –2656.18, AIC = 5396.36). We estimated phylogenetic relationships using maximum-likelihood (ML) as implemented in RAxML v.7.0.4 (Stamatakis 2006). This analysis used a random starting tree, the faster rapid hill-climbing algorithm proposed by Stamatakis et al. (2007), and the GTR + I +G model of sequence evolution. The ML analysis used 100 search rep- etitions and we used the phylogenetic estimate with the smallest –ln likelihood score as the preferred ML phylog- eny. We performed one thousand nonparametric bootstrap replicates in RAxML using the same model of sequence 2 · Zootaxa 3095 © 2011 Magnolia Press DUELLMAN 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. evolution with one search replicate per bootstrap replicate, a random starting tree, and optimizing branch lengths and model parameters during the bootstrap analysis. We calculated split support using SumTrees in DendroPy (Sukumaran & Holder 2010). Based on the analysis of Wiens et al. (2007), we rooted the resulting phylogeny using Gastrotheca zeugocystis. Description of new species Gastrotheca nebulanastes new species Holotype: KU 173210, an adult female (Fig. 2A), from Buenos Aires (Fig. 5A), on Paucartambo–Pilcopata road, 2280 m, 13˚09'16" S, 71˚35'11"W, Distrito de Kosñipata, Provincia de Paucartambo, Región de Cusco, Peru, one of a series obtained on 19 January 1977, by W. E.
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