The Unexpected Trans-Andean Distribution of Cochranella Resplendens (Lynch and Duellman, 1978) (Anura: Centrolenidae)
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Zootaxa 4238 (2): 268–274 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Correspondence ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4238.2.8 http://zoobank.org/urn:lsid:zoobank.org:pub:3404BFA9-4297-4408-8DF2-03CB0A9E9CE3 Out of Amazonia: the unexpected trans-Andean distribution of Cochranella resplendens (Lynch and Duellman, 1978) (Anura: Centrolenidae) CLAUDIA MOLINA-ZULUAGA1, ESTEFANY CANO1, ADRIANA RESTREPO1, MARCO RADA2 & JUAN M. DAZA1,3 1Grupo Herpetológico de Antioquia, Instituto de Biología, Universidad de Antioquia, A.A. 1226, Medellín, Colombia 2Departamento de Zoologia, Instituto de Biociências; Universidade de São Paulo, Rua do Matão, trav. 14, 321, Cidade Universitária, 05508-090 São Paulo, Brazil 3Corresponding author. E-mail: [email protected] The glassfrog genus Cochranella, with nine recognized species, is distributed in the lowlands and mid elevation of the Neotropical forests, from Nicaragua to Bolivia (Guayasamin et al. 2009; Twomey et al. 2014). Four species are trans- Andean—C. granulosa (Taylor 1949) occurs in the lowlands and mountains, at mid elevation, of Central America, C. litoralis (Ruiz-Carranza & Lynch 1996) and C. mache Guayasamin & Bonaccorso 2004 occur in the Pacific lowlands and the western cloud forests of Colombia and Ecuador, and C. euknemos (Savage & Starrett 1967) occurs both in Central America and South America (northwestern Colombia).—The other five species have cis-Andean distributions in the Amazonian slopes and lowlands, from Colombia to Bolivia: C. nola Harvey 1996, C. guayasamini Twomey, Delia & Castroviejo-Fisher 2014, C. resplendens (Lynch & Duellman 1973), C. erminea Torres-Gastello, Suárez-Segovia & Cisneros-Heredia 2007, and C. phryxa Aguayo-Vedia & Harvey 2006. In Colombia, C. resplendens is known from the foothills of the Amazon versant in Caquetá (Malambo et al. 2013) and Putumayo (Lynch & Duellman 1973; Ruiz- Carranza et al. 1996). The species is also known from Ecuador (Lynch & Duellman 1973) and Peru (Twomey et al. 2014). Here, we report two new records of Cochranella resplendens, extending the species distribution beyond the Amazonian lowlands into the northern Cordillera Central in Colombia. During recent fieldwork on the eastern versant of the northern Cordillera Central in the Department of Antioquia, we found two specimens that we assigned to Cochranella resplendens. The first record corresponds to an adult female (MHUA-A 9140; Fig. 1A) collected on June 13, 2015 at Finca El Chaquiral, vereda El Retiro, municipality of Anorí, Department of Antioquia (6°58’56.06”N, 75°7’48.72” W, 1699 m). The specimen was found at night, on a bush leaf next to a 2 meter-wide stream inside a regrowth forest. The second record, an adult male (MHUA-A 9540; Fig. 1B) was collected on August 27, 2015 at El Eden, vereda San Antonio, municipality of Alejandría, department of Antioquia (6°22’2.1” N, 75°1’38.03” W, 1309 m). The individual was captured during the day inside a mature secondary forest after falling from a tree next to a stream. Both localities correspond to Premontane Perhumid Forest life zone (Holdridge 1987). These records extend northward the species known distribution to the trans-andean region approximately 618 km north from its nearest published locality in Belén de los Andaquíes, departament of Caquetá (Malambo et al. 2013; Fig. 2). Cochranella resplendes is a rare species in herpetological collections. Because the unusual distribution of the specimens reported herein (i.e., outside the Amazon basin), and the high similarity among some other species (i.e C. mache, C. euknemos and C. phryxa), we present more information regarding how the specimens were identified as C. resplendens. We used molecular and morphological evidence to allocate the two specimens to C. resplendens. We sequenced a mitochondrial fragment corresponding to the 16S region following the protocols described in Guayasamin et al. (2008b). Sequence alignment was conducted using MAFFT under default parameters (Katoh 2013). We inferred a 16S gene tree using Maximum Likelihood using the TIM2+G+I model (Figure 3). Model of nucleotide evolution and phylogeny were estimated using the program IQ-TREE (Nguyen et al. 2015). Nodal support was estimated using the ultrafast bootstrap method implemented in IQ-TREE (Minh et al., 2013). Included taxa are shown in Table 1. Using Mega 7 (Tamura et al. 2013) we estimated uncorrected genetic distances between species in Cochranella (Table 2). The phylogenetic hypothesis and the genetic distances show that the two specimens can be unambiguously assigned to C. resplendens. A remarkable finding is that the sample of C. resplendens from the Amazon versant in Ecuador and the ones from the Central Andes in Colombia show very little genetic variation (0.4 %; Table 2), despite being located in two different biogeographic areas (Fig. 2). 268 Accepted by S. Castroviejo-Fisher: 6 Jan. 2017; published: 3 Mar. 2017 TABLE 1. Taxa and Genbank accession numbers included in the phylogenetic analysis. Species Voucher Genbank accesion number Cochranella erminea CORBIDI 10477 KM068264 Cochranella erminea MHNC 13932 KM068258 Cochranella erminea MHNC 7247 KF534360 Cochranella erminea MNCN 45949 KM068257 Cochranella euknemos AJC 1687 KR863138 Cochranella euknemos AJC 1688 KR863137 Cochranella euknemos AJC 1850 KR863135 Cochranella euknemos CH5109 EU663008 Cochranella euknemos CH 6423 KR863136 Cochranella euknemos CH 6440 KR863139 Cochranella euknemos CH 6696 KR863141 Cochranella euknemos CH 6766 KR863134 Cochranella euknemos CH 6864 KR863140 Cochranella euknemos KRL 0803 FJ784377 Cochranella euknemos KRL 0841 FJ784396 Cochranella euknemos KRL 0937 FJ784443 Cochranella euknemos KRL 1054 FJ784458 Cochranella euknemos KRL 1055 FJ784459 Cochranella granulosa KRL 1011 FJ784455 Cochranella granulosa USNM 559082 EU663010 Cochranella guayasamini CORBIDI 8956 KM068265 Cochranella guayasamini MHNC 13930 KM068259 Cochranella litoralis QCAZ 27693 EU662990 Cochranella mache QCAZ 27747 EU663013 Cochranella nola CBG 814 EU663015 Cochranella nola CBG 1094 EU663016 Cochranella resplendens MHUAA 9140 KY347026 Cochranella resplendens MHUAA 9540 KY347027 Cochranella resplendens QCAZ 38088 KF534361 Cochranella sp. 1 AJC 2718 KF534362 Cochranella sp. CBG 1096 EU663021 Espadarana prosoblepon UCR 17102 EU662995 Both specimens referred to here as Cochranella resplendens exhibited the diagnostic characteristics described by Lynch & Duellman (1973): i) white parietal and visceral peritonea; ii) color in life green to dark green with white to bluish-green flecks; iii) round snout in dorsal view and gradually inclined anteroventrally in lateral view; iv) extensive webbing between outer fingers; v) dorsal skin shagreened with elevated warts corresponding to flecks; vi) crenulated dermal fold on the outer edge of hands, forearms, and elbows, on heels, tarsi, and feet; vii) pigmented nuptial excrescence, Type I; concealed prepollex; viii) humeral spines in adult males absent (crista ventralis not protruding); ix) cloacal ornamentation composed of enameled tubercules, folds, and crenulated flaps. Within the genus, the only species that lack humeral spines and have inclined snouts in lateral view are Cochranella euknemos (Savage & Starrett 1967), C. mache (Guayasamin & Bonaccorso 2004), and C. phryxa (Aguayo-Vedia & Harvey 2006). However, C. resplendens is distinguished by having well crenulated folds on hands, ulnae, tarsi and feet (Fig. 1A, C; less crenulated fold in C. euknemos; Fig. 1D), color in life green to dark green with white to bluish-green flecks (green or bluish-green dorsum with yellow flecks in C. euknemos and C. mache), well defined and visible tympanum (Fig. 1A, C; not visible externally in C. phryxa), and lacking a large dull yellow-colored patch on the top of the head (present in C. mache). COCHRANELLA RESPLENDENS Zootaxa 4238 (2) © 2017 Magnolia Press · 269 TABLE 2. Percentage of uncorrected genetic distances among Cochranella species based on DNA sequences of a 864 bp fragment of the mitochondrial gene 16S. Bold numbers in the diagonal indicate intraspecific genetic variation, interspecific genetic distances are shown below and above the diagonal, mean and range, respectively. Taxon sampling shown in Figure 3. 12345 1 erminea 2.8 0.4–5.5 3.5–5.0 3.7–4.4 5.9–6.2 2 euknemos 4.6 1.1 3.5–4.8 3.7–4.5 5.6–6.0 3 granulosa 4.3 4.3 0.2 3.3–4.3 4.5–5.2 4 guayasamini 4.1 4.0 3.6 0.0 5.1–5.6 5 litoralis 6.1 5.8 4.9 5.4 --- 6 mache 4.3 2.8 4.1 3.6 5.9 7 nola 6.1 5.2 5.4 5.1 5.5 8 resplendens 4.8 5.5 3.6 4.9 5.7 9 AJC2718 3.8 4.7 3.8 2.0 6.0 10 CBG1096 6.1 5.8 5.4 5.9 5.5 continued. 678910 1 erminea 4.1–4.5 5.6–6.6 4.4–5.2 3.5–4.0 5.4–6.5 2 euknemos 2.5–2.9 4.8–5.3 4.5–5.9 4.3–5.2 5.2–6.1 3 granulosa 3.8–4.4 4.8–6.0 3.2–4.1 3.2–4.4 5.3–5.5 4 guayasamini 3.5–3.8 4.8–5.5 4.7–5.4 1.8–2.2 5.9–5.9 5 litoralis --- 5.4–5.5 5.5–5.8 --- --- 6 mache --- 5.1–5.3 4.3–4.8 --- --- 7 nola 5.2 0.4 5.3–6.3 5.9–6.0 2.9–3.0 8 resplendens 4.5 5.7 0.4 4.8–5.5 5.8–6.3 9 AJC2718 4.6 5.9 5.1 --- --- 10 CBG1096 4.9 2.9 6.0 6.2 --- For decades, the frog fauna from the eastern side of the Andes has been considered very distinct from the trans- Andean region (Duellman 1979). Few examples of species inhabiting both sides of the Andes (mid to high elevations, including internal valleys in Colombia) are: Centrolene buckleyi (Ruiz-Carranza et al.