Anurans From Yanachaga 2012 Peru

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    EDITORIAL COMMITTEE /CONSEJO DE REDACCIN

    FOUNDER-MANAGING EDITOR /EDITOR FUNDADOR-GERENTEEnrique La Marca. Laboratorio de Biogeografa, Escuela de Geografa, Universidad de Los Andes, Mrida, Venezuela

    CHIEF EDITOR /EDITOR JEFE Jonh Jairo Mueses-Cisneros. Fundacin FIBA, Coln Putumayo, Colombia

    ASSOCIATE EDITOR /EDITOR ASOCIADOCarlos Martnez Rivera. Philadelphia Zoological Gardens, Pennsylvania, Estados Unidos

    SECRETARY / SECRETARIAIngrid Vanessa Perdomo Castillo. Fundacin FIBA, Coln Putumayo, Colombia

    EDITORIAL ADVISORS / ASESORES EDITORIALES Abraham Mijares-Urrutia.Grif th University, Gold Coast, Queensland, AustraliaJuan Elas Garca-Prez. Universidad de Los Llanos Ezequiel Zamora, Guanare, VenezuelaLuis Felipe Esqueda. Investigador independiente, Santiago de Chile, ChileLuis Fernando Navarrete. Bioreptilia, and Serpentario, Universidad Central de Venezuela,Caracas,Stefan Ltters. Trier University, Trier, GermanySergio Potsch de Carvalho e Silva. Universidade Federal do Rio de Janeiro, R.J., BrazilFernando Castro Herrera. Universidad del Valle, Cali, ColombiaMiguel Trefaut Rodrigues. Universidade de So Paulo, So Paulo, BrazilMarinus Hoogmoed. Museu Paraense Emlio Goeldi, Belem, Par, BrazilDiego Cisneros-Heredia. Universidad San Francisco de Quito, Quito, Ecuador John D. Lynch. Universidad Nacional de Colombia, Bogot, ColombiaGustavo Casas-Andreu. Universidad Nacional Autnoma de Mxico, Mexico City, MxicoFederico Bolaos. Universidad de Costa Rica, San Jos, Costa RicaThasun Amarasinghe. Taprobanica Nature Conservation Society, Kendalanda, Homagama,Sri LankaIndraneil Das. Institute of Biodiversity and Environmental Conservation, Universiti Malaysia,SarawakAaron Bauer. Villanova University, Pennsylvania, United States of AmericaAlan Channing. University of the Western Cape, Bellville, South AfricaSantiago J. Snchez Pacheco. University of Toronto, Toronto, CanadaH. Mauricio Ortega Andrade. Fundacin EcoCiencia, Quito, Ecuador Rudolf von May. University of California, Berkeley, United States of AmericaFelipe Franco Curcio. Universida de de So Paulo, Brazil

    EDITOR ASSISTANT / ASISTENTE DE EDITOR Miguel A. Bastidas. Fundacin BIOGEOS, Mrida, Venezuela

    ISSN 1690-7930 (Printed edition /Edicin impresa)ISSN 1856-9285 (Online edition /Edicin electrnica)

    Legal Deposit / HECHO EL DEPSITO DE LEY PP200402ME2957 (Printed edition /Edicin impresa)

    PPI200802ME2958 (Online edition /Edicin electrnica)

    HERPETOTROPICOS

    Founded in 2004JOURNAL ON TROPICAL AMPHIBIANS AND REPTILES

    SPECIES ON COVER

    Herpetotropicos is indexed and abstracted in / Herpetotropicos es indizada en:Zoological Record (BIOSIS), Wildlife & Ecology Studies Worldwide (University of Delaware Library), and REVENCYT (Directorio ndice de Revistas Venezolanas de Ciencia

    Tecnologa. Biblioteca Digital de la Universidad de Los Andes).

    LIBRARY ACCESS / ACCESO EN BIBLIOTECAS Copies deposited for public access in / Copias depositadas para acceso pblico en:

    American Museum of Natural History (Library), New York, USA. Biblioteca Nacional (Caracas, Venezuela), Biblioteca Febres Cordero, Mrida, Venezuela. Biblioteca Integrada d Arquitectura, Ciencias e Ingeniera de la Universidad de Los Andes, Mrida, Venezuela.

    Finantial support / Financiamiento: Consejo de Desarrollo, Cient co, Humanstico y Tecnolgico de la Universidad de Los Andes (CDCHT-ULA) Mrida, Venezuela.Sponsorship / Patrocinio: Fundacin BIOGEOS para el estudio de la diversidad biolgica.

    INDEXED AND ABSTRACTED IN /INDIZACIN

    PUBLISHER / CASA EDITORA(Institutional address) Direccin institucional

    Laboratorio de Biogeografa, Escuela de Geografa,Universidad de Los Andes (ULABG), va Chorros de Milla,

    Mrida 5101, Venezuela.Tel.: + 58 - 274 - 2401647 Fax: + 58 - 274 - 2401635

    E-mail:[email protected]

    The Vietnamese Mossy Frog,Theloderma corticale (Boulenger, 1903) is a robust, attractively camou aged

    arboreal frog in the family Rhacophoridae. The commonname of the unusual species is derived from its roughlytextured, green and brownish-red skin, which allows it toconceal itself within with its mossy forest habitat. To date,the Vietnamese Mossy Frog has only been found fromthe humid forests of the Mao Son and Tam Dao mountainranges in northern Vietnam; however, the secretive speciesmay actually be distributed more widely.

    The Vietnamese Mossy Frog, like many amphibian speciesin Southeast Asia, is poorly known, and is listed as DataDe cient according to the IUCN Red List of ThreatenedSpecies. Much of the information we have on the biology

    of the species is derived from captive individuals. Maleshave a quiet, melodious advertisement call. Femalesdeposit eggs above water- lled rock cavities and tree-holesin which their tadpoles develop. The characteristic roughskin texture of the species is already evident in tadpolesprior to metamorphosis.

    The greatest threat to the Vietnamese Mossy Frog, andother forest-dependent amphibians in Southeast Asia, ishabitat loss, which is occurring at higher relative rates inSoutheast Asia than in other tropical regions. An additionalconcern may be collection for the international pet trade, forwhich the species is in high demand, but the availability ofcaptive bred individuals may negate this potential threat.

    Text and Photo by Jodi J. L. Rowley ( All rights reserved)

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    23G. CHVEZ et al . - ANURANS FROM YANACHAGA CHEMILLEN NATIONAL PARK

    ANNOTATED CHECKLIST AND ECOLOGICAL NOTES OFANURANS FROM THE SOUTHERN REGION OF YANACHAGACHEMILLEN NATIONAL PARK, CENTRAL ANDES OF PERUGERMN CHVEZ1,3, CATERINA H. COSMPOLIS2and LESLY LUJN1

    1 Centro de Ornitologa y Biodiversidad (CORBIDI), Calle Santa Rita 103, Los Huertos de San Antonio, Surco, Lima, Per.

    2 Universidad Nacional Agraria La Molina (UNALM), Avenida La Molina, La Molina, Lima, Per.

    Abstract: Distribution data, ecological and taxonomical notes of twenty-one species of anurans are presented. This information was collectedon a survey conducted on the southern border of Yanachaga Chemillen National Park, central Andes of Peru, during 2011. Furthermore,preliminary speci c richness and additional eld data on the species occurring in the area are presented in this article.

    Keywords: Amphibia, distribution data, taxonomical notes, ecological notes, survey, speci c richness, Llamaquizu-San Alberto river basin.

    Resumen: G. Chvez, C.H. Cosmpolis y L. Lujn. Lista comentada y notas ecolgicas de los anuros de la regin suNacional Yanachaga Chemillen, Andes centrales de Pru. El registro de veintin especies de anuros, datos de distribucin, ecologa ytaxonoma son el resultado de una expedicin realizada en la regin sur del Parque Nacional Yanachaga Chemillen, en los Andes centralesde Per durante el ao 2011. Adicionalmente, datos preliminares acerca de la riqueza espec ca y otros datos tomados en campo acercade las especies que ocurren aqu son presentados en este artculo.

    Palabras clave: Amphibia, datos de distribucin, notas taxonmicas, notas ecolgicas, inventario, riqueza espec ca, Cuenca del roLlamaquiz-San Alberto.

    233 Send correspondence to /Enviar correspondencia a:vamp [email protected]

    HERPETOTROPICOS Vol. 8(1-2):23-38 ISSN 1690-7930 (Printed) ISSN 1856-9285 (Online)Printed in Venezuela. All rights reserved

    Copyright 2012 BIOGEOS

    INTRODUCTION

    The eastern slopes of the Andes are characterized for its endemism

    and speci c richness. Many expeditions have explored this area,but its great amphibians biodiversity was only noticed in the early1980s (Lynch 1979, Lynch and Duellman 1980) in Ecuador andthe late 1990s in Peru (Duellman and Pramuk 1999).

    Many other biological expeditions followed on the Peruvianmontane forest, resulting on the discovery of several new frogspecies (Duellman and Chaparro 2008), especially on the vicinityof Yanachaga-Chemillen National Park (YCNP), in the provinceof Pasco. Some of these species include a harlequin frog of thegenus Atelopus (Lehret al . 2008); aHypsiboas hylid frog (Lehretal . 2010); fourPhrynopusspecies (Chaparro et al . 2008, Duellmanand Hedges 2008), thirteenPristimantis species (Boano et al . 2008;Duellman and Hedges 2005, 2007; Lehret al . 2004a, 2006), onespecies of Hypodactylus (Lehr 2005) and one species ofNoblella (Lehret al . 2004b). This list involves some critically endangeredfrogs (e.g. Ameerega planipaleae, Morales and Velazco 1998; Atelopus oxapampae, Lehr et al . 2008). These ndings highlightthe importance of the YCNP area that might be one of the lastrefugia for amphibian species on the eastern slopes of the Andes.

    Unfortunately, this area is constantly threatened by agricultural

    activities that seem to be invading the border of the national park.Consequently, a long term study was initiated in the southern borderof the YCNP and this article presents the results obtained from theeld trip performed from July to August 2011.

    STUDY AREA AND METHODSStudy AreaThe study area is located at the southern border of YCNP, on thecentral Peruvian Andes (Fig. 1), characterized by montane and cloudforest vegetation. Some arboreal and shrub species of the genera Acacia, Chusquea, Clusia, Cecropia, Hedyiosmum, Heliconia,Podocarpus and Weinmannia, and herbaceous species of thegenera Bomarea, Carex , Oreobolus; as well as lichens and manyspecies of orchids and bromeliads are present in the area (Foster1985, 1986). Several waterfalls and dissected mountains shape theterrain and the soil structure consists of clay and rocks (mainly alongriversides). Agricultural land and human settlements are found onthe lower basin of the region.

    Amphibians were surveyed on the two main basins within the

    Received / Recibido:24 M AY 2012 Accepted / Aceptado:17 SEP 2012Published / Publicado:15 DEC 2012

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    study area: San Alberto stream and Llamaquizu river. For practicalreasons, these basins were classi ed in four sampling locations(Table 1): (i) Lower Llamaquiz (LLB), located at the YCNP bufferarea and with agricultural areas, riparian forest and secondaryforest; (ii) Upper Llamaquiz (ULB), at the southern border of YCNP,characterized by riverside forest, primary and secondary forest; (iii)Lower San Alberto (LSAB), on the buffer area with the presence ofsecondary forest and human settlements; and 4) Upper San Alberto(USAB), inside YCNP and only with primary forest.

    METHODS

    Diurnal and nocturnal transects were used to survey amphibiansin the area. Seventeen transects of 200 m2 were surveyed on eachlocality, with a minimum distance of 200 meters between transects(Heyeret al . 1994). Two researchers walked along each transectat low speed during the day (08:00 - 13:00h) and night (20:00 -02:00h). All anurans heard calling were identi ed and registered.

    Deep-water sampling was also performed by using a dip net andmoving it slowly through the water. Caught animals were identi ed,measured, photographed and released at the site of capture, if nofurther identi cation was required. For further identi cation, somespecimens were preserved in 10% buffered formalin, and keptin 70% ethanol. Detailed examination of collected material was

    conducted by GC and LL on preserved material. Acronym for voucherspecimens is CORBIDI= Centro de Ornitologia y Biodiversidad(Lima, Peru).

    Capture success and speci c richness (S) was tested usingrarefaction analysis (PAST statistical program, Hammeret al . 2001)in order to avoid potential bias due to differences on sampling size(Adrainet al. 2000). Hammeret al . (2001) based his algorithmon Krebs (1989) analysis, which used a log Gamma function forcomputing combinatorial terms. We used this method to obtain theexpected S value on the study area.

    RESULTS A total of 21 species were recorded, corresponding to eightgenera and ve families: Bufonidae (3 spp.), Centrolenidae (1 sp.),Dendrobatidae (1 sp.), Hylidae (2 spp.) and Strabomantidae (14spp.) (Table 2). A total of 305 individuals were recorded, showing arelative high abundance in the study area. However, rarefaction curvetends to be in nite (see Discussion), as an evidence that the totalsampling is not enough to represent the real richness in the area.Our results also show a high speci c richness in ULB and USAB;both localities are away from human settlements. Only two specieswere common in all localities: the strabomantid frogsPristimantisbromeliaceus and Pristimantis saggitulus.

    FIG. 1.Map showing the study area and the four localities sampled.Mapa que muestra el rea de estudio y las cuatro localidades evaluadas.

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    25G. CHVEZ et al . - ANURANS FROM YANACHAGA CHEMILLEN NATIONAL PARK

    Species Accounts

    FAMILY BUFONIDAE Atelopus oxapampae (Lehr, Ltterset Lundberg, 2008)

    (Fig. 2, A-B).

    This relatively recent described frog is one of the rarest speciesin this study. Only one individual was recorded (CORBIDI 010270)in the Lower San Alberto basin, within the type locality. It is the onlyharlequin frog that occurs in this area. The individual was caughtat night showing low activity at that time. The habitat where it wascaptured matches the data presented by Lehr et al. (2008) whorecorded the six type specimensin a narrow creek. This creek ischaracterized by a dense cloud forest which vegetation covers thecreek, and direct sunlight rarely reached the ground. The individualwas found on leaves at 30 cm above the ground, very close to thecreek (less than 20 cm of distance). The predominant vegetationon the area is secondary forest and crops. Based on our eld dataand habitat described by Lehret al. (2008), the major threat for thisspecies is the human impact in the area; thus, extensive agricultureis practiced for plantations of granadilla or coffee. In consequence ofthis practice, agrochemical substances for plantations are used hereand the farmers usually throw the remnants to creeks or streams,places where Atelopus oxapampae could breed. Sympatric speciesare Pristimantis saggitulus and Hyalinobatrachium carlesvilai ; bothare nocturnal species and therefore their niches do not overlap with A. oxapampae. Remarkably, this harlequin frog is the only diurnalamphibian recorded in the entire San Alberto basin.

    Rhinella cf. leptoscelis (Fig. 2, C-D).

    The recognized species Rhinella leptoscelis Boulenger, 1912, wasrediscovered and redescribed by Padialet al . (2009). Individualswere captured in LLB (CORBIDI 010166, 010175-76, 010192,

    010194) and agree with Padialet al. (2009). They have tympanumdistinct, large paratoids glands -separated from eyelid and tympanumby the supratympanic crests-, body surface covered by spinytubercles and the basal webbings are serrated. Nevertheless, theyare different because they have small tubercles on the dorsum (largeinR. leptoscelis) and yellow iris (green inR. leptoscelis). Furthermore,Padial et al . (2009) defined its distribution from Cordillera deVilcabamba, Cusco Region, southern Peru, to Chapare region, inBolivia. If we can con rm the identity of these specimens consideredas Rhinellacf. leptoscelis in this study, it would be the rst record forthe species at central Peru.

    Cordillera de Vilcabamba is approximately 400 km from YCNP. Although it is a long distance, other records for species fromCordillera de Yanachaga or surrounding areas, even most northerlythan the locality of this study, have been reported in the past years(Chvez et al . 2008). Therefore, it would not be strange to recordspecies from Cordillera de Vilcabamba in Yanachaga; nevertheless,more studies need to be performed and more specimens should becaptured to con rm this record. The habitat de ned for this speciesis humid montane forest, same habitat whereRhinellacf. leptosceliswas recorded. In addition, we have records of juveniles in theriverside vegetation of LLB, surrounded by agriculture zones andopen areas, which evidence that R. cf. leptoscelis tadpoles coulddevelop in disturbed rivers and open territories. The only sympatricspecies with this toad wasPristimantis bipunctatus, a species highlyrecorded in open areas and agriculture zones. Despite the presenceofP. bipunctatus, there is no evidence about the type of competitionbetween both species. Apparently, the great differences in length(Rhinellacf. leptoscelis is much larger thanP. bipunctatus) could becaused by a separation of niches.

    Rhinella yanachaga (Lehr, Pramuk, Hedgeset Cordova, 2007)

    (Fig. 2, E-F).

    This arboreal toad belongs, according to morphological characters,to the Rhinella veraguensis group, having orbito-tympanic andpostorbital crests weak and parotoid glands moderately large.However, it is distinguished from other similar species of theveraguensis group by having a small snout-to-vent lenght (maximumSVL 45.7 mm) and males with vocal slits. The specimens recordedin this research (CORBIDI 010236, 010279010283) agree very wellwith all morphological characters recognized forRhinella yanachaga

    and with the color pattern described for this species. Sites weresampled at the surroundings of the type locality forR. yanachaga and this species was recorded only on the primary forests of ULBand USAB, being abundant on the latest. Nevertheless, the specieswas not recorded in the secondary forest or other habitats presentin the other sampling sites; this is apparently a remarkable habitatquality preference, not mentioned by Lehret al . (2007). Theseauthors de ned the type locality inside YCNP as cloud forests,mainly primary, with low levels of human impact, which match oureld data for the species. All individuals were recorded in bushes,between 0.5-2 m above the ground, although it is not possible to

    TABLE 1.Coordinates of the sampled localities. LLB=LowerLlamaquizu Basin; ULB=Upper Llamaquizu Basin; LSAB=LowerSan Alberto Basin; USAB=Upper San Alberto basin.TABLA 1. Coordenadas de las Localidades evaluadas. LLB=Cuencabaja de Llamaquiz; ULB= Cuenca alta de Llamaquiz;LSAB=Cuenca baja de San Alberto; USAB=Cuenca alta de San Alberto.

    Locality

    LLB

    ULB

    LSAB

    USAB

    Southern

    1003652.06

    1003646.62

    1003240.62

    1003202.24

    Western

    7502112.52

    7501701.90

    7502203.89

    7502058.31

    Elevation (m)

    2080

    2148

    2231

    2765

    Coordinates

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    con rm their habitat preferences. Individuals were captured at night,between 19:0023:00h, resting on leaves.Rhinella yanachagawas found sympatric withPhrynopus bracki, Prhynopus tribulosus,Pristimantis bromeliaceus, Pristimantis lucasi, Pristimantissp. 1and Pristimantis stictogaster . With the exception ofP. stictogaster ,these Pristimantis species were recorded in the same place, usedbushes as microhabitat, and were active at the same time thanR. yanachaga. This overlap of niches could suggest a potentialcompetition between the species and, in that case, probably themicrohabitat use would be the determinant for the relationship.

    FAMILY CENTROLENIDAEHyalinobatrachium carlesvilai (Castroviejo-Fisher, Padial,

    Chaparro, Aguayoet de la Riva, 2009)(Fig. 3, A-B).

    This is the rst record of this glassfrog for the Pasco region. The

    presence of this species involves a new locality but not an extensionof its distribution. Specimens recorded (CORBIDI 010178-010183,

    010272-010274) exhibit all diagnostic characters described byCastroviejo-Fisheret al . (2009) such as: iridophorous absent on theventral parietal peritoneum, snout truncate in dorsal and lateral view,liver bulbous; humeral spine is absent in adult males, nger webbingabsent between ngers I and II and basal between ngers II andIII.Hyalinobatrachium carlesvilaiis the most abundant species inhabitats related to streams and it was found in bushes, bromeliads,Heliconiaspp. and some high herbs. Most of the specimens recordedwere males and captures were performed during the night, whenmany male adults were calling on leaves above water. This datacorrespond with the ndings of Castroviejo-Fischeret al. (2009).Even though the vegetation in streams was dense, the places wherethis glassfrog was recorded are inside an area strongly impacted byagricultural and construction activities similar to the habitat forH.carlesvilai in Brazil (Cisneros-Herediaet al . 2009). The level of theimpact in the type locality or localities for paratypes is not mentionedin the description. In the study area,H. carlesvilai is sympatric with

    Pristimantis saggitulus, P. bromeliaceus and P. cruciocularis. Thesespecies are mainly nocturnal, likeH. carlevilai , but the strata of the

    TABLE 2.Species recorded by each locality. LLB=Lower Llamaquizu Basin; ULB=Upper Llamaquizu Basin; LSAB=Lower San AlbeBasin; USAB=Upper San Alberto basin.TABLA 2.Especies registradas por cada localidad. LLB=Cuenca baja de Llamaquiz; ULB= Cuenca alta de Llamaquiz; LSAB=Cuencabaja de San Alberto; USAB=Cuenca alta de San Alberto.

    Clase

    Amphibia

    Orden

    Anura

    Family

    BufonidaeBufonidae

    BufonidaeCentrolenidaeDendrobatidaeHylidaeHylidaeStrabomantidaeStrabomantidaeStrabomantidaeStrabomantidaeStrabomantidaeStrabomantidaeStrabomantidaeStrabomantidae

    StrabomantidaeStrabomantidaeStrabomantidaeStrabomantidaeStrabomantidaeStrabomantidae

    Species

    Atelopus oxapampaeRhinella leptoscelis

    Rhinella yanachagaHyalinobatrachium carlesvilai Amerega planipaleaeHypsiboas aguilari Scinax oreitesPhrynopus tribulosusPhrynopus bracki Phrynopus auriculatusPristimantis adiastolusPristimimantis albertusPristimantis bipunctatusPristimimantis bromeliaceusPristimantis cruciocularis

    Pristimantis leucorrhinusPristimantis lucasi Pristimantis rhabdocnemusPristimantis sagittulusPristimantissp.Pristimantis stictogaster

    LLB

    X

    XX

    XXX

    X

    ULB

    X

    XX

    X

    XXXXX

    XXXX

    X

    LSAB

    X

    X

    X

    XXX

    XX

    USAB

    X

    XXX

    X

    X

    XXXXX

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    forest that they use is lower than the one used by the mentionedspecies.

    FAMILY DENDROBATIDAE Ameerega planipaleae (Moraleset Velazco, 1998)

    (Fig. 3, C-D).

    Seven specimens were recorded and none of them were collected.They all presented the diagnostic characters for this species(Morales and Velazco 1998) such as: skin on dorsum granular,

    dorsal coloration yellow with two dorsolateral yellow stripes andventral coloration pale blue or turquoise with black reticulations.Coloration on groins and anterior surfaces of thighs is black with redor orange spots. Some individuals presented red spots on ventralsurfaces of shanks, a color variation previously described by Medina-Muller and Chvez (2009). This species is known only from thetype locality, a location that, unfortunately, lacks precise geographicinformation. Data based on eld observations performed by GC,de ne A. planipaleae distribution as very restricted, justifying its

    FIG. 2.Dorsal and ventral view of: AB) Atelopus oxapampae; CD)Rhinellacf. leptoscelis; EF)Rhinella yanachaga.Vista dorsal y ventral de: AB) Atelopus oxapampae; CD)Rhinellacf. leptoscelis; EF)Rhinella yanachaga.

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    inclusion under the Critically Endangered category of IUCN (2004).Ecological and natural history data are scarce for this frog. Our eldobservations evidenced that it is a typical diurnal species, being veryactive in the afternoon, between 17:00-18:30 h. Habitat recordedfor this frog is secondary forests, where individuals use mainlyleaf litter at day and night. Despite this species being sympatricwith Pristimantis cruciocularis, P. bipunctatus, P. bromeliaceus and Hyalinobatrachium carlesvilai , there is not overlapping ofmicrohabitats or activities. Consequently, the chances of ecologicalcompetition with other frogs are scarce, because it is the only diurnalspecies recorded in the LLB. Our eld data reveals that this frog isthreatened by habitat loss and degradation, since the surroundedarea is completely covered by farms and human buildings.

    FAMILY HYLIDAEHypsiboas aguilari (Lehr, Faivovichet Jngfer, 2010)

    (Fig. 3, E-F).

    Many individuals of this species wererecorded but only three werecollected (CORBIDI 010196-98). Specimens exhibit a head widerthan the body, a snout rounded in dorsal view and slightly truncatein lateral view, prominent eyes with the upper half of palpebralmembrane translucent and the lower half with brown reticulations,hypertrophied arms in males, and dorsolateral folds; all of thesecharacters differ from other similar species (Lehret al . 2010).This species was extremely abundant in impacted areas. In fact,tadpoles, juveniles an adult were recorded massively in creeks orstreams along farms or towns, and were absent on streams insideprimary forest or areas without evidence of human impact. Thisspecies does not climb trees or bushes, and is recorded frequently

    hidden between low vegetation. Although no adults of other specieswere recorded sympatrically withH. aguilari , tadpoles of Scinaxoreites were recorded in the same creeks; these data is coincidentwith the data given in the description by Lehret al . (2010) whichmention the fact thatS. oreites tadpoles occurs syntopically withH.aguilari tadpoles in the Llamaquizu basin. All males were recordedactive at night, from 18:00 24:00 h and the males density wasconsiderably higher than that for females.

    Scinax oreites (Duellmanet Wiens, 1993)(Fig. 4, A-B).

    Although this species can be abundant in some areas at northernPeru (GC, unpublished data), in the study area only three individualswere recorded (CORBIDI 010264-65, 010269). Both juvenilescorresponded with the species description (Duellman and Wiens,1993) presenting yellow groins, the main diagnostical character forthis species. The individuals were captured on riparian vegetation,where they used bushes and some trees as shelter and were usuallyfound on leaves 12 meters above the ground. They were activeduring the nights (20:00 23:00 h). Lehret al. (2010) mentionedthis species occurs simpatrically withHypsiboas aguilari in openareas and even agricultural zones; this data was corroborated inthe present research.

    FAMILY STRABOMANTIDAEPhrynopus auriculatus (Duellman et Hedges, 2008)

    (Fig. 4, C-D).This species was recorded only on primary forest, at the USAB, insideYCNP. The only individual captured (CORBIDI 010304) exhibits themost remarkable traits for the species: bearing a tympanic membraneand tympanic annulus, and toe V was slightly longer than toe III. Thetype locality is the same wherePhrynopus auriculatuswas recorded.The temperature in this habitat is the lowest for the whole studyarea and the elevation was the highest.P. auriculatus was recordedamong lichens and sharing microhabitat withPhrynopus bracki and Phrynopus tribulosus, species that have a very similar length.Other frog recorded using the same microhabitat wasPristimantisstictogaster , but this species is longer than the others, and all thesespecies are nocturnal. Based on these observations, it is possible tosuppose that the niches are overlapping betweenP. auriculatus, P.bracki and P. tribulosusand, in consequence, a potential ecologicalcompetition occurs between these species.

    Phrynopus bracki (Hedges, 1990)(Fig. 4, E-F).

    This species is (likePhrynopus auriculatus) one of the rarest in thestudy area and was only found at the type locality. Two individualswere captured (CORBIDI 010302-03). Both of them showed a smalllength (19.00 mm or less) a vestigial rst toe with the width equalto the length, and a fourth toe long and broad. This frog was foundin primary forest with cold temperatures. The species is sympatricand syntopic withP. auriculatus and Phrynopus tribulosus, probablyestablishing a relationship of competition because the overlappingof the niches. Males were calling inside the lichens or leaf-litter that

    they inhabit, and the species is active at night.

    Phrynopus tribulosus (Duellmanet Hedges, 2008) (Fig. 5, A-B).

    This small frog was recorded sympatrically withPhrynopus auriculatus and Phrynopus bracki in a primary forest. Individuals captured(CORBIDI 010299-010301) exhibit diagnostic characteristicsmentioned by Duellman and Hedges (2008) such as: tympanicmembrane not differenced and tympanic annulus absent, uppereyelid bearing small tubercles, vocal slits and nuptial pads absent,subconical tubercle on the heel and outer edge of tarsus with rowof subconical tubercles and a green dorsal coloration. This specieswas previously known just by a single individual found deep within amossy bank by day (Duellman and Hedges, 2008). We found threespecimens, all of them using lichens and leaf litter as microhabitat forshelter; two of them were calling inside leaf-litter at night, at 20:30 h.

    Pristimantis adiastolus (Duellmanet Hedges, 2008)(Fig. 5, C-D).

    This frog, a member of thePristimantis peruvianus group, wasrecorded by only one individual (CORBIDI 010221). This species is

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    cryptic withPristimantis bipunctatus and in the original descriptionofP. bipunctatus (Duellman and Hedges, 2005) several individualsof Pristimantis adiastoluswere included but were later separatedthrough morphological and phylogenetic analyses. Diagnosticcharacters for P. adiastolus are present in the specimen capturedin this study, such as skin on dorsum shagreened and weaklyareolate on venter, dorsolateral folds present, tympanic membrane

    differentiated and posterior surfaces of thighs brown. The specimenwas found in a primary forest in the ULB, on a leaf 1.5 meters abovethe ground, in simpatry withPristimantis bromeliaceus, P. saggitulus and P. rhabdocnemus. It is dif cult to establish a relationshipbetween P. adiastolus and the rest of species co-inhabiting this area,but this species is probably closer toP. rhabdocnemus which usesthe same microhabitats. The species is different fromP. saggitulus,

    FIG. 3.Dorsal and ventral view of: AB) Hyalinobatrachium carlesvilai ; CD) Ameerega planipaleae; EF) Hypsiboas aguilari.Vista dorsal y ventral de: AB)Hyalinobatrachium carlesvilai; CD) Ameerega planipaleae; EF)Hypsiboas aguilari.

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    because it uses vegetation (trees, bushes or herbs) very close tobody waters; andP. bromeliaceus uses a higher stratum in the forestthan P. adiastolus. According to the description (Duellman andHedges 2008) this species inhabits humid montane forest at 1200meters of elevation; however, this study presents a new altitudinalrecord: 2140 m, a difference almost one thousand meters higherthan previous known records.

    Pristimantis albertus (Duellmanet Hedges, 2007)(Fig. 5, E-F).

    This species was intensively recorded in three out of four sampled

    areas, increasing the known number of individuals, because it wasdescribed only with two specimens. The voucher frogs (CORBIDI010202-010203, 010219, 010255, 010261, 010263, 010285,010291) show characters such as shagreened skin on dorsumwith scattered small tubercles, low dorsolateral folds roughenedand discontinuous, tympanic membrane differentiated, tympanicannulus distinct, upper eyelid lacking tubercles and the color on theposterior surfaces of thighs is brown, all of them coincident with thediagnosis of Pristimantis albertus. Furthermore, this species wasrecorded not only in the type locality (USAB) but also in the Lowerand Upper Llamaquiz basins. The habitat where this species wasfound includes primary and secondary forests. All the individuals were

    FIG. 4.Dorsal and ventral view of: AB)Scinax oreites; CD) Phrynopus auriculatus; EF)Phrynopus bracki.Vista dorsal y ventral de: AB)Scinax oreites; CD)Phrynopus auriculatus; EF)Phrynopus bracki.

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    captured on leaves, far from any body of water, between 0.3 to 1.5 mabove the ground, and were active at night. The species is sympatricwith Pristimantis lucasi , P. bromeliaceus, P. rhabdocnemus, P.bipunctatusand P. saggitulus, but presents an overlapping of nicheswithP. lucasi, P. rhabdocnemus and P. bipunctatus, which probablycauses competition between the species.

    Pristimantis bipunctatus (Duellmanet Hedges, 2005) (Fig. 6, A-B).

    This species is a member of thePristimantis conspicillatus groupand was recorded remarkably along the Llamaquizu basin (lower andupper) and in Lower San Alberto basin, where it was one of the mostabundant species. Collected specimens (CORBIDI 010170-010174,010201, 010204, 010251, 010289) show the diagnostic charactersdescribed by Duellman and Hedges (2005), such as: skin on dorsumnely shagreened with a couple of black warts on the scapular region,dorsolateral folds absent and color on the posterior surface of thighsis brown with cream ecks.Pristimantis bipunctatus was recordedon riverside low vegetation, being the predominant species in this

    FIG. 5.Dorsal and ventral view of: AB)Phrynopus tribulosus; CD)Pristimantis adiastolus; EF)Pristimantis albertus.Vista dorsal y ventral de: AB)Phrynopus tribulosus; CD)Pristimantis adiastolus; EF)Pristimantis albertus.

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    habitat and, in fact, the most abundant anuran inhabiting nearbody waters in LLB. In lower and upper San Alberto this frog wasrecorded simpatrically withPristimantis saggitulus; however, theirmicrohabitats do not overlap.Pristimantis bipunctatus use the lowerstrata in the vegetation next to a stream or river, whileP. saggitulus use the medium strata (bushes or young trees).

    Pristimantis bromeliaceus (Lynch, 1979)(Fig. 6, C-D).

    This frog, a member of thePristimantis lacrimosus group, was themost abundant species in the entire study area. The specimens(CORBIDI 010184-010185, 010200, 010217, 010222010230,010248-010249, 010253, 010294-010295) correspond very wellwith the description: bearing a sub-acuminate snout, small tubercleson the upper eyelid, dentigerous processes of vomers oblique,prominent and small tubercles on heels and outer side of tarsi.The coloration exhibited by these specimens was yellowish, pale

    reddish or green and some individuals have an interorbital bar thatcorresponds with the description and with the colors mentioned byDuellman and Lehr (2009). This arboreal species uses the higherstrata of primary forest, secondary forest, and riparian vegetation.It is usually recorded active at night, with males calling actively from19:00 to midnight or beyond. The distance between males in somecases was only 0.1 m, and only some females were found. SeveralPristimantis species were recorded simpatrically withPristimantisbromeliaceus, but only a few were using the same microhabitat. OnlyHyalinobatrachium carlesvilai could overlap in microhabitat and timeof activity. But according to our eld observations, when males ofP. bromeliaceus and H. carlesvilai are using the same site (leavesof bush or trees), males of H.carlesviali call underneath leaves

    and P. bromeliaceus on top of the leaves. Although bromeliadsare reported as a usual shelter, we caught P. bromeliaceus by dayon leaves and inside some young leaves of Heliconiaspp. Thisspecies is considered under the Vulnerable category by IUCN (2008)having as justi cation its occurrence area, which is probably lessthan 20.000 Km2 in a very fragmented region of the Andes in Peru.Despite this,P. bromeliaceus shows healthy populations in most ofthe habitats present here, even in areas strongly impacted by humanactivity or buildings. However, more studies are needed to increasethe knowledge about the real conservation status of this species.

    Pristimantis cruciocularis (Lehr, Lundberg, Aguilaret vonMay, 2006)(Fig. 6, E-F).

    This frog was recorded only in the Llamaquizu basin. Although it wasnot an abundant species, it had a moderate abundance in upperLlamaquizu basin. The specimens (CORBIDI 010186-010191,010238) exhibit diagnostic characters for the species: two tubercleson the upper eyelid, skin on dorsum shagreened with scattered smallspicules, males lack vocal sacs, red or orange and white spots ongroins and anterior surfaces of thighs. Only one captured individual(CORBIDI 010238) exhibits a different color on dorsum and groins(Fig. 7), presenting a uniform red color on groins and anteriorsurfaces of thighs, and a longitudinal brown and pale brown pattern

    on dorsum.Pristimantis cruciocularis was recorded at medium strataon primary and secondary forests, on leaves and 1.5 m above theground. The species was always recorded during night and was foundon leaf-litter, in vegetation close to streams or creeks. These datacorrespond with the species description information (Lehret al . 2006).Sympatric species wereHyalinobatrachium carlesvilai, Pristimantisbipunctatus, P. saggitulus and P. bromeliaceus. Of these, onlyP.saggitulus is overlapping the microhabitat withP. cruciocularis andboth are nocturnal, which indicates a potential source of competitionbetween these frogs.

    Pristimantis leucorrhinus (Boano, Mazzottiet Sindaco,2008)(Fig. 8, A-B).

    This species, a member of thePristimantis unistrigatus group, wasrecorded in the study area by only two individuals (CORBIDI 010237,010277). These specimens agree with the species description

    (Boano et al . 2008) by having skin tuberculate on dorsum withprominent pointed tubercles on limbs and areolate with scatteredrounded tubercles on venter, several tubercles on the upper eyelid,one of them is large and conical and groins and anterior surfaces ofthighs coloration black with white spots. According to the charactersmentioned by Boanoet al . (2008) the speci c name leucorrhinus isderived from the latin leucos meaning white and the Greek nounrhinos meaning snout because the holotype (the only previouslyknown specimen for this species) has a white spot in the pit of thenose, like the specimens collected in this research. However, thischaracter is present or absent in several individuals of many speciesof the Pristimantis unistrigatus group. Although the records arescarce to de ne any ecological data, both individuals ofPristimantis

    leucorrhinus were recorded at primary forest, on leaves at 1.5 mabove the ground and during the night. The species is sympatricwithPristimantis bromeliaceus and P. rhabdocnemus, and probablyhas a niche overlap withP. rhabdocnemus, recorded in the samemicrohabitat and having the same activity periods. Therefore, it isprobable that both taxa have a potential competition relationship.

    Pristimantis lucasi (Duellmanet Chaparro, 2008)(Fig. 8, C-D).

    This frog, a member of thePristimantis unistrigatus group, wasrecorded in a few localities, all of them in the upper basins(Llamaquizu and San Alberto). Four individuals were recorded(CORBIDI 010206-010207, 010297-010298), everyone showeddiagnostic characters given by Duellman and Chaparro (2008) suchas: skin on dorsum smooth with scattered subconical tubercles,tympanic membrane not differentiated, tympanic annulus absent,upper eyelid bearing two large and conical tubercles, heel bearinga single and conical tubercle, males with vocal slits, coloration ongroins, posterior surfaces of thighs and ventral surfaces of shankspale greenish tan. This species was only known from the montaneforest, below the crest of Abra Esperanza, therefore we add a newplace for this species occurrence: in the ULB, approximately 20 Kmfrom the type locality. The habitat where this species was found is

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    FIG. 6.Dorsal and ventral view of: AB) Pristimantis bipunctatus; CD)Pristimantis bromeliaceus; EF) Pristimantis cruciocularisVista dorsal y ventral de: AB)Pristimantis bipunctatus; CD) Pristimantis bromeliaceus; EF)Pristimantis cruciocularis.

    mainly primary forest and only one individual was recorded in a trailinside secondary forest, although very close to a primary forest.Pristimantis lucasi uses microhabitats in the medium strata of thevegetation, was captured on leaves, 11.5 m above the ground andduring the night. Sympatric species werePhrynopus auriculatus.P. tribulosus, P. bracki, Pristimantis albertus, P. bipunctatus, P.bromeliaceus, P. cruciocularis, P. leucorrhinus, P. stictogaster,P. rhabdocnemus and Pristimantis sp. From these species, P.lucasi microhabitat overlaps withP. albertus, P. cruciocularis, P.leucorrhinus and P. rhabdocnemus. These species use the samestrata of the vegetation and have similar length; accordingly, a

    probable trophic competition could exist among these frogs.

    Pristimantis rhabdocnemus (Duellmanet Hedges, 2005)(Fig. 8, E-F).

    This species was one of the three most abundant species in thestudy area. It was recorded in three out of four sampling sites.Seventeen individuals were captured (CORBIDI 010205, 010220,010239-010240, 010242-010247, 010252, 010254, 010262, 010275,010292-010293) and they correspond very well with the speciesdescription (Duellman and Hedges, 2005): by having shagreened

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    FIG. 8.Dorsal and ventral view of: AB)Pristimantis leucorrhinus; CD) Pristimantis lucasi ; EF)Pristimantis rhabdocnemus.Vista dorsal y ventral de: AB)Pristimantis leucorrhinus; CD)Pristimantis lucasi; EF)Pristimantis rhabdocnemus.

    DISCUSSIONThe collection of 22 species presented in this study provides apreliminary view of the amphibian fauna from the southern regionof YCNP. This collection also contains records of great importancefor their biogeographical and taxonomical significance withinthe context of the current knowledge of Andean herpetofauna.Several specimens represent records of species that used to beconsidered endemic for a single or few localities inside YCNP(e.g . new localities are given here forPristimantis lucasi and P.

    leucorrhinus). Additionally, other species have been found outsidetheir type localities (e.g. Ameerega planipaleae, Phrynopus bracki,Phrynopus tribulosus and Phrynopus auriculatus).

    Speci c richness was tested with a rarefaction analyses on thetotal study area. According to Fig. 10, the lack of asymptote on thespecies accumulation curve suggests that more species might bepresent in the study area. This result is supported on the remarkableincrement of speci c richness for Strabomantid frogs at the end of theeldwork. Species described from Cordillera de Yanachaga, absentin this study belong, mainly, to the family Strabomantidae (Duellman

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    and Hedges 2005; Duellman and Chaparro 2008), being just a fewexceptions. The possibility to nd new taxa is open, and evidencedon the record of Pristimantissp., thus, additional survey effortsare likely to reveal species missed in this survey. Furthermore,some frequently encountered species (e.g., Hypsiboas aguilari ,Hyalinobatrachium carlesvilai , Pristimantis bromeliaceusand P.saggitulus) are as well considered moderately common by otherherpetologists (pers. comm. with J.C. Chaparro and J. Icochea). Although our data is too limited to provide an analysis of abundance,

    temporal or elevational variation in herpetofaunal communities;ecological data presented here is a preliminary result that precedesfuture contributions based on the same study area.

    ACKNOWLEDGEMENTSWe would thank the Instituto del Bien Comn (IBC), Oxapampa of ce,Peru, for trust in GC work; and the Birdlife International and RAREConservation, by their support on the eldwork. For the permits, we

    FIG. 9.Dorsal and ventral view of: AB) Pristimantis saggitulus; CD) Pristimantis sp; EF) Pristimantis stictogaster.Vista dorsal y ventral de: AB)Pristimantis saggitulus; CD)Pristimantis sp; EF)Pristimantis stictogaster.

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    FIG. 10.Rarefaction curve of the study area.Curva de rarefaction del rea de estudio.

    are indebted to the Servicio Nacional de Areas Naturales Protegidas,Oxapampa Of ce and Direccin General de Flora y Fauna Silvestre,Peru. And in particular, we are indebted with gratitude to AimyCaceres for her valuable help, recommendations and contributionsto the manuscript; Cesar Laura, Federico Rizopatrn, Alex Samarand Guido Casimiro, staff of IBC, for their valuable and unbelievablehelp and logistic assistance; with Florencia Trama by her hospitalityin San Alberto; with Mecho by her friendship and hospitality inLlamaquizu, and with Daniel, Alex and Nildo Ermitao by their

    enthusiastic eld assistance.

    REFERENCES

    Adrain, J.M., S.R. Westrop and D.E. Chatterton. 2000. Siluriantrilobite alpha diversity and the end-Ordovician mass extinction.Paleobiology 26:625646.

    Boano, G., S. Mazzotti and R. Sindaco. 2008. A new peculiar frogspecies of the genus Pristimantisfrom Yanachaga-ChemillnNational Park, Peru. Zootaxa 1674:5157.

    Castroviejo-Fisher,S., J.M. Padial, J.C. Chaparro, R. Aguayoand I. de la Riva. 2009. A new species ofHyalinobatrachium(Anura: Centrolenidae) from the Amazonian slopes of the central Andes, with comments on the diversity of the genus in the area.Zootaxa 2143:24-44.

    Chaparro, J.C., J.M. Padial and I. de la Riva. 2008. Two sympatricnew species of Phrynopus (Anura: Strabomantidae) fromYanachaga Chemilln National Park (central Peruvian Andes).Zootaxa 1761:4958.

    Chvez, G., M. Medina-Mller and A. Pereyra. 2008. Amphibia, Anura, Hylidae,Osteocephalus leoniae: Distribution extension.Check List 4:401403.

    Cisneros-Heredia, D.F., C. Strsmann, R.B. vila and R.A.Kawashita-Ribeyro. 2009. Amphibia, Anura, Centrolenidae,

    Hyalinobatrachium carlesvilaiCastroviejo-Fisher, Padial,Chaparro, Aguayo and de la Riva 2009: First country record,Brazil. Checklist 6(2):225-226.

    Duellman, W.E. and J.J. Wiens. 1993. Hylid frogs of the genusScinax Wagler, 1830, in Amazonian Ecuador and Peru.Occasional Papers of the Museum of Natural History of theUniversity of Kansas 153:1-57.

    Duellman, W.E. and J.B. Pramuk. 1999. Frogs of the genusEleutherodactylus (Anura: Leptodactylidae) in the Andes ofnorthern Peru. Scienti c Papers of the Natural History Museum.The University of Kansas 13:178.

    Duellman, W.E and S.B. Hedges. 2005. Eleutherodactyline frogs(Anura: Leptodactylidae) from the CordilleraYanachaga in centralPeru. Copeia 2005:526538.

    Duellman, W.E and S.B. Hedges. 2007. Three new species ofPristimantis(Lissamphibia: Anura) from montane forests of theCordillera Yanachaga in central Peru. Phyllomedusa 6:119135.

    Duellman, W.E. and J.C. Chaparro. 2008. Two distinctive newspecies of Pristimantis (Anura: Strabomantidae) from theCordillera Oriental with a distributional synopsis of strabomantidsin Central Peru. Zootaxa 1918:1325.

    Duellman, W.E and Hedges, S.B. 2008. Two new minute species ofPhrynopus(Anura: Strabomantidae) from the Cordillera Orientalof Peru. Zootaxa 1675:5966.

    Duellman, W.E. and E. Lehr. 2009.Terrestrial-breeding frogs(Strabomantidae) in Peru. Munster, Germany: Nature und TierVerlag, 382 pp.

    Foster, R. 1985. Principales especies de ora del Parque NacionalYanachaga Chemilln. Pp. 148-176.In: 1987. A.J. Brack (ed.).Plan Maestro del Parque Nacional Yanachaga Chemilln. Pro

    Naturaleza, Lima, Per.Foster, R. 1986. Preliminary checklist. Plants of Parque Yanachaga(including the Palcazu Valley and Cordillera San Matas) Selvacentral, Per. Field Museum of Natural History, Chicago. 49 pp.

    Hammer, O., D.A.T. Harper and P.D. Ryan. 2001. Past:Paleontological Statistics Software Package for Education andData Analysis. Palaeontologia Electronica 4(1):9 Pp.

    Hedges, S.B. 1990. A new species of Phrynopus (Anura:Leptodactylidae) from Per. Copeia 1990:108112.

    Heyer, W.R., M.A. Donnelly, R.W. McDiarmid, L.C. HayekFoster. 1994. Methods for Measuring and Monitoring BiologicalDiversity: Standard Methods for Amphibians. SmithsonianInstitution Press. Washington, D.C.

    Krebs, C.J. 1989. Ecological Methodology. Harper and Row, NewYork. 425 pp.

    Lehr, E., C. Aguilar and W.E. Duellman. 2004a. A strikingnew species of Eleutherodactylus from Andean Peru (Anura:Leptodactylidae). Herpetologica 60:275280.

    Lehr, E., C. Aguilar and M. Lundberg. 2004b. A new species ofPhyllonastes from Peru (Amphibia, Anura, Leptodactylidae).Journal of Herpetology 38:214218.

    Lehr, E. 2005. A new species of theEleutherodactylus nigrovittatus group (Anura: Leptodactylidae) from Andean Peru. Herpetologica61:199208.

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