Notes on the taxonomy of some Glassfrogs from the Andes of Peru and Ecuador (Amphibia: Centrolenidae)

Notes on the taxonomy of some Glassfrogs from the Andes of Peru and Ecuador (Amphibia: Centrolenidae)

Diego F. Cisneros-Heredia^I,III; Juan M. Guayasamin^II

^IUniversidad San Francisco de Quito USFQ, Colegio de Ciencias Biológicas y Ambientales, Laboratorio de Zoología Terrestre, calle Diego de Robles y Ave. Interoceánica, Campus Cumbayá, edif. Darwin, DW010-A, Casilla Postal 17-1200-841, Quito, Ecuador. E-mail: [e-mail]

^IIUniversidad Tecnológica Indoamérica, Centro de Investigación de la Biodiversidad y el Cambio Climático, Av. Machala y Sabanilla, Quito, Ecuador

^IIIKing's College London, Department of Geography, London, England, United Kingdom

ABSTRACT

We present new information on several species of centrolenid frogs from Ecuador and Peru that justify the placement of Centrolene fernandoi Duellman and Schulte as a junior synonym of Centrolenella audax Lynch and Duellman; Centrolenella puyoensis Flores & McDiarmid as a synonym of Centrolenella mariae Duellman & Toft; and Cochranella tangarana Duellman & Schulte as a synonym of Cochranella saxiscandens Duellman & Schulte.

Key-words:Centrolene fernandoi; Centrolene audax; Nymphargus puyoensis; Nymphargus mariae; Rulyrana saxiscandens; Rulyrana tangarana; Rulyrana spiculata; Synonymy.

RESUMEN

Presentamos nueva información sobre algunas especies de ranas centrolénidas de Ecuador y Perú que justifica colocar a Centrolene fernandoi Duellman and Schulte como sinónimo junior de Centrolenella audax Lynch and Duellman, Centrolenella puyoensis Flores & McDiarmid como sinónimo de Centrolenella mariae Duellman & Toft, y Cochranella tangarana Duellman & Schulte como sinónimo de Cochranella saxiscandens Duellman & Schulte.

Palabras-clave:Centrolene fernandoi; Centrolene audax; Nymphargus puyoensis; Nymphargus mariae; Rulyrana saxiscandens; Rulyrana tangarana; Rulyrana spiculata; Sinonimia.

INTRODUCTION

Glassfrogs are conspicuous members of riverine communities across Neotropical America and have more than 140 described species (Frost, 2013). Twenty-nine species of glassfrogs have been reported from Peru (see below). Nymphargus ocellatus Boulenger was the first centrolenid species to be described from Peru (Boulenger, 1918), and no further glassfrogs were reported from the country until Duellman's (1976) description of Nymphargus truebae Duellman and Rulyrana spiculata Duellman, who also reported Nymphargus siren (Lynch & Duellman) and Hyalinobatrachium munozorum Lynch & Duellman. In 1979, Nymphargus mariae Duellman & Toft was described from the Serranía del Sira. Cannatella & Duellman (1982) re-evaluated the Peruvian specimens assigned to N. siren and regarded them as a different species: Nymphargus phenax (Cannatella & Duellman). They also described Nymphargus pluvialis (Cannatella & Duellman), and provided the first Peruvian records for Teratohyla midas (Lynch & Duellman) and Hyalinobatrachium bergeri (Cannatella). Centrolene azulae (Flores & McDiarmid) was described in subsequent years from an isolated mountain range on the eastern Andes of Peru, while Centrolene hesperium (Cadle & McDiarmid) and Cochranella euhystrix (Cadle & McDiarmid) were described from the Pacific slopes of northwestern Andean Peru (Flores & McDiarmid, 1989; Cadle & McDiarmid, 1990). Duellman & Wild (1993) provided the first country record of Centrolene buckleyi Boulenger. Duellman & Schulte (1993) almost doubled the number of Peruvian centrolenids with the description of eight species from the eastern slopes of Cordillera Central and adjacent ridges in the department of San Martín: Centrolene fernandoi Duellman & Schulte, Centrolene lemniscatum Duellman & Schulte, Centrolene muelleri Duellman & Schulte, Nymphargus chancas (Duellman & Schulte), Cochranella croceopodes Duellman & Schulte, Rulyrana saxiscandens (Duellman & Schulte), R. tangarana (Duellman & Schulte), and Hyalinobatrachium lemur Duellman & Schulte. Nymphargus mixomaculatus (Guayasamin, Lehr, Rodríguez & Aguilar) was described from central Andean Peru (Guayasamin et al., 2006). Torres-Gastello et al. (2007) described Rulyrana erminea Torres-Gastello, Suárez-Segovia & Cisneros-Heredia, and reported the first records of Cochranella resplendens (Lynch & Duellman) and Vitreorana oyampiensis Lescure from Amazonian Peru. Cisneros-Heredia et al. (2008) described Rulyrana mcdiarmidi from Ecuador and Peru, and presented the first record of Nymphargus posadae from Peru. Yánez-Muñoz et al. (2009) reported the first Peruvian record of Hyalinobatrachium iaspidiense Ayarzagüena from Amazonian Peru. Castroviejo-Fisher et al. (2009) described Hyalinobatrachium carlesvilai Castroviejo-Fisher, Padial, Chaparro, Aguayo & de la Riva from the Amazonian slopes of central Andean Peru, assigned all previous records of H. munozorum from Peru either to H. carlesvilai or to H. bergeri, synonymized H. lemur with H. pellucidum Lynch & Duellman, and extended the distribution of the latter south to the department of Cusco, southern Peru. Catenazzi et al. (2012) described Centrolene sabini Catenazzi, von May, Lehr, Gagliardi-Urrutia & Guayasamin from the Amazonian slopes of southeastern Andean Peru. Catenazzi & Venegas (2012) presented photographs of Chimerella mariaelenae (Cisneros-Heredia & McDiarmid) from Kampankis, in the Amazonian slopes of northeastern Peru.

While developing our extensive reviews of Centrolenidae (Cisneros-Heredia & McDiarmid, 2007; Guayasamin et al., 2009), we cooperatively found that there is no evidence to support specific recognition of several populations of Peruvian and Ecuadorian centrolenids currently hypothesised as different species. Herein, we present our findings in an effort to enhance the understanding on the diversity and conservation of Neotropical amphibians.

MATERIALS AND METHODS

Characters and terminology are standardized following the definitions provided by Cisneros-Heredia & McDiarmid (2007). Taxonomy and systematics follow Guayasamin et al. (2009). The following measurements (in millimetres) were taken with electronic digital callipers (0.05 mm accuracy, rounded to the nearest 0.1 mm): snout-vent length, SVL; head width, HW; head length, HL; horizontal eye diameter, ED; inter-orbital distance, IOD; eye-nostril distance, EN; inter-narial distance, IN; width of disc on the third finger, 3DW; tibia length, TL; foot length, FL. Upper eyelid width was not measured because of its limited utility due to preservation bias. We use the notational device for webbing formulae of Savage & Heyer (1967), as modified by Savage & Heyer (1997). Sex and sexual maturity was determined by direct examination of the condition of gonads and development of secondary sexual characters (vocal slits and nuptial pads). We examined specimens (^{Appendix I}) deposited in the following collections: DHMECN – División de Herpetología, Museo Ecuatoriano de Ciencias Naturales, Quito; DFCH-USFQ – Universidad San Francisco de Quito, Quito; QCAZ – Museo de Zoología, Pontificia Universidad Católica del Ecuador, Quito; BMNH – Natural History Museum, London; KU – The University of Kansas, Natural History Museum, Lawrence; USNM – National Museum of Natural History, Washington, D.C.; MCZ – Museum of Comparative Zoology, Harvard University.

RESULTS AND CONCLUSIONS

Centrolene audax Lynch & Duellman, 1973

(Fig. 1)

Centrolenella audax Lynch & Duellman, 1973.

Centrolene audax – Ruiz-Carranza & Lynch, 1991.

"Centrolene" audax – Guayasamin et al., 2009.

Centrolene fernandoi – Duellman & Schulte, 1993. Holotype: KU 211770. Type locality: west slope of Abra Tangarana, 7 km (by road) northeast of San Juan de Pacaysapa (06º12'S, 76º44'W, 1080 m), Provincia Lamas, Departamento San Martín, Perú. New synonymy.

Lynch & Duellman (1973) described Centrolenella audax for glassfrog populations diagnosed as having small yellow spots on the dorsum, short and distally-curved humeral spines in males, and extensive webbing between outer fingers from the Amazonian versant of the northern Andes. Centrolene audax is currently known in Colombia and Ecuador from few localities in Low Montane Evergreen Forest on the Amazonian versant of the Andes, between 1350 and 1800 m (Mueses-Cisneros, 2005; Cisneros-Heredia & McDiarmid, 2007; Yánez-Muñoz et al., 2010).

Centrolene fernandoi Duellman & Schulte (1993) was described based on nine specimens collected on the western slope of Abra Tangarana, Amazonian versant of the Andes of Peru. Duellman & Schulte (1993) compared C. fernandoi with Centrolene audax and reported its high similarity, but differentiated them by its snout form (bluntly rounded in C. fernandoi, truncate in C. audax), dorsal skin texture (with scattered small spicules in C. fernandoi, without spicules in C. audax), dorsal fleck colouration (bluish-white in C. fernandoi, golden in C. audax), finger colouration (pale green in C. fernandoi, pale yellow in C. audax), and iris background colouration (silvery green in C. fernandoi, pale bronze in C. audax). Centrolene fernandoi remains known only from its type locality (Frost, 2011).

We examined six specimens of Centrolene fernandoi (type-series) and 42 specimens of Centrolene audax (including the type-series). We found that all differences used to separate them are intraspecifically variable within C. audax. Snout form of C. audax varies between rounded to truncate in profile, presence of spicules shows sexual and ontogenic variation (visible in reproductive males, and absent in non-reproductive males and females), dorsal flecks vary from pale yellow to golden yellow (furthermore, the photograph of C. fernandoi in the original description shows pale yellow spots), finger colouration varies from pale green to bright yellow, and iris colouration varies from pale bronze to silvery green or mustard with thin black reticulation. Since no discrete differences are evident and their populations have no obvious biogeographic barriers, we place Centrolene fernandoi Duellman & Schulte, 1993 as a junior synonym of Centrolenella audax Lynch & Duellman, 1973. Therefore, Centrolene audax inhabits Low Montane Evergreen Forest on the Amazonian versant of the Andes of southern Colombia, Ecuador, and northern Peru, between 1080 and 1800 m (Duellman & Schulte, 1993; Mueses-Cisneros, 2005; Cisneros-Heredia & McDiarmid, 2007; Yánez-Muñoz et al., 2010).

Nymphargus mariae (Duellman & Toft, 1979)

(Fig. 2)

Centrolenella mariae Duellman & Toft, 1979.

Centrolenella puyoensis – Flores & McDiarmid, 1989. Holotype: MCZ 91187, by original designation. Type locality: "1.0 km W Puyo, Provincia de Pastaza, Ecuador, between 1000-1050 m elevation". New synonymy.

Cochranella mariae – Ruiz-Carranza & Lynch, 1991. Cisneros-Heredia & McDiarmid, 2007.

Cochranella puyoensis – Ruiz-Carranza & Lynch, 1991. Cisneros-Heredia & McDiarmid, 2006.

Centrolene mariae – Duellman & Schulte, 1993.

Centrolene puyoensis – Duellman & Schulte, 1993.

Centrolene puyoense – Stuart et al., 2008.

Nymphargus mariae – Guayasamin et al., 2009.

Nymphargus puyoensis – Guayasamin et al., 2009.

Centrolenella mariae Duellman & Toft was described based on one female specimen collected at Serranía de Sira, department of Huánuco, Peru (Duellman & Toft, 1979). Flores & McDiarmid (1989) described Centrolenella puyoensis and Centrolenella azulae, hypothesising that, together with C. mariae, they formed a monophyletic group (the C. mariae species-group). Subsequent authors (Ruiz-Carranza & Lynch, 1991, 1995; Duellman & Schulte, 1993) followed this hypothesis, but Cisneros-Heredia & McDiarmid (2006, 2007) questioned the validity of the C. mariae species-group, further pointing out that although C. azulae is diagnosable, C. mariae and C. puyoensis are very similar and probably conspecific (Cisneros-Heredia & McDiarmid, 2007). Centrolenella mariae and C. puyoensis were placed in the genus Nymphargus by Guayasamin et al. (2009) based on morphological and molecular data, respectively. Flores & McDiarmid (1989) separated N. puyoensis from N. mariae by tympanum exposure (three-quarters in N. puyoensis, one-half in N. mariae), hand and foot webbing (slightly more extensive in N. mariae), ulnar fold (present in N. puyoensis, absent in N. mariae), intricate cloacal ornamentation (present in N. puyoensis, absent in N. mariae) and differences in proportions (greater eye-nostril/eye diameter and shank length/snout-vent length in N. puyoensis).

We examined 16 specimens assignable to Nymphargus puyoensis from Ecuador and the holotype of Nymphargus mariae and found no evidence to support their differentiation. All characters used to diagnose these two species can be attributed to subtle differences that fall inside the intraspecific variation of a single species. Flores & McDiarmid (1989) proposed a polarization of characters that is rather subjective and biased due to their small sample size (one specimen for each of their "species"). Variation of tympanum exposure, webbing, and body proportions observed among N. puyoensis and N. mariae is continuous and similar, or even lower than the natural differences observed within populations of other species of centrolenids. The absence of ulnar folds and cloacal ornamentations in the type specimen of N. mariae could be attributed to natural variation, but also due to preservation artifacts (see Cisneros-Heredia & McDiarmid, 2006 for information on the variation of specimens of N. puyoensis). Either way it has also been observed in specimens of N. puyoensis.

In the absence of valid discriminating evidence to support the hypothesis that Nymphargus puyoensis and Nymphargus mariae are different lineages, we place Centrolenella puyoensis Flores & McDiarmid, 1989 as a synonym of Centrolenella mariae Duellman & Toft, 1979. Thus, Nymphargus mariae, as herein redefined, inhabits Foothill Evergreen Forest and Lowland Evergreen Forest flooded by White-water Rivers on the Amazonian versant of the Andes of Ecuador and Peru (Cordillera del Sira), between 300 and 1550 m (Flores & McDiarmid, 1989; Cisneros-Heredia & McDiarmid, 2006, 2007; Yánez-Muñoz et al., 2010).

This synonym reflects the absence of evidence to support the hypothesis that the population from the Serranía del Sira in eastern Amazonian Peru (type-locality of Nymphargus mariae) is different from those of eastern Amazonian Ecuador. Although it might be argued that the Serranía del Sira is a rather isolated mountain range that likely contains several amphibian endemics, our decision to place Nymphargus puyoensis in the synonymy of N. mariae is based on the fact that there are no morphological traits that support the existence of two putative species, and that potential biogeographic barriers cannot justify specific status without the corroboration of traits intrinsic to the organisms. We encourage future researchers to analyse other lines of evidence to evaluate the status of these populations.

Rulyrana saxiscandens (Duellman & Schulte, 1993)

(Fig. 3)

Cochranella saxiscandens Duellman & Schulte, 1993.

Cochranella tangarana Duellman & Schulte, 1993. Type locality: "west slope of Abra Tangarana, 7 km (by road) northeast of San Juan de Pacaysapa (06º12'S, 76º44'W), 1080 m), Provincia Lamas, Departamento San Martín, Perú". New synonymy.

Rulyrana saxiscandens – Guayasamin et al., 2009.

Rulyrana tangarana – Guayasamin et al., 2009.

Duellman & Schulte (1993) described Cochranella saxiscandens and Cochranella tangarana based on specimens collected at two nearby localities of the Mayo River, Tarapoto region, department of San Martín, Peru. Duellman & Schulte (1993) differentiated these two species (now placed in the genus Rulyrana) by their snout form (bluntly round in Rulyrana saxiscandens, truncate in Rulyrana tangarana), dorsal colouration in preservative (dark-grey to black in R. saxiscandens, lavender in R. tangarana), melanophores on the ventral surfaces of shanks and tarsi (present in R. saxiscandens, absent in R. tangarana); presence of spicules on dorsal surfaces (absent in R. saxiscandens, present in R. tangarana); and inner tarsal fold (absent in R. saxiscandens, present in R. tangarana).

We examined 22 specimens of Rulyrana saxiscandens and two of Rulyrana tangarana (including all type-specimens) and found that all stated differences between them actually correspond to intraspecific variation. Snout shape varies continuously from round to truncate; tympanic annulus is conspicuous at different degrees due to the supratympanic fold; dorsal colouration in preservative varies continuously from dark-purple, purplish-grey, dark-lavender, to light-lavender (similar colour variation has been observed in Rulyrana flavopunctata); melanophores are always present on ventral surfaces although sometimes scarce; spicule presence and appearance varies ontogenically and sexually (see Cisneros-Heredia & McDiarmid, 2007); and inner tarsal fold is always present but sometimes poorly noticeable.

In the absence of evidence to support the hypothesis that two species are involved in the populations of the Mayo River, we place Cochranella tangarana Duellman & Schulte, 1993 as synonym of Cochranella saxiscandens Duellman & Schulte, 1993.

Rulyrana saxiscandens remains very similar to Rulyrana spiculata (Duellman, 1976), which is known from forests on the Amazonian versant of the Andes of central and southern Peru and eastern Bolivia, between 1200 and 1700 m (Frost, 2011; Rodríguez et al., 2004). Duellman & Schulte (1993) reported that Centronella saxiscandens (and Centrolenella tangarana) were similar to Centrolenella spiculata, but differed due to snout form, tympanum and inner tarsal fold appearance, coloration, and presence of spicules on dorsal surfaces in Rulyrana tangarana. Evan Twomey and associates are studying these species, and we refer to them for a definitive conclusion.

With the present changes, the diversity of glassfrogs of Peru currently includes 29 species: Centrolene audax, C. azulae, C. buckleyi, C. hesperium, C. lemniscatum, C. muelleri, C. sabini, Chimerella mariaelenae, Cochranella croceopodes, C. euhystrix, C. resplendens, Hyalinobatrachium bergeri, H. carlesvilai, H. iaspidiense, H. pellucidum, Rulyrana erminea, R. mcdiarmidi, R. saxiscandens, R. spiculata, Vitreorana oyampiensis, Nymphargus chancas, N. mariae, N. mixomaculatus, N. ocellatus, N. phenax, N. pluvialis, N. posadae, Teratohyla amelie, T. midas.

Hyalinobatrachium munozorum and Centrolene condor are expected to occur in Peru. Hyalinobatrachium munozorum occurs in Ecuador and Bolivia (Cisneros-Heredia & McDiarmid 2007, Castroviejo-Fisher et al., 2011), and C. condor is known from several localities in the Cordillera del Condor, just a few kilometres from Peruvian territory (Cisneros-Heredia & Morales-Mite, 2008; Almendáriz & Batallas, 2012).

ACKNOWLEDGMENTS

Funding for the development of diverse stages of this study was obtained from the following: Russell E. Train Education for Nature Program/WWF, Conservation International, Smithsonian Women's Committee, Research Training Program of the National Museum of Natural History – Smithsonian Institution, María Elena Heredia, Laura Heredia, and Universidad San Francisco de Quito to DFCH; and from the project "Patrones de diversidad de los anfibios andinos del Ecuador" of the Universidad Tecnológica Indoamérica to JMG. We are thankful to Linda Trueb (KU), Mario Yánez-Muñoz (DHMECN), Roy W. McDiarmid, W. Ron Heyer and George R. Zug (USNM), David Gower and Mark Wilkinson (BMNH), and James Hanken and José Rosado (MCZ) for granting access to specimens under their care. We are grateful to Santiago Castroviejo, Marco Rada, and Evan Twomey for discussions on the taxonomy of Peruvian centrolenids.

Aceito em: 27/08/2013

Publicado em: 30/06/2014

Examined specimens

Centrolene audax: ECUADOR: Napo: KU 146624 (holotype of Centrolenella audax): Salto de Agua; KU 155502-03: 7 km SW of Río Azuela; KU 164496: Azuela; KU 164497-504: 2 km SSW of Río Reventador; USNM 286620-22: Cascada de San Rafael; KU 178018-27: Río Salado; USNM 286623-25, MCZ A97807-8: 14.6 km (by road) NE of Río Salado; KU 190015: 43 km NE of Santa Rosa; KU 190016: 8.9 km NE Santa Rosa; KU 143290, 143292: 16,5 km NNE Santa Rosa; DHMECN 06788-89: Reserva Biológica Narupa. PERU: San Martín: KU 211770 (holotype of Centrolene fernandoi), 211771-5: W slope Abra Tangarana.

Nymphargus mariae: ECUADOR: Napo: DFCH-USFQ D285: ca. 45 km E of Narupa. PASTAZA: MCZ 91187 (holotype of Centrolenella puyoensis): 1.0 km W Puyo; USNM 291298: Río Pucayacu. QCAZ 37932: stream tributary of Río Lliquino; QCAZ 39293: near Villano; DHMECN 04752-53, 04756: Conambo; Orellana: QCAZ 7104, 7499: Río Huataracu; Sucumbíos: DHMECN 06190: Río Verde. PERU: Huanuco: KU 174713 (holotype of Centrolenella mariae): Serranía de Sira.

Rulyrana saxiscandens: PERU: KU 211776 (holotype of Cochranella tangarana), 211777, 217299: W slope of Abra Tangarana; KU 211779 (holotype of Cochranella saxiscandens), 211780-88, 211789-98, 211800-01: Cataratas Ahaushiyacu; KU 211802-03: 15 km NE of Tarapoto.

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APPENDIX I 

Publication Dates

• Publication in this collection
  02 July 2014
• Date of issue
  2014

History

• Received
  27 Aug 2013
• Accepted
  30 June 2014