Accepted by S. Carranza: 19 May 2006; published: 3 Aug. 2006 49[EN] We report new distributional information for Eleutherodactylus mercedesae in Bolivia, and provide the first record for Peru based on an adult female. This species, previously endemic to Bolivia, now ranges across about 1000 km in cloud forests on the Amazonian slopes of the Andes from southern Peru to central Bolivia. We provide the first morphological description of females based on two specimens, compare them with the male type and paratype, add some observations to the original description, and comment on variation in the species.[ES] Aportamos información novedosa sobre la distribución de Eleutherodactylus mercedesae en Bolivia y la primera cita para Perú, basada en una hembra adulta. Esta especie se consideraba hasta ahora endémica para Bolivia. Su rango conocido de distribución comprende actualmente unos 1000 km de los bosques nublados de las laderas amazónicas de los Andes, desde el sur de Perú al centro de Bolivia. Se describe por primera vez la hembra de esta especie en base a dos ejemplares, que son comparados con el holotipo y el paratipo. Añadimos algunas observaciones sobre la descripción original y la variación morfológica de la especie.This work was partially funded by projects REN/GLO 2001-1046 and CGL2005-03156 of the Spanish Ministry of Education and Science (I. De la Riva, Principal Investigator).Peer reviewe

De la Riva, Ignacio

McDiarmid, Roy

Padial, Manuel M.

Zootaxa

Padial, José M., Roy, Riva, Ignacio De La (2006): Distribution and morphological variation of Eleutherodactylus mercedesae Lynch & McDiarmid, 1987 (Amphibia, Anura, Leptodactylidae) with first record for Peru. Zootaxa 1278: 49-56, DOI: 10.5281/zenodo.27351

Padial, José M.

Riva, Ignacio De La

NEUROSURGERY ENTHUSIASTIC WOMEN SOCIETY

Distribution and morphological variation of Eleutherodactylus mercedesae Lynch & McDiarmid, 1987 (Amphibia, Anura, Leptodactylidae) with first record for Peru

FIGURE 2. Map of Southern Peru and Northern­Central Bolivia showing the collecting sites of Eleutherodactylus mercedesae. Squares indicate the localities mentioned in the original description.Published as part of Padial, José M., Roy & Riva, Ignacio De La, 2006, Distribution and morphological variation of Eleutherodactylus mercedesae Lynch & McDiarmid, 1987 (Amphibia, Anura, Leptodactylidae) with first record for Peru, pp. 49-56 in Zootaxa 1278 on page 52, DOI: 10.5281/zenodo.27351

FIGURE 2 in Distribution and morphological variation of Eleutherodactylus mercedesae Lynch & McDiarmid, 1987 (Amphibia, Anura, Leptodactylidae) with first record for Peru

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1278Accepted by S. Carranza: 19 May 2006; published: 3 Aug. 2006 49ZOOTAXAISSN 1175-5326  (print edition)ISSN 1175-5334 (online edition)Copyright © 2006  Magnolia PressZootaxa 1278: 49–56  (2006) www.mapress.com/zootaxa/Distribution and morphological variation of Eleutherodactylus mercedesae Lynch & McDiarmid, 1987 (Amphibia, Anura, Leptodactylidae) with first record for PeruJOSÉ M. PADIAL1, ROY McDIARMID2 & IGNACIO DE LA RIVA1*1Department of Biodiversity and Evolutionary Biology, Museo Nacional de Ciencias Naturales, CSIC. C/ José Gutiérrez Abascal 2, 28006 Madrid, Spain2USGS Patuxent Wildlife Research Center. Smithsonian Institution, National Museum of Natural History, PO Box 37012, Washington, DC 20013-7012 USA*Corresponding author. E-mail: iriva@mncn.csic.esAbstractWe report new distributional information for Eleutherodactylus mercedesae in Bolivia, and providethe first record for Peru based on an adult female. This species, previously endemic to Bolivia, nowranges across about 1000 km in cloud forests on the Amazonian slopes of the Andes from southernPeru to central Bolivia. We provide the first morphological description of females based on twospecimens, compare them with the male type and paratype, add some observations to the originaldescription, and comment on variation in the species.Key words: Eleutherodactylus mercedesae, Anura, Leptodactylidae, Bolivia, Peru, distribution,variationResumenAportamos información novedosa sobre la distribución de Eleutherodactylus mercedesae enBolivia y la primera cita para Perú, basada en una hembra adulta. Esta especie se consideraba hastaahora endémica para Bolivia. Su rango conocido de distribución comprende actualmente unos 1000km de los bosques nublados de las laderas amazónicas de los Andes, desde el sur de Perú al centrode Bolivia. Se describe por primera vez la hembra de esta especie en base a dos ejemplares, que soncomparados con el holotipo y el paratipo. Añadimos algunas observaciones sobre la descripciónoriginal y la variación morfológica de la especie.PADIAL ET AL.50                                       © 2006 Magnolia Press1278ZOOTAXA IntroductionEleutherodactylus mercedesae Lynch & McDiarmid, 1987 is an uncommon speciesinhabiting the Yungas and Humid Montane Forests of Central Bolivia (Lynch &McDiarmid 1987; De la Riva 1990). Until recently, it was exclusively known from twoclose localities (De la Riva 1993), but additional findings increased its known latitudinalrange, first around the type locality, and subsequently further to the northwest, inDepartment La Paz, Bolivia (De la Riva et al. 2000; Köhler 2000). Thus, our informationhas shifted from considering this species to be restricted to the surroundings of the typelocality to consider it a rare species with a distribution of 300 km wide in the AndeanCloud Forests of Bolivia (De la Riva et al. 2000). Despite recent intense fieldwork (Padialet al. 2004), including surveys of the type locality, few specimens have been collected andour knowledge of this species is still rudimentary. Nevertheless, recent collectionsexamined (CBG and USNM) by us yielded new distributional information of this speciesfrom Bolivia and a record from Peru, where it was previously unrecorded (Lehr 2002;Frost 2004; AmphibiaWeb 2006). Eleutherodactylus mercedesae was described from two adult males. Since then, theonly additional information on the species’ morphology was based on juveniles (Köhler2000). Females of Eleutherodactylus mercedesae were unknown (Lynch and McDiarmid1987). Since      Eleutherodactylus is a large and complex frog genus, with most species inthe same species group diagnosed only by subtle differences, some inter-populationalvariation could be misinterpreted as specific diagnostic characters if several specimens arenot studied. Moreover, with scant distributional data and with long distances betweencollecting sites, this variation could lead to erroneous recognition of populations asdifferent species. Therefore, we think it is important to provide any new morphologicaland distributional data.MethodsMorphological characters studied mainly follow Lynch and Duellman (1997) although,following Padial & De la Riva (2005) and Padial et al. (2005), abbreviations and somemeasurements described below differ from those used by the above-mentioned authors.Measurements and their respective abbreviations are described as follows: snout-ventlength, SVL; head length (from posterior margin of the lower jaw to tip of snout), HL;head width (measured at level of rictus), HW; eye length, EL; eye to nostril distance, EN;inter-narial distance, IND; eye to eye distance (distance between the anterior margins ofeyes), EE; tympanic membrane height, TYH; tympanic membrane length, TYL; width ofterminal disk of third finger, FIII; width of terminal disk of fourth finger, FIV; forearmlength (from elbow to the proximal margin of thenar tubercle), ARM; tibia length, TL;thigh length, TH; foot length (from proximal border of inner metatarsal tubercle to tip of © 2006 Magnolia Press                                                               51ELEUTHERODACTYLUS1278ZOOTAXAfourth toe), FL; width of terminal disk of fourth toe, TIV. We do not include values ofinterorbital distance (IOD) and upper eyelid width (EW). Our experience indicates thatthese parameters are usually of scarce utility because the preservation condition of aspecimen highly influences the measurements and makes it difficult to have precise andcomparable values. Specimens were measured by JMP with a digital calliper to the nearest0.01 mm. To avoid pseudo precision we rounded all measurements following Hayek et al.(2001). Specimens studied are deposited in the following collections: Colección Bolivianade Fauna, La Paz, Bolivia (CBF); Museo de Historia Natural de Cochabamba,Cochabamba, Bolivia (MHNC); National Museum of Natural History, Washington DC.USA (USNM); Zoologisches Forschungsinstitut und Museum Alexander Koenig, Bonn,Germany (ZFMK). All specimens examined are listed in the Appendix.ResultsA single adult female (USNM-346140; Figure 1) collected by A. Salas in a cloud forest at1700 m in Department Cusco, on 22 September 1991, represents the first and only recordfor Peru (see appendix). New records in Bolivia are: an adult male from SerraníaBellavista in Department La Paz (CBF-3701); a juvenile from Cotapata National Park,also in Department La Paz (CBF-4120); and an adult female from Pampa Grande,Altamachi, Department Cochabamba (MHNC-AMS-196). Thus, the known distribution ofE. mercedesae comprises a narrow band of Montane Forest from 1400–1950 m altitude,and about 300 km in length in Bolivia, and a single record in Peru (1700 m), separated bya distance of 480 airline km from the closest Bolivian records (see Figure 2). The Peruvianrecord, together with recent taxonomic discoveries (e.g. Lehr et al. 2004), clearly showshow incomplete our knowledge on the diversity of Eleutherodactylus in Peru is.FIGURE 1. Dorsal and ventral views of adult female Eleutherodactylus mercedesae (USNM-346140) from Peru (SVL =62.1).PADIAL ET AL.52                                       © 2006 Magnolia Press1278ZOOTAXAFIGURE 2. Map of Southern Peru and Northern-Central Bolivia showing the collecting sites ofEleutherodactylus mercedesae. Squares indicate the localities mentioned in the original description.We herein provide information on some adult characteristics not mentioned for thetypes and a more detailed morphometric account (Tables 1 and 2). The description ofjuvenile Eleutherodactylus mercedesae (ZFMK-72571-3, 72597-99) by Köhler (2000) isadequate and coincident with our observations; thus, we do not describe juveniles herein.Coloration in alcohol. Adult females are very similar to males (both types and CBF-3701) in coloration and pattern (Figure 1). The dorsum is greenish-grey to brownish-greywith cream irregular marks; the belly and throat show a cream background with moderateto intense brown mottling. All specimens have an inverted “V” or a rhomboidal creammark on brown background on the throat (as the holotype and some specimens ofEleutherodactylus rhabdolaemus). Another characteristic is a butterfly-like cream mark onthe scapular region (also observed in the holotype). Some black bold marks are present inboth sexes, in the anterior margin of flanks, scapular region, interocular region, lorealregion, subocular region, supratympanic fold, and dorsal surfaces of forearm.Skin texture. Although in the original description the dorsal skin was described asfinely granular, some specimens have a smooth skin (e.g. MHNC-AMS-196). The numberof warts and granules on dorsum, head and flanks is also variable. For example, eyelidtubercles and occipital warts are absent in MHNC-AMS-196 and some of these structureshave almost disappeared in the types. Dorsolateral and lateral folds are only present in theparatype.Sexual characters. USNM-346140 (Figure 1) is a gravid female with unpigmentedeggs, grouped in five round and concatenated packets in each oviduct. This morphology of © 2006 Magnolia Press                                                               53ELEUTHERODACTYLUS1278ZOOTAXAoviducts has been not previously observed in any of the other Bolivian species (pers. obs.).Males have white testis, as all other Bolivian species (pers. obs.). Lynch & McDiarmid(1987) diagnosed the absence of nuptial pads in males of E. mercedesae, but a re-examination revealed a single, faint, low, white glandular nuptial pad on the dorsal surfaceof the thumb, more evident in the paratype (USNM-165753) than in the holotype (USNM-257848) and the specimen CBF-3701. The sexual dimorphism in size is evident, beingfemales larger than males, although there are not differences in proportions (Tables 1 and2).TABLE 1. Measurements (in mm) and proportions of adult specimens of Eleutherodactylusmercedesae.Males FemalesUSNM257848 (Holotype)USNM165753 (Paratype)CBF3701 USNM346140 MHNC-AMS-196SVL 41.7 52.2 51.5 62.1 63.3HL 16.1 19.8 18.2 24.6 24.5HW 16.5 18.2 18.2 26.1 24.6EL 5.1 6.2 5.5 7.0 5.9EN 4.9 6.5 6.4 7.5 6.4IND 4.0 4.7 4.4 5.7 5.5EE 7.4 8.4 8.0 10.8 10.3TYH 2.3 2.9 2.5 4.0 3.6TYL 1.9 2.6 2.7 3.4 3.3FIII 2.2 2.9 2.3 3.4 2.8FIV 2.3 2.9 2.1 3.6 2.8ARM 9.4 12.0 12.6 15.7 15.1TL 28.0 35.4 35.5 43.6 42.6TH 24.8 30.8 31.7 40.8 38.7FL 23.5 29.3 29.4 38.9 36.5TIV 1.8 2.2 1.8 2.6 1.9TL/SVL 0.7 0.7 0.7 0.7 0.7FL/SVL 0.6 0.6 0.6 0.6 0.6HL/SVL 0.4 0.4 0.4 0.4 0.4HW/SVL 0.4 0.3 0.4 0.4 0.4HW/HL 1.0 0.9 1.0 1. 1 1.0EN/EL 1.0 1.0 1.2 1.1 1.1TYL/TYH 0.8 0.9 1.1 0.9 0.9PADIAL ET AL.54                                       © 2006 Magnolia Press1278ZOOTAXA TABLE 2. Means and ranges of morphometric proportions of adult specimens of Eleutherodactylusmercedesae.Remarks. In most Eleutherodactylus from Bolivia, males have more rounded tympanicmembranes and females more elliptical ones (JMP pers. obs.). In E. mercedesae, bothsexes have an elliptical tympanic membrane, being the paratype the only exception. Fingerlengths (3>4>1>2), toe lengths (4>3>5>2>1), and disc shapes are similar in males andfemales. In the female USNM-346140, low occipital “W” shaped crests are also evident,what is exclusive of this specimen. The tarsal fold is absent in MHNC-AMS-196. Fingerand toe fringes are present in different degrees in all adult specimens. Development ofsupernumerary tubercles in the hand is also variable, being big and prominent in theholotype and less prominent in MHNC-AMS-196, which could be an artifact ofpreservation.No relevant differences were found among males and females, besides from the usualsexual dimorphism present in other Eleutherodactylus. More relevant aspects are that bothmales and females are identical in the proportions (see Tables 1 and 2) and no remarkabledifferences were found among specimens from different localities. The observed discontinuities in the distribution of E. mercedesae are surely due to thelack of adequate sampling.AcknowledgementsJMP’s work in Bolivia was financed by a grant from the Mutis program of the MAE-AECI(Spain), and his visits to US museums were funded by an Ernst Mayr Travel Grant inAnimal Systematics (Museum of Comparative Zoology, Harvard University). Specialthanks to A. Muñoz and R. Aguayo (CBG) for allowing JMP the study of their specimens.JMP and IDlR are grateful to the following persons for the help and space provided at theirrespective institutions: Dr. M. Suárez, A. Justiniano, R. Vespa, R. Montaño, and L.González (Museo de Historia Natural Noel Kempff Mercado, Santa Cruz, Bolivia), J.Aparicio (Colección Boliviana de Fauna, La Paz), J. Rosado (MCZ), W. R. HeyerProportions Males (n = 3) Females (n = 2)TL/SVL 0.7 (0.7–0.7) 0.7 (0.7–0.7)FL/SVL 0.6 (0.6–0.6) 0.6 (0.6–0.6)HL/SVL 0.4 (0.4–0.4) 0.4 (0.4–0.4)HW/SVL 0.4 (0.3–0.4) 0.4 (0.4–0.4)HW/HL 1.0 (0.9–1.0) 1.0 (1.0–1.1)EN/EL 1.1 (1.0–1.2) 1.1 (1.1–1.1)TYL/TYH 0.9 (0.8–1.1) 0.9 (0.9–0.9) © 2006 Magnolia Press                                                               55ELEUTHERODACTYLUS1278ZOOTAXA(USNM), and W. Böhme and J. Köhler (ZFMK). This work was partially funded byprojects REN/GLO 2001-1046 and CGL2005-03156 of the Spanish Ministry of Educationand Science (I. De la Riva, Principal Investigator). We are grateful to Fernando Palaciosfor his decisive help.Literature citedAmphibiaWeb. (2006) Available from: http://elib.cs.berkeley.edu/aw/ (Date of access 24 January2006).De la Riva, I. (1990) Lista preliminar comentada de los anfibios de Bolivia con datos sobre su dis-tribución. Bollettino del Museo regionale di Scienze Naturali Torino, 8, 261–319.De la Riva, I. (1993) Sinopsis del género Eleutherodactylus (Amphibia, Anura, Leptodactylidae) enBolivia y adición de tres especies nuevas para el país. Revista Española de Herpetología, 7,97–105.De la Riva, I., Köhler, J., Lötters, S. & Reichle, S. (2000) Ten years of research on Bolivianamphibians: updated checklist, distribution, taxonomic problems, literature and iconography.Revista Española de Herpetología, 14, 19–164.Frost, D.R. (2004) Amphibian Species of the World: an online reference. Available from: http://research.amnh.org/herpetology/amphibia/index.html (Date of access 24 January 2006).Hayek, L.A., Heyer, W.R. & Gascon, C. (2001) Frog morphometrics: a cautionary tale. Alytes, 18,153–177.Köhler, J. (2000) Amphibian diversity in Bolivia: a study with special reference to montane forestregions. Bonner zoologische Monographien, 48, 1–244.Lehr, E., (2002) Amphibien und Reptilien in Peru. Die Herpetofauna entlang des 10. Breitengradesvon Peru: Arterfassung, Taxonomie, ökologische Bemerkungen und biogeographische Bezie-hungen. Natur und Tier Verlag (NTV Wissenschaft), Münster, 208 pp.Lehr, E., Aguilar, C. & Duellman, W.E. (2004) A striking new species of Eleutherodactylus fromAndean Peru (Anura: Leptodactylidae). Herpetologica, 2, 275–280.Lynch, J.D. & McDiarmid, R.W. (1987) Two new species of Eleutherodactylus (Amphibia: Anura:Leptodactylidae) from Bolivia. Proceedings of the Biological Society of Washington, 100,337–346.Lynch, J.D. & Duellman, W.E. (1997) Frogs of the genus Eleutherodactylus in Western Ecuador:systematics, ecology, and biogeography. University of Kansas Natural History Museum Spe-cial Publications, 23, 1–236.Padial, J.M., Gonzáles, L., Reichle, S., Aguayo, R. & De la Riva, I. (2004) First records of five spe-cies of the genus Eleutherodactylus (Anura: Leptodactylidae) for Bolivia. Graellsia, 60,167–174.Padial, J.M. & De la Riva, I. (2005) Rediscovery, redescription and advertisement call of Eleuth-erodactylus heterodactylus (Miranda Ribeiro, 1937) (Anura: Leptodactylidae), and notes onother Eleutherodactylus. Journal of Herpetology, 39, 372–379.Padial, J.M., Gonzáles, L. & De la Riva, I. (2005) A new species of the Eleutherodactylus discoida-lis Group (Anura: Leptodactylidae) from Andean Humid Montane Forest of Bolivia. Herpeto-logica, 61, 318–325.PADIAL ET AL.56                                       © 2006 Magnolia Press1278ZOOTAXA AppendixSpecimens examinedEleutherodactylus mercedesae: BOLIVIA. Department Cochabamba: “Old”Chapare road, 1400–1500 m, (ZFMK-72571-73); “Old” Chapare road, 1650 m, (ZFMK-72597–99); Pampa Grande, Departmental Protected Area Altamachi, (MHNC-AMS–196); 3.3 km N of Cochabamba to Villa Tunari road on road to San Onofre, at apoint 97.5 km from Cochabamba on road from Cochabamba to Villa Tunari, 1690 m,17º10’S/65º46’W (USNM-257848, holotype); Limbo, 73.5 Km from Cochabamba to VillaTunari, 1950 m, 17º10’S/65º48’W (USNM–165753, paratype). Department La Paz:Hornuni, Cotapata National Park, Prov. Nor Yungas, 16º12’50’’S/67º53’10’’W (CBF-4120); road to Coroico, Serranía Bella Vista, Province Nor Yungas, 16º14’S/67º43’W(CBF-3701). PERU. Department Cusco: Paucartambo, 68 km by road NE of PuenteUnión on río Tachila (Bosque de las Nubes, km 150 on Paucartambo-Atalaya road), 1700m, 13º14’13’’S/71º34’00’’W (USNM-346140).

Distribution and morphological variation of Eleutherodactylus mercedesae Lynch & McDiarmid, 1987 (Amphibia, Anura, Leptodactylidae) with first record for Peru

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