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Volume 6 Number 1 June 2007Volume 6 Number 1 June 2007

Journal of HerpetologyJournal of Herpetology

Correspondence to:Jaime BertoluciDepartamento de Ciências Biológicas – ESALQ – USPAv. Pádua Dias, 11 Caixa Postal 913418-900 Piracicaba – SP – BRAZILE-mail: bertoluc@esalq.usp.br

Subscriptions and back issues:www.phyllomedusa.esalq.usp.br

Cover: A male Atelopus zeteki from PanamaPhoto: Michael Redmer

ContentsVolume 6 Number 1 - June 2007

Articles

A striking new species of Anolis lizard (Squamata, Iguania) from PanamaErik Hulebak, Steven Poe, Roberto Ibáñez, and Ernest E. Williams .................................................................. 5

Ecology of Alopoglossus angulatus and A. atriventris (Squamata, Gymnophthalmidae)in western AmazoniaLaurie J. Vitt, Teresa Cristina S. Ávila-Pires, Maria Cristina Espósito, Shawn S. Sartorius,and Peter A. Zani ............................................................................................................................................... 11

A new species of the Eleutherodactylus lacrimosus assemblage (Anura, Brachycephalidae)from the lowland rainforest canopy of Yasuni National Park, Amazonian EcuadorShawn F. McCracken, Michael R. J. Forstner and James R. Dixon ................................................................. 23

Nocturnal position in the Panamanian Golden Frog, Atelopus zeteki (Anura, Bufonidae),with notes on fluorescent pigment trackingErik D. Lindquist, Scott A. Sapoznick, Edgardo J. Griffith Rodriguez, Peter B. Johantgen,and Joni M. Criswell .......................................................................................................................................... 37

Morphological variation in Leptodactylus lutzi (Anura, Leptodactylidae) with descriptionof its advertisement call and notes on its courtship behaviorPhilippe J. R. Kok, Marcelo N. C. Kokubum, Ross D. MacCulloch, and Amy Lathrop ..................................... 45

Visual and acoustic signaling in three species of Brazilian nocturnal tree frogs (Anura, Hylidae)Luís F. Toledo, Olívia G. S. Araújo, Lorena D. Guimarães, Rodrigo Lingnau, and Célio F. B. Haddad ........... 61

Short Communication

Coelomic helminths of five colubrid snake species (Serpentes, Colubridae) from Costa RicaStephen R. Goldberg and Charles R. Bursey ................................................................................................... 69

Book Review

Lima, A. P., W. E. Magnusson, M. Menin, L. K. Erdtmann, D. J. Rodrigues, C. Keller, and W. Hödl. 2006.Guide to the Frogs of Reserva Adolpho DuckeBy Janalee P. Caldwell ...................................................................................................................................... 73

Phyllomedusa - Journal of HerpetologyAll material originally published in Phyllomedusabelongs to Departamento de Ciências Biológicas -ESALQ - USP, and may not be reproduced, stored in aretrieval system, or transmitted in any form or by anymeans, electronics, mechanical, photocopying,recording, or otherwise, without prior written permissionof the publishers.ISSN 1519-1397Printed in Brazil in June 2007

Journal of Herpetology

VOLUME 6 - NUMBER 1

JANUARY/JUNE - 2007

Phyllomedusa

IS PUBLISHED BY DEPARTAMENTO DE CIÊNCIAS BIOLÓGICAS,ESCOLA SUPERIOR DE AGRICULTURA “LUIZ DE QUEIROZ”,

UNIVERSIDADE DE SÃO PAULO, AND PARTIALLY SUPPORTED BYFUNDAÇÃO DE AMPARO À PESQUISA DO ESTADO DA BAHIA

(process number APR0019/2007)

BIANNUAL

Phyllomedusa Piracicaba v.6 n.1 pp. 1-80 Jan/Jun 2007

ISSN 1519-1397

Journal of Herpetology

PIRACICABA

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Phyllomedusa: Journal of Herpetology – vol. 6, n. 1, 2007. – Piracicaba, SP, Brazil: Departamento de Ciências Biológicas,Escola Superior de Agricultura “Luiz de Queiroz”, Universidade de São Paulo.

v.; ilVol. 1 (2002) to Vol. 3 (2004) published by Melopsittacus Publicações Científicas, Belo Horizonte, MG, Brazil.Vol. 1 (2002) to Vol. 5 (2006) Phyllomedusa: Journal of Neotropical HerpetologyBiannualArticles and abstracts in English; additional abstracts in Portuguese, Spanish, French, Italian, or German are optional.ISSN 1519-13971. Herpetology CDU - 598

Editorial BoardJournal of Herpetology

Augusto Shinya AbeUniversidade Estadual Paulista, Brazil

Rogério Pereira BastosUniversidade Federal de Goiás, Brazil

Janalee P. CaldwellUniversity of Oklahoma, USA

Ulisses CaramaschiMuseu Nacional, Brazil

Guarino R. ColliUniversidade de Brasília, Brazil

Carlos A. G. CruzMuseu Nacional, BrazilWilliam E. Duellman

University of Kansas, USAPaula Cabral Eterovick

Pontifícia Universidade Católica de Minas Gerais, BrazilJulián Faivovich

American Museum of Natural History, USARenato Neves Feio

Universidade Federal de Viçosa, Brazil

Editor-in-ChiefJaime Bertoluci

Universidade de São Paulo, Brazil

First Associate EditorPedro Luís Bernardo da Rocha

Universidade Federal da Bahia, Brazil

Ross AlfordJames Cook University, Australia

Franco AndreoneMuseo Regionale di Scienze Naturali di Torino, Italy

Teresa Cristina Sauer de Ávila PiresMuseu Paraense Emilio Goeldi, Brazil

Néstor G. BassoCentro Nacional Patagónico, Argentina

James BogartUniversity of Guelph, Canada

Ignacio De la RivaMuseo Nacional de Ciencias Naturales, Spain

Alain DuboisMuséum National d’Histoire Naturelle, France

Stephen GoldbergWhittier College, USA

Tim HallidayOpen University, UKPhilippe J. R. Kok

Institut Royal des Sciences Naturelles de Belgique, Belgium

Axel KwetStaatliches Museum für Naturkunde Stuttgart, Germany

Ross D. MacCullochRoyal Ontario Museum, Canada

Peter A. MeylanEckerd College NAS, USA

Carlos Arturo NavasUniversidade de São Paulo, Brazil

Carlos I. PiñaCONICET, ArgentinaStephen J. Richards

South Australia Museum, AustraliaChristine Strüssmann

Universidade Federal de Mato Grosso, BrazilLinda Trueb

University of Kansas, USAVanessa Kruth Verdade

Universidade de São Paulo, BrazilRichard Carl Vogt

Instituto Nacional de Pesquisas da Amazônia, Brazil

Associate Editors

Board Members

Ronaldo FernandesMuseu Nacional, Brazil

Darrel R. FrostAmerican Museum of Natural History, USA

Célio Fernando Batista HaddadUniversidade Estadual Paulista, Brazil

Walter HödlUniversität Wien, Austria

Flora Acuña JuncáUniversidade Estadual de Feira de Santana, Brazil

Arturo I. KehrCONICET, ArgentinaWilliam Magnusson

Instituto Nacional de Pesquisas da Amazônia, BrazilOtávio Augusto Vuolo Marques

Instituto Butantan, BrazilJosé Peres Pombal Jr.Museu Nacional, Brazil

Carlos Frederico Duarte da RochaUniversidade Estadual do Rio de Janeiro, Brazil

Miguel Trefaut RodriguesUniversidade de São Paulo, Brazil

Catherine A. ToftUniversity of California, Davis, USA

Monique Van SluysUniversidade Estadual do Rio de Janeiro, Brazil

Luciano Martins VerdadeUniversidade de São Paulo, Brazil

Oscar Flores VillelaUniversidad Nacional Autónoma de México

Laurie J. VittUniversity of Oklahoma, USA

Hussam ZaherMuseu de Zoologia, Univ. de São Paulo, Brazil

Barbara ZimmermanUniversity of Toronto, Canada

Web MasterFábio A. Bazanelli

Universidade de São Paulo, Brazil

Phyllomedusa - 6(1), June 2007

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Phyllomedusa – Journal of Neotropical Herpetology celebrates the completion of 5 successful yearsof uninterrupted contributions to the field of Herpetology. During this period, the journal has attracted theattention of herpetologists from Latin America, as well as from North America, Europe, and Australia. Atotal of 170 authors from 16 countries have chosen PHYLLOMEDUSA for their scientific communications. Ofthe 14 new species were described in our pages, three were frogs, two amphisbaenians, three lizards, andsix snakes.

The journal’s success seems to have resulted from (1) the high quality of its editorial board, whichincludes active members of scientific community from several different countries, such as Argentina,Austria, Brazil, Canada, Mexico, and USA; (2) the fact that PHYLLOMEDUSA papers can be retrieved frommany important reference indexes and databases, such as Biological Abstracts, Zoological Record, CABIPublishing, Elsevier Science Bibliographic Databases, The Reptile Database, Bibliomania’sHerpetological Contents, and Herpetological Literature Database; (3) the wide scope of the journal, whichpublishes papers in all fields of Herpetology; and (4) the availability of all papers at journal’s website priorto the distribution of its printed version.

PHYLLOMEDUSA is broadening its geographical scope to reflect the globalization of its contributors andtheir interests. From now on, papers will be published on taxa from the Neotropics, as well as the rest ofthe world. Accordingly, the name of the journal is changing to PHYLLOMEDUSA – Journal of Herpetology,and the Editorial Board has incorporated new Associate Editors to represent scientific communities fromother zoogeographic regions. There is a total of 20 internationally recognized herpetologists from 11countries and three continents. The diversity of this group will enhance scientific communication amongherpetologists around the world, and this should be the primary goal of any scientific periodical.

Many people greatly assisted the staff of PHYLLOMEDUSA during its first 5 years of publication, and weowe a debt of gratitude to each of them. I am especially indebted to two colleagues in particular. AndréNemésio convinced me to create PHYLLOMEDUSA and published it for the first 3 years during difficulttimes. Pedro Rocha, applied his intelligence, discipline, and expertise to the process of manuscriptevaluation, and immeasurably enhanced the quality of the published material. A first-class journal dependsentirely on the quality of its authors, editorial board members, and ad hoc referees, and PHYLLOMEDUSAalways counted on many of the best ones indeed. Associate editors who have served during these past 5years have contributed greatly to the quality of the journal and made my life every so much easier. Finally,I wish to thank subscribers for their support and many libraries around the world that have includedPHYLLOMEDUSA in their collections. Idmar Pedro is the designer responsible for the extraordinary graphicquality of the journal, and Fábio A. Bazanelli is gratefully acknowledged for his voluntary work as webdesigner and webmaster.

Financial support was provided by UFMG – Universidade Federal de Minas Gerais (2002–2004),USP – Universidade de São Paulo (since 2005), FEALQ – Fundação de Estudos Agrários Luiz de Queiroz(since 2005), CNPq – Conselho Nacional de Desenvolvimento Científico e Tecnológico (2003), FAPESB– Fundação de Amparo à Pesquisa do Estado da Bahia (2007) and some private institutions. Richard Vogthelped us obtain funds on several occasions. Breck Bartholomew deserves my most sincereacknowledgements for his continuous help accepting international subscriptions and divulgingPHYLLOMEDUSA since 2003.

My last acknowledgements are extended to the new Associate Editors, who have accepted myinvitation and have joined PHYLLOMEDUSA in its mission of serving as an international outlet for originalherpetological research.

Jaime BertoluciEditor

Editorial

Phyllomedusa - 6(1), June 2007

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Introduction

Visual communication was first described fordiurnal anurans and seems to be widelydistributed among them (review in Hödl andAmézquita 2001). However, it has been recently

Visual and acoustic signaling in three species ofBrazilian nocturnal tree frogs (Anura, Hylidae)Luís Felipe Toledo1, Olívia G. S. Araújo1, Lorena D. Guimarães2, Rodrigo Lingnau3, andCélio F. B. Haddad1

1 Departamento de Zoologia, Instituto de Biociências, Universidade Estadual Paulista. Caixa Postal 199, 13506-970, RioClaro, SP, Brazil. E-mail: toledolf2@yahoo.com.

2 Departamento de Biologia, Instituto de Biociências, Universidade Federal de Goiás. Caixa Postal 131, 74001-970,Goiânia, GO, Brazil.

3 Laboratório de Herpetologia, Museu de Ciências e Tecnologia & Faculdade de Biociências, Pontifícia UniversidadeCatólica do Rio Grande do Sul. Av. Ipiranga, 6681, 90619-900, Porto Alegre, RS, Brazil.

Received 3 August 2006.Accepted 7 February 2007.Distributed June 2007.

Phyllomedusa 6(1):61-68, 2007© 2007 Departamento de Ciências Biológicas - ESALQ - USP

ISSN 1519-1397

AbstractVisual and acoustic signaling in three species of Brazilian nocturnal tree frogs(Anura, Hylidae). Visual communication seems to be widespread among nocturnalanurans, however, reports of these behaviors in many Neotropical species are lacking.Therefore, we gathered information collected during several sporadic field expeditionsin central and southern Brazil with three nocturnal tree frogs: Aplastodiscus perviridis,Hypsiboas albopunctatus and H. bischoffi. These species displayed various aggressivebehaviors, both visual and acoustic, towards other males. For A. perviridis wedescribed arm lifting and leg kicking; for H. albopunctatus we described theadvertisement and territorial calls, visual signalizations, including a previouslyunreported behavior (short leg kicking), and male-male combat; and for H. bischoffiwe described the advertisement and fighting calls, toes and fingers trembling, leglifting, and leg kicking. We speculate about the evolution of some behaviors andconcluded that the use of visual signals among Neotropical anurans may be much morecommon than suggested by the current knowledge.

Keywords: Anura, Hylidae, Aplastodiscus perviridis, Hypsiboas albopunctatus,Hypsiboas bischoffi, visual and acoustic signaling, agonistic interactions, physicalcombats.

observed that some nocturnal species alsoperform visual communication (e.g., Bertoluci2002; Abrunhosa and Wogel 2004; Hartmann etal. 2005, Giasson and Haddad 2006). Visualcommunication is usually associated withreproduction (e.g., for female attraction) oraggressive interactions related to territoriality(Hödl and Amézquita 2001, Rosenthal et al.2004), but also with predation (Bertoluci 2002)and intraspecific communication (Abrunhosa

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and Wogel 2004). Currently, a total of 27 visualdisplays have been described, including legstretching, throat display, toe/fingers trembling,and body rising (Hödl and Amézquita 2001,Bertoluci 2002; Abrunhosa and Wogel 2004;Hartmann et al. 2005, Hirschmann and Hödl2006). Although these behaviors are complexand apparently widespread, their origin is oftendifficult to access, despite the efforts of Hödland Amézquita (2001).

We present new observations on the noctur-nal visual signaling of three hylid species:Aplastodiscus perviridis, Hypsiboas albo-punctatus, and H. bischoffi. Although territorialvocalizations have been reported to exist in therepertoire of H. albopunctatus (Haddad et al.1988, Bastos et al. 2003), no other informationis available for the remaining taxa. Visualsignaling has not been described for any of thesespecies. Only one report of visual signaling hasbeen made to date for Aplastodiscus [A.eugenioi in Hartmann et al. 2004 as Hyla sp.(aff. ehrhardti)], and one for Hypsiboas (H.albomarginatus in Giasson and Haddad 2006).Both were observed during aggressive inte-ractions. Besides providing direct observationsof visual and aggressive interactions for thestudied taxa, we described novel vocalizations(territorial and fighting calls) for two of thosespecies, and suggest new hypotheses for theorigin of the visual signals herein described.

Material and Methods

We gathered information collected duringseveral sporadic field expeditions in central andsouthern Brazil. Observations of Aplastodiscusperviridis were made on 8 March 2006 atFazenda São Francisco, Municipality of PonteSerrada, State of Santa Catarina. Observationsof Hypsiboas albopunctatus occurred in March2002 at Municipality of Mineiros, State ofGoiás, in June 2005 at Municipality of Santanado Riacho (Serra do Cipó), State of MinasGerais, and in February 2006 at Municipality ofSanta Rita do Passa Quatro (Parque Estadual de

Vassununga - PEV), State of São Paulo.Observations of H. bischoffi occurred inDecember 2005 at Municipality of Treviso,State of Santa Catarina, and in September 2003at Municipality of Iporanga (Parque EstadualTurístico do Alto Ribeira - PETAR), State ofSão Paulo. Sampling methods included adlibitum, focal-animal, all occurrences of somebehaviors, and sequences (Altmann 1974;Lehner 1996). For Hypsiboas albopunctatus,aggressive behaviors were stimulated byplayback of advertisement and territorial calls,or by placing a conspecific male about 20 cmfrom the resident calling male. With theremaining species no manipulation was made.The terminology for describing visual signalingfollows that presented by Hödl and Amézquita(2001) complemented by Hartman et al. (2005).

Aggressive vocalizations were recorded witha Marantz cassette tape recorder (PMD222),equipped with an external directionalmicrophone (Audiotecnica AT835b) positionedca. 50 cm from the calling male. We usedchrome cassette tapes at 4.75 cm/s. We analyzedthe calls using Raven 1.2 software (16 bits ofresolution, 44 kHz of frequency sampling, FFTand frame length of 256 samples). Theterminology for the vocal analysis followed thatpresented in Toledo and Haddad (2005), exceptfor “pseudonotes”, which was considered groupof pulses within a note, accordingly to Heyer etal. (1990). Visual signaling of all species wasrecorded in video cameras and the files areavailable upon request.

Results

Under natural conditions, all speciesdisplayed various aggressive behaviors, bothvisual and acoustic, towards other males. Thesame happened in response to encountersprovoked by the introduction of a conspecificmale, as well as when males were apparentlyisolated from a group or potential competitor.Playback of advertisement and/or territorial callsalso stimulated aggressive reactions. The visual

Toledo et al.

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signals observed were leg kicking, limb lifting(sensu Hartmann et al. 2005), and toe/fingerstrembling (sensu Hödl and Amézquita 2001).

Two males of A. perviridis were observedcalling in temporary water bodies inside anartificial channel at the edge of a small road.One of the males displayed visual signals whileperched on a leaf 20 cm above the ground. Thesecond male was calling while perched on a leaf15 cm above the ground and 3 m apart from thefirst one. Male number one emitted 44advertisement calls within a 43-second periodand displayed leg kicks with both hind limbs, aswell as arm lifting, in no particular order. Itstarted by kicking its left leg, and repeated itfour, eight, and 19 seconds later. It then liftedthe left arm and kicked its right leg. After 18

seconds, it kicked its left leg (Figure 1). Noother male or female was seen in the vicinity.

Males of H. albopunctatus were callingwhile perched 1 m from the ground on themargins of temporary ponds, permanent ponds,and rivulets with very low water flow. A higherdensity of males was usually observed the dayafter a rain. When calling, males exhibitedbright white to yellow colorful spots and stripeson the femoral region and on the inguinalregion, respectively (Figure 2A). We observedvisual displays when males of H. albopunctatuswere introduced near conspecific males andduring playbacks. The visual signaling of H.albopunctatus seems to be performed in anescalated ordination: initially males trembledtheir fingers (n = 5), than turned toward the

Figure 1 - Adult male of Aplastodiscus perviridis performing leg kicking (initial position and visual signal: A and B,respectively) and arm lifting (initial position and visual signal: C and D, respectively). Observations madeat Fazenda São Francisco, Municipality of Ponte Serrada, State of Santa Catarina, Southern Brazil.

Visual and acoustic signaling in Brazilian nocturnal tree frogs

A

B

C

D

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intruder male (introduced by us in the territoryof the resident male, n = 2), lifted an arm (n =2), performed a short leg kick (n = 11), andfinally initiated physical combat with theintruder male (n = 2 in provoked encounters andn = 2 in natural conditions). This sequence ofdisplays was observed twice in the same nightwith two different resident males. Physicalcombats occurred among males perched on thebushes (n = 3) and on the floor (n = 1). In onecase, the resident male pursuited the intruder

before combat. During the fight, both maleshugged face-to-face (Figure 2D) and tried tothrow the opponent out of its calling site throughhind leg kicking. Males also used their pre-pollical spines to cause injuries in the opponent(see Figure 2D). Scars were commonly observedin males of H. albopunctatus. The short legkicks observed seems to correspond to thatdescribed by Hartmann et al. (2005). However,the males do not extend the entire legbackwards, but only partially. In its maximum

Figure 2 - Adult males of Hypsiboas albopunctatus performing visual displays (A-C) and fighting (D). Male callingand displaying bright spots in femoral region and bright stripes in inguinal region (A); male during agonisticinteraction prior to (B) and during short leg kicking (C). Observations were made in the Municipality ofSanta Rita do Passa Quatro (B-C), State of São Paulo, and Municipality of Santana do Riacho (A and D),State of Minas Gerais, southeastern Brazil.

Toledo et al.

A B

D C

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“A”, “B”, and “T”) in the vocalizations of H.albopunctatus (Figure 3). A fourth type of note(note “C”) seems to be one of the twoadvertisement notes (A or B) emitted weaklyand in isolation. Notes “A” and “B” had lowenergy frequencies above 3.6 kHz (Table 1;Figure 3), and can be considered side bandseries due to the pulsatile nature of the call (seeHeyer et al. 1990). These notes were describedby different authors: Haddad et al. (1988),Heyer et al. (1990), De la Riva et al. (1997),Kwet et al. (2002), and Bastos et al. (2003).

Figure 3 - Waveform (above) and spectrogram (below) ofthe advertisement call (notes “A” to “C”) andterritorial call (note “T”) of Hypsiboasalbopunctatus, recorded at Municipality ofSanta Rita do Passa Quatro, State of São Pau-lo, Southeastern Brazil; air temperature: 24ºC.

Table 1 - Notes of Hypsiboas albopunctatus calls recorded at Parque Estadual de Vassununga, Municipality of SantaRita do Passa Quatro, State of São Paulo, southeastern Brazil (air temperature: 24ºC). Values presented asmean ± standard deviation (range).

extension, the tibia achieved an angle ofapproximately 45º with the tarsus, and the foottouched the leave where the frog was located(Figures 2B-C).

We recognized three types of notes (notes

Figure 4 - Spectrogram of the initial note of the adver-tisement call (A) and the fighting call (B) ofHypsiboas bischoffi, recorded at Parque Esta-dual Turístico do Alto Ribeira (PETAR),Municipality of Iporanga, State of São Paulo,Southeastern Brazil; air temperature: 22ºC.

Visual and acoustic signaling in Brazilian nocturnal tree frogs

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Figure 5 - Two sequences of visual displays in Hypsiboas bischoffi stimulated by playback of advertisement call of thesame individual. A) Initial position of the first sequence; B) trembling of right toes; C) trembling of leftfingers; D) initial position of the second sequence; E) right leg lifting; F) left leg kicking. Observations weremade at the Municipality of Treviso, State of Santa Catarina, Southern Brazil.

Note “T” was considered to have a territorialfunction since its rate of emission in provokedencounters increased considerably when anothermale was placed nearby. Its emission rate alsoincreased in response to playbacks of any typeof vocalizations, even those including the othertypes of notes. Territorial notes were pulsed (ca.80 pulses per note) and emitted in a low fre-quency (ca. 1 kHz). The amplitude of frequen-cies ranged from 0.58 to 3.41 kHz, with thenotes been emitted during 0.43 seconds onaverage (Table 1; Figure 3). In two cases, malesemitted several sequences of notes “T”. Malesalso were able to make multiple combinations ofthese notes. The most commonly heard phraseswere “AB”, but “TB”, “TBB”, and “ABT” werealso heard.

Males of H. bischoffi called while observedat about 1 m from the ground, around the edges

of lentic water bodies (temporary or permanentponds, and rivulets with very low water flow).Their advertisement call includes two types ofnotes. An initial note (Figure 4A), of meandominant frequency of 1.894 kHz (frequencyrange = 1.572 to 2.092; N = 8), was emitted inisolation or preceding a warbling series of 15 to20 short notes (see descriptions in Bokermann1967; Heyer et al. 1990 as Hyla multilineata).When stimulated by playback of their ownadvertisement calls, two males displayed legkicking, foot lifting, and finger and toetrembling (Figure 5). Physical combat was alsoobserved between two males on a second night.When the intruder male approached a resident ofa calling site, the later started to emit a group ofinitial notes (Figure 4A) and chased the intruderinto a shrub over a flooded ground. Both malesunderwent physical interactions, assuming the

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same position described for H. albopunctatus.During the fight both individuals emitted twoseries of seven notes as a fighting call (Figure4B). These notes were spaced by 147milliseconds (range = 112 – 119; SD = 22.9; n =12). Each note had a mean dominant frequencyof 1.47 kHz (range = 1.29 – 1.55; SD = 0.06; n= 14), mean minimum frequency of 1.168 kHz(range = 1.063 – 1.262; SD = 0.06; n = 14), andmean maximum frequency of 1.765 kHz (range= 1.609 –1.871; SD = 0.07; n = 14).

Discussion

Our observations provide new evidences thatnocturnal visual signaling may be more frequentthan previously believed (see also Hartmann etal. 2005, Giasson and Haddad 2006). More datashould be generated in the near future, enablingbroader analyses and inferences (Hödl andAmézquita 2001). By now, we can suggest somenew hypotheses about the origin of these visualbehaviors and propose a new nomenclature for apreviously undescribed behavior. Visual signalsare believed to be derived from pre-existingmovements (Hödl and Amézquita 2001). Forexample, arm waving, hind-feet lifting, limblifting, and leg kicking may be related tomovements produced during physical inte-ractions between males (Hödl and Amézquita2001, Giasson and Haddad 2006). Following thesame criteria, we suggest that fingers tremblingmay be originated from movements of comfort.Similar movements can be observed when theindividuals are changing their positions andpostures on the vegetation (pers. obs.). Malesusually turn towards their opponents just beforeinitiating a physical fight (Abrunhosa and Wogel2004, present study). Therefore, fingerstrembling could indicate a change in the positionof the anuran as a prelude to the movementtoward the opponent’s direction. Interpretingthis signal as one of the last stages of non-physical agonistic interactions could preventfights and the costs associated to them (e.g.,Kluge 1981, Martins and Haddad 1988, Wells

1988). Short leg kicking, as described here forH. albopunctatus, may be referred as a newvisual display since it does not fit in anypreviously proposed category (Hödl andAmézquita 2001, Hartmann et al. 2005). Itsorigin and function seem to be the same as thosesuggested for leg kicking or arm waving, forexample (see above), therefore it may alsoprevent physical combats.

Visual signaling tied to complex courtshipbehavior has been reported to occur inAplastodiscus eugenioi (Hartmann et al. 2004).Complex courtship behavior was also reportedfor A. perviridis, which involves tactile stimuliand vocalizations, but without reports of visualsignals (Haddad et al. 2005). Moreover, therehave been no reports of signal use betweenmales in the absence of females as observed byus for A. perviridis. We suggest that the visualsignaling observed here for A. perviridis couldbe related to inter-male territoriality, as observedin other species discussed by us.

All the three species herein studied (Aplasto-discus perviridis, Hypsiboas albopunctatus, andH. bischoffi) make use of a combination ofvisual and acoustic signaling. The aggressi-veness of the males can be escalated, rangingfrom territorial vocalizations to signals thatprelude fights, such as fingers trembling and legkicking. By having complex calls with differentnotes, males can intensify the signal that theywant to communicate (e.g. Cardoso & Haddad1984, Lingnau et al. 2004). In the present study,H. albopunctatus increased the emission ofaggressive notes when another male was nearby.The same situation has been observed forseveral other hylid species, such as Den-dropsophus minutus (Cardoso & Haddad1984), Hypsiboas albomarginatus (Giassonand Haddad 2006), and Scinax fusco-marginatus (Toledo and Haddad 2005). Theuse of visual signals among Neotropicalanurans may be much more common thansuggested by currently knowledge, reinforcingthe need of extensive studies and reports, suchas the present one.

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Acknowledgements

We are grateful to Luís O. M. Giasson, IvanSazima, and two anonymous referees for thediscussion and suggestions on early versions ofthe manuscript. Luís O. M. Giasson assistedwith field activities. Fabrício Oda provided thepicture of the fighting males of Hypsiboasalbopunctatus. RAN-IBAMA (IBAMA/RAN183/05) and COTEC-IF provided necessarycollecting permits. CAPES and CNPq grantedscholarships; Biota-FAPESP (proc. n. 01/13341-3) and CNPq (bolsa de produtividade em pes-quisa to CFBH) funded the CFBH’s herpetologylaboratory. LFT also tanks Idea Wild, Neotro-pical Grassland Conservancy for equipmentdonation, and the Bioacustics Research Programof the Cornell Laboratory of Ornithology for thesoftware license granted.

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Altmann, J. 1974. Observational study of behavior:sampling methods. Behaviour 15: 227–267.

Bastos R. P., M. A. F. Bueno, S. L. Dutra, and L. P. Lima.2003. Padrões de vocalização de anúncio em cincoespécies de Hylidae (Amphibia: Anura) do Brasil cen-tral. Comunicação do Museu de Ciências e TecnologiaPUCRS, Série Zoologia 16: 39–51.

Bertoluci, J. 2002. Pedal luring in the leaf-frogPhyllomedusa burmeisteri (Anura, Hylidae,Phyllomedusinae). Phyllomedusa 1: 93–95.

Bokermann, W. C. A. 1967. Notas sôbre cantos nupciaisde anfíbios brasileiros (Anura). III. Anais da Acade-mia Brasileira de Ciências 39: 491–493.

Cardoso, A. J. and C. F. B. Haddad. 1984. Variabilidadeacústica em diferentes populações e interações agres-sivas de Hyla minuta (Amphibia, Anura). Ciência eCultura 36: 1393–1399.

De la Riva I., R. Márquez and J. Bosch. 1997. Descriptionof the advertisement calls of some South AmericanHylidae (Amphibia, Anura): taxonomic andmethodological consequences. Bonner ZoologischeBeiträge 47: 175–185.

Giasson, L. O. M. and C. F. B. Haddad. 2006. SocialInteractions in Hypsiboas albomarginatus (Anura:Hylidae) and the significance of acoustic and visualsignals. Journal of Herpetology 40: 171–180.

Haddad C. F. B., G. V. Andrade and A. J. Cardoso. 1988.Anfíbios anuros no Parque Nacional da Serra da Canas-tra, Estado de Minas Gerais. Brasil Florestal 64: 9–20.

Haddad, C. F. B., J. Faivovich and P. C. A. Garcia. 2005.The specialized reproductive mode of the treefrogAplastodiscus perviridis (Anura: Hylidae). Amphibia-Reptilia 26: 87–92.

Hartmann, M. T., P. A. Hartmann and C. F. B. Haddad.2004. Visual signaling and reproductive biology in thenocturnal treefrog, Hyla sp. (aff. ehrhardti) (Anura:Hylidae). Amphibia-Reptilia 25: 395–406.

Hartmann, M. T., L. O. M. Giasson, P. A. Hartmann, andC. F. B. Haddad. 2005. Visual communication inBrazilian species of anurans from the Atlantic forest.Journal of Natural History 39: 1675–1685.

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On behalf of the 6th World Congress ofHerpetology Organizing Committee, we wouldlike to invite colleagues to organize and proposesymposia. Proposals should state clearly that thesymposium organizer is ready to assume theorganization of the symposium. Proposalsubmissions will be open until 30 September2007 and should include the following items:

1) A symposium title; 2) The full name ofthe organizer or chair of the proposedsymposium; 3) Institutional affiliation of theorganizer and contacts (full address, e-mail,phone and FAX numbers; 4) A brief text (200words) explaining the goal of the symposium; 5)A list of potential speakers and their topics.

Presentations should be of 20 minuteduration for each participant (this time shouldinclude time for questions and discussions). Ifthe Symposium organizer has compellingreasons for other time allotments, thesymposium organizer must make his/her casewith the Organizing Committee for an alternatescheme of time allotments that would allow fortime coordination with the other symposia beingheld simultaneously.

Call for Symposium Proposals

Organizers should indicate the participantsto prepare their presentations in Power Point forPC. Below are the names and contact of themembers the Symposium Committee of the 6thWorld Congress of Herpetology - please sendyour proposal to one of the members:

Dr. Carlos Frederico D. Rocha (Head ofSymposium Committee) (Rio de Janeiro, Brazil)– cfdrocha@uerj.br

Dr. Claudia Keller (Manaus, Brazil) –keller@inpa.gov.br

Dr. Jean-Marc Hero (Queensland, Australia)- m.hero@griffith.edu.au

Dr. Martin Whiting (Johannesburg, SouthAfrica) - martin@gecko.wits.ac.za

Dr. Miguel Trefaut Rodrigues (São Paulo,Brazil) – mturodri@usp.br

Dr. Richard Griffiths (Kent, England) -R.A.Griffiths@kent.ac.uk

Dr. W. Ronald Heyer (Washington, D.C.,U.S.A.) - HEYERR@si.edu

More information and details about themeeting may be found on the WCH web page:http://www.worldcongressofherpetology.org/index.php?section=11

Phyllomedusa - 6(1), June 2007

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INSTRUCTIONS TO AUTHORSGeneral information . PHYLLOMEDUSApublishes Articles concerning the whole field ofHerpetology. PHYLLOMEDUSA also maintainssections for Short Communications and BookReviews. Manuscripts will be considered oncondition that they have not been publishedelsewhere or are not under consideration forpublication, in whole or in part, in another journalor book. Manuscripts must be submitted by e-mailor in the digital format (diskette or CD). Highquality color photographs are also accepted.Manuscripts must be written in English. If Englishis not your primary language, please be assured itis reviewed before submission. Please direct anyquestion about submission to the main editor.Publication in PHYLLOMEDUSA, includingcolor pictures, is free of charge. Scope. Manuscripts must contain significant newfindings of fundamental and general herpetologicalinterest and may not be considered if they do notmeet these criteria. Surveys and descriptions ofnew species are published only where there issufficient new biological information or taxonomicrevision also involved to render the paper of gene-ral herpetological interest. Low priority is given toconfirmatory studies, works that are primarilydescriptive in nature, investigations primarily oflocal or regional interest, techniques unless ofbroad application, species range extensions, anddescriptions of phenomena based on scanty data.Manuscripts should include a clear statement ofpurpose and/or hypothesis to be tested by the work,and may be rejected if this is lacking.

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José Wellington Alves dos Santos1,2, RobertaPacheco Damasceno1,2, Pedro Luís Bernardoda Rocha 2,31 Departamento de Zoologia, Instituto deBiociências, Universidade de São Paulo, 05508-900, São Paulo, SP, Brazil. E-mails:wly@ib.usp.br, rpdama@usp.br.2 Departamento de Zoologia, Instituto de Biolo-gia, Universidade Federal da Bahia, 40170-210,Salvador, BA, Brazil. E-mail: peurocha@ufba.br.3 Author for correspondence.

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(Riassunto), or German (Zusammenfassung)could be included

4) Keywords in English and (optional) in one ofthe languages above cited

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the authors to describe the study area andinclude a location map)

7) Results8) Discussion9) Acknowledgements10) References11) Tables with their legends12) Figures with their legendsSubtitles (items 5 to 8) should not be used in ShortCommunications. Please refer to a recent issue forformat details.

Tables. Tables should be numbered consecutivelyin Arabic numerals and placed after References.Very large tables should be placed at the end of thearticle as Appendixes. Tables should be formattedwith horizontal, but not vertical, rules. In the text,tables should be referred as Table 1, Tables 2 and3, Tables 2-5.

Figures (including color photographs). Figuresshould be numbered consecutively and placed afterthe Tables. Please embed them in the text file. Fi-gures should be referred to in the text as Figure 1,Figures 2 and 3, Figures 2-5, Figure 4A (not Fig. 1,figure 1, fig. 1, or Figure 4a). After acceptance allfigures must be sent as separate files (notembedded in a text file) and their names shouldfollow the model: author(s) name(s) Figure X (e.g.Pritchard et al. Figure 1A). High quality color orblack and white photographs, or computer-generated figures (e.g. maps) are preferable.

Nomenclature. For the first mention of a species,give its complete binomial name; for taxonomicpapers give the authority and date. Scientificnames must be shown in italics. Also italicizewords not adopted into the original language of themanuscript. Use SI units, e.g.

Time: “08:16 h”, “17:52 h”.Distances and areas: “12 km2”, “55 m”, “17,840ha”.Geographic coordinates: “04o43’23’’ S,45o58’04’’ W”.Temperatures: 24°C

Citations. Author citations in the text must followthe pattern: “Vitt (2000)” or “(Wassersug 1975)”.For two-author publications, both authors must becited separated by “and”; e.g. “Bastos and Haddad(1996)” or “(Zimmerman and Simberloff 1996)”.For more than two authors, only the first one mustbe cited followed by et al. (in italics); e.g. “Heyeret al. (1990)” or “(Inger et al. 1986)”. Severalpublications must be cited in chronological order;e.g. “(Crump 1974, Duellman 1978, 1988,Duellman and Trueb 1986)”. Two or morepublications by the same author must be cited as“(Vanzolini 1991, 1992)” or “(Cadle 1984a, b, c,1985)”. Author names should be separated by“and”, not by &.

References. In the reference list, the citedpublications must be included in alphabetical orderin the following format (“gray literature” must beavoided):

Journal articles with usual volume and issuenumber

Vanzolini, P. E. 1993. A new species of turtle,genus Trachemys, from the state of Maranhão,Brazil (Testudines, Emydidae). Revista Brasi-leira de Biologia 55: 111–125.

Two authors in a journal seriesZamudio, K. R. and H. W. Greene. 1997. Phylo-

geography of the bushmaster (Lachesis muta:Viperidae): implications for Neotropicalbiogeography, systematics, and conservation.Biological Journal of the Linnean Society 62:421–442.

More than two authors in a journal seriesHero, J.-M., W. E. Magnusson, C. F. D. Rocha and

C. P. Catterall. 2001. Antipredator defensesinfluence the distribution of amphibian preyspecies in the Central Amazon rain forest.Biotropica 33: 131–141.

Chapter in an edited volumeHedges, S. B. 1999. Distribution patterns of

amphibians in the West Indies. Pp. 211–254 inW. E. Duellman (ed.), Patterns of Distributionof Amphibians – a global perspective. Balti-more and London. The Johns Hopkins Univer-sity Press.

Unpublished M.Sc. Dissertation or Ph.D. ThesisVerdade, V. K. 2001. Revisão das espécies de

Colostethus Cope, 1866 da Mata Atlântica(Anura, Dendrobatidae). Unpublished M.Sc.Dissertation. Universidade de São Paulo, Brazil.

BookMcDiarmid, R. W. and R. Altig (eds.). 1999.

Tadpoles – the biology of anuran larvae. Chi-cago and London. The University of ChicagoPress. 633 pp.

Material from the World Wide WebFrost, D. R. (ed.). 2000. Amphibian Species of the

World – an online reference . URL: http://reaserch.amnh.org/herpetology/amphibia/index.html. Captured on 23 December 2002.

Animal Care and License for Collection:PHYLLOMEDUSA editors feel that animals used inresearch be treated humanely and ethically, and allcontributors to PHYLLOMEDUSA are expected tocomply with this feeling. In addition, we requirethat all required state and federal permits havebeen obtained (IBAMA license for Brazil). Thenumber of the license must be included in theAcknowledgements.

Proofs. The publisher will undertake proofreading.Unless specifically requested when the paper issubmitted, proofs will not be sent to authors.

Reprints. Authors will receive 30 reprints free ofcharge.

Submission . Manuscripts must be sent inelectronic format (.doc) to the Editor via e-mailbertoluc@esalq.usp.bror through the homepagewww.phyllomedusa.esalq.usp.br

or by mail (diskette or CD-ROM) to

Jaime BertoluciDepartamento de Ciências BiológicasESALQ – USPAv. Pádua Dias, 11 Caixa Postal 913418-900 Piracicaba – SP BRAZIL

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