The genus Hypsiboas is composed of 90 species distributed in Central and South America from Nicaragua to Argentina, as well as on Trinidad and Tobago (Frost 2014). Previous studies on the reproductive biology of species in this genus have reported that female mature oocytes or eggs generally have pigmented animal poles (Tab. 1), which has been generally suggested as a protection of the exposed embryos from solar radiation (Wells 2007).Fil: Nali, Renato C.. Universidade Estadual Paulista Julio de Mesquita Filho; BrasilFil: Faivovich, Julián. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales "Bernardino Rivadavia"; ArgentinaFil: Prado, Cynthia P. A.. Universidade Estadual Paulista Julio de Mesquita Filho; Brasi

Faivovich, Julián

Nali, Renato C.

Prado, Cynthia P. A.

Faivovich, Julian

Acervo Digital da Unesp

The occurrence of unpigmented mature oocytes in Hypsiboas (Anura: Hylidae)

CONICET Digital

53CorrespondenceAll articles available online at http://www.salamandra-journal.com© 2014 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, GermanyCorrespondenceSALAMANDRA 50(1) 53–56 30 April 2014 ISSN 0036–3375The occurrence of unpigmented mature oocytes in Hypsiboas (Anura: Hylidae) Renato C. Nali¹, Julián Faivovich² & Cynthia P. A. Prado³¹) Programa de Pós-graduação em Ciências Biológicas (Zoologia), Universidade Estadual Paulista “Júlio de Mesquita Filho”, Rio Claro, São Paulo, Brazil ²) División Herpetología, Museo Argentino de Ciencias Naturales-CONICET, Ángel Gallardo 470, C1405DJR, Buenos Aires, Argentina & Departamento de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales,  Universidad de Buenos Aires³) Departamento de Morfologia e Fisiologia Animal, FCAV, Universidade Estadual Paulista “Júlio de Mesquita Filho”, Jaboticabal, São Paulo, BrazilCorresponding author: Renato C. Nali, e-mail: r_nali@yahoo.com.brManuscript received: 25 October 2013Accepted: 10 December 2013 by Jörn KöhlerThe genus Hypsiboas is composed of 90 species distribut-ed in Central and South America from Nicaragua to Ar-gentina, as well as on Trinidad and Tobago (Frost 2014). Previous studies on the reproductive biology of species in this genus have reported that female mature oocytes or eggs generally have pigmented animal poles (Tab. 1), which has been generally suggested as a protection of the exposed embryos from solar radiation (Wells 2007). The Hispaniolan green treefrog, Hypsiboas heilprini (Noble, 1923) (Fig. 1A) is a vulnerable species assigned to the Hypsiboas albopunctatus group (Hedges et al. 2004, Faivovich et al. 2005), and occurs in the Dominican Re-public and Haiti (Hispaniola). While its tadpole was de-scribed by Noble (1927), its reproductive biology remains largely unknown even after 90 years since its original de-scription (Schwartz & Henderson 1991, Landestoy 2013). Herein, we report on the analysis of the ovaries of a gravid female of Hypsiboas heilprini, collected in Haiti in October 2007 and deposited as number CUMV 15991 at the Cornell University Museum of Vertebrates, Ithaca, NY, USA. The ovaries contained 332 unpigmented mature oocytes (Fig. 1B) with a mean diameter of 2.38 ± 0.10 mm (range = 2.19 – 2.51 mm; N = 25 oocytes), as measured un-der a Leica stereomicroscope with photo-capturing soft-ware (Leica Application Suite).The presence of unpigmented oocytes is a rare charac-ter state for the genus Hypsiboas. Currently, the only oth-er species reported to have them are H. lemai (Duellman 1997), H. nympha, and H. roraima (Faivovich et al. 2006; Tab. 1) from the Hypsiboas benitezi group (Faivovich et al., 2005), with stream-associated reproduction. Hypsiboas is included in the well-supported clade Copho mantini (Faivovich et al. 2005). In this clade un-pigmented oocytes or eggs are known to occur in Aplasto­discus, Hyloscirtus, and most Myersiohyla (Faivovich et al. 2013). However, with the exception of Aplasto discus (Hart-mann et al. 2004, Haddad et al. 2005, Zina & Haddad 2007) and a species of Hyloscirtus (H. platy dacty lus, La Marca 1985), little is known about their reproductive modes. In Aplastodiscus, the eggs are deposited inside sub-terranean nests constructed by the males in the muddy side of streams and ponds, where larvae spend the early stages of their development (Hartmann et al. 2004, Haddad et al. 2005, Zina & Haddad 2007). In Hylo scirtus platydacty­lus, the unpigmented eggs are deposited on leaves and ma-ternal care might be present (La Marca 1985). Among the few Hypsiboas known to have unpigmented eggs, observations for H. lemai suggest that eggs are nor-mally deposited on the vegetation overhanging streams, from where the tadpoles would drop and develop in the water (Duellman 1997). For H. heilprini, previous stud-ies and our observations indicate that eggs are deposit-ed in concealed and protected places, such as the wall of a streamside cave or other natural caves (Schwartz & Henderson 1991, Landestoy 2013). Considering the lim-ited knowledge on the diversity of the reproductive bio-lo gy of other Cophomantini with unpigmented eggs, the record of this species depositing its eggs in hidden places is 54Correspondencereminiscent of the reproductive strategy of Aplasto discus, although in the latter nests are generally constructed by males (e.g. Haddad et al. 2005). Future observations will hopefully provide more details. Hypsiboas heilprini has been consistently recovered as the sister taxon of all other species of the H. albopuncta­tus group (Faivovich et al. 2005, 2013, Wiens et al. 2006, 2010, Pyron & Wiens 2011) although usually with low sup-port values. Its phylogenetic relationships indicate that un-pigmented eggs evolved independently from those present in species of the H. benitezi group.Table 1. Occurrence of egg/oocyte pigmentation in species of the genus Hypsiboas. Species Pigmented animal poleReferenceH. heilprini – This studyH. lemai – Duellman 1997H. nympha – Faivovich et al. 2006H. roraima – Faivovich et al. 2006H. aguilari + Lehr et al. 2010H. albomarginatus + L. O. M. Giasson, pers. comm.H. albopunctatus + Muniz et al. 2008H. atlanticus + Camurugi & Juncá 2013H. bischoffi + Haddad 1991H. boans + Duellman 1978, Burger et al. 2002H. caipora + Antunes 2007H. calcaratus + Duellman 1978H. cinerascens + Duellman 1978, Telles et al. 2013H. cipoensis + Eterovick et al. 2002H. cordobae + Fernandez 1927H. crepitans + Hoogmoed & Gorzula 1979H. curupi + Garcia et al. 2007H. faber + Martins et al. 1998H. fasciatus + Duellman 1978H. geographicus + Kenny 1969, Duellman 1978, Lima et al. 2006H. goianus + Menin et al. 2004H. hutchinsi + Pyburn & Hall 1984H. joaquini + Kwet et al. 2010H. lanciformis + Duellman 1978H. latistriatus + Orrico et al. 2007H. leptolineatus + Kwet et al. 2010H. lundii + C. S. Ribeiro, pers. comm.H. marginatus + Kwet et al. 2010H. marianitae + Duellman et al. 1997H. picturatus + E. E. Tapia & L. A. Coloma, pers. comm.; Hoogmoed 1979H. poaju + Garcia et al. 2008H. polytaenius + Lutz 1973H. prasinus + Haddad 1991H. pulchellus + Barrio 1962H. punctatus + Kenny 1969, Duellman 1978H. raniceps + C. P. A. Prado, pers. obs.H. riojanus + Caruso 1949H. rosenbergi + Kluge 1981H. rufitelus + Duellman 1970H. sibleszi + Hoogmoed 1979H. wavrini + Martins & Moreira 199155CorrespondenceAcknowledgementsThe authors are grateful to L.O.M. Giasson, C.S. Ribeiro, E.E. Tapia and L.A. Coloma for their personal communications, and P.D.P. Pinheiro for his comments.. R.C. Nali thanks the São Paulo Research Foundation (FAPESP proc. 2012/06228-0) for a doctoral fellowship and C.P.A. Prado acknowledges the CNPq (proc. 471106/2010-0) and São Paulo Research Founda-tion (FAPESP proc. 2009/12013-4) for their financial support. J. Faivovich thanks ANPCyT 2007-2202, 2011-1895, UBACyT 20020090200727, PIP CONICET 11220110100889, and São Paulo Research Foundation (FAPESP proc. 2012/10000-5). The authors are also grateful to H.W. Greene for the photograph of Hypsiboas heilprini.ReferencesAntunes, A. P. (2007): Descrição taxonômica e história natural de uma nova espécie de Hypsiboas do alto da Serra de Para-napiacaba, estado de São Paulo. – Unpublished Master’s Dis-sertation (in Portuguese), Universidade Estadual Paulista, SP, Brazil.Barrio, A. (1962): Los Hylidae de Punta Lara, provincia de Bue-nos Aires. – Physis, 23: 129–142.Burger, J., W. Arizabal & M. Gochfeld (2002): Nesting be-havior of a gladiator frog Hyla boans in Peru. – Journal of Her-petology, 36: 640–648. Camurugi, F. & F. Juncá (2013): Reproductive biology of Hypsi­boas atlanticus (Anura: Hylidae). – Herpetology Notes, 6: 489–495.Caruso, M. A. (1949): Sobre el ciclo sexual anual de algunos “Hylidae” del norte Argentino (“Phyllomedusa sauvagii” e “Hyla raddiana”). – Acta Zoologica Lilloana, 8: 83–103.Duellman, W. E. (1970): Hylid frogs of Middle America. – Mono graphs of the Museum of Natural History, University of Kansas, 1–2: 1–753.Duellman, W. E. (1978): The biology of an equatorial herpeto-fauna in Amazonian Ecuador. – Miscellaneous Publications of the Museum of Natural History of the University of Kansas, 65: 1–352.Duellman, W. E. (1997): Amphibians of La Escalera region, southeastern Venezuela: taxonomy, ecology, and biogeogra-phy. – Scientiﬁc Papers, Natural History Museum, University of Kansas, 2: 1–52.Duellman, W. E., I. De La Riva & E. R. Wild (1997): Frogs of the Hyla armata and Hyla pulchella groups in the Andes of South America, with definitions and analyses of phylogenetic rela-tionships of Andean groups of Hyla. – Scientific Papers of the Natural History Museum, The University of Kansas, 3: 1–41.Eterovick, P. C., I. S. Barros & I. Sazima (2002): Tadpoles of two species in the Hyla polytaenia species group and compari-son with other tadpoles of Hyla polytaenia and Hyla pulchel­la groups (Anura, Hylidae). – Journal of Herpetology, 36: 512–515.Faivovich, J., C. F. B. Haddad, P. C. A. Garcia, D. R. Frost, J. A. Campbell & W. C. Wheeler (2005): Systematic review of the frog family Hylidae, with special reference to Hylinae: Phylogenetic analysis and taxonomic revision. – Bulletin of the American Museum of Natural History, 294: 1–240.Faivovich, J., J. Moravec, D. F. Cisneros-Heredia & J. Köh-ler (2006): A new species of the Hypsiboas benitezi group from the western Amazon basin (Amphibia: Anura: Hylidae). – Herpetologica, 52: 96–108.Faivovich, J., R. W. McDiarmid & C. W. Myers (2013): Two new species of Myersiohyla (Anura: Hylidae) from Cerro de la Ne-blina, Venezuela, with comments on other species of the ge-nus. – American Museum Novitates, 3792: 63 pp.Fernandez, K. (1927): Sobre la biología y reproducción de batra-cios argentinos (segunda parte). – Boletin de la Academia Na-cional de Ciencias en Córdoba, 29: 271–238.Frost, D. R. (2014): Amphibian Species of the World: an Online Reference. Version 6.0 (25 January 2014). – Electronic Da-tabase accessible at http://research.amnh.org/herpetology/amphibia/index.html. American Museum of Natural History, New York, USA.Garcia, P. C. A., J. Faivovich & C. F. B. Haddad (2007): Rede-scription of Hypsiboas semiguttatus, with the description of a new species of the Hypsiboas pulchellus group. – Copeia, 2007: 933–951.Garcia, P. C. A., O. L. Peixoto & C. F. B. Haddad (2008): A new species of Hypsiboas (Anura: Hylidae) from the Atlantic Forest of Santa Catarina, Southern Brazil, with comments on its conservation status. – South American Journal of Herpeto-logy, 3: 27–35.Figure 1. (A) Hispaniolan green treefrog, Hypsiboas heilprini (photograph by H. W. Greene) and (B) segment from a female’s ovary (accession number CUMV 15991), showing both mature and immature unpigmented oocytes.56CorrespondenceHaddad, C. F. B. (1991): Ecologia reprodutiva de uma comuni-dade de anfíbios anuros na Serra do Japi, Sudeste do Brasil. – Unpublished PhD Thesis (in Portuguese), Universidade Es-tadual de Campinas, SP, Brazil.Haddad, C. F. B., J. Faivovich & P. C. A. Garcia (2005): The specialized reproductive mode of the treefrog Aplastodiscus perviridis (Anura: Hylidae). – Amphibia-Reptilia, 26: 87–92.Hartmann, M. T., P. A. Hartmann & C. F. B. Haddad (2004): Visual signaling and reproductive biology in a nocturnal tree-frog, genus Hyla (Anura: Hylidae). – Amphibia-Reptilia, 25: 395–406.Hedges, B., S. Inchaustegui, M. Hernandez & R. Powell (2004): Hypsiboas heilprini. – in: IUCN 2012: IUCN Red List of Threatened Species. Version 2012.2. – www.iucnredlist.org. Accessed on 11 March 2013.Hoogmoed, M. S. (1979): Resurrection of Hyla ornatissima No-ble (Amphibia, Hylidae) and remarks on related species of green tree frogs from the Guiana area. Notes on the herpeto-fauna of Surinam VI. – Zoologische Verhandelingen 172: 1–46.Hoogmoed, M. S. & S. Gorzula (1979): Checklist of the savan-na inhabiting frogs of the El Manteco region with notes on their ecology and the description of a new species of treefrog. – Zoo logische Mededelingen, 54: 183–216.Kenny, J. S. (1969): The Amphibia of Trinidad. – Studies on the fauna of Curaçao and other Caribbean islands, 108: 1–78.Kluge, A. G. (1981): The life history, social organization, and parental behavior of Hyla rosenbergi Boulenger, a nestbuild-ing gladiator frog. – Miscellaneous Publications University of Michigan, Museum of Zoology, 160: 1–170.Kwet, A., R. Lingnau & M. Di-Bernardo (2010): Anfíbios da Serra Gaucha Sul do Brasil. – Tübingen: Brasilien Zentrum der Universität Tübingen.La Marca, E. (1985): Systematics and ecological observations on the Neotropical frogs Hyla jahni and Hyla platydactyla. – Jour-nal of Herpetology, 19: 227– 237.Landestoy, M. A. (2013): Observations on the breeding behavior of the Hispaniolan green treefrog, Hypsiboas heilprini. – IRCF Reptiles & Amphibians, 20: 160–165.Lehr, E., J. Faivovich & K.-H. Jungfer (2010): A new Andean species of the Hypsiboas pulchellus group: Adults, calls, and phylogenetic relationships. – Herpetologica, 66: 296–307.Lima, A. P., W. E. Magnusson, M. Menin, L. K. Erdtmann, D. J. Rodrigues, C. Keller & W. Hödl (2006): Guia de sapos da Reserva Adolph Ducke – Amazônia Central. – Manaus: Edi-tora Attema. Lutz, B. (1973): Brazilian species of Hyla. – Austin: University of Texas Press. Martins, M. & G. Moreira (1991): The nest and the tadpole of Hyla wavrini  Parker (Amphibia, Anura).  – Memórias do Instituto Butantan, São Paulo, 53: 197–204.Martins, M., J. P. J. Pombal, & C. F. B. Haddad (1998): Esca-lated aggressive behaviour and facultative parental care in the nest building gladiator frog, Hyla faber. – Amphibia-Reptilia, 19: 65–73.Menin, M., R. A. Silva & A. A. Giaretta (2004): Reproductive biology of Hyla goiana  (Anura, Hylidae).  – Iheringia, Série Zoologia, 94: 49–52.Muniz, K. P. R., A. A. Giaretta, W. R. Silva & K. G. Facure (2008): Autoecologia de Hypsiboas albopunctatus (Anura, Hylidae) em área de Cerrado no sudeste do Brasil. – Iheringia, Série Zoologia, 98: 254–259.Noble, G. K. (1927): The value of life history data in the study of evolution of the Amphibia. – Annals of the New York Acad-emy of Sciences, 30: 31–128.Orrico, V. G. D., M. M. Mongin & A. M. F. T. de Carvalho-e-Silva (2007): The tadpole of Hypsiboas latistriatus (Caramas-chi and Cruz, 2004), a species of the Hypsiboas polytaenius (Cope, 1870) clade (Amphibia, Anura, Hylidae). – Zootaxa, 1531: 25–37.Pyburn, W. F. & D. H. Hall (1984): A new stream-inhabiting treefrog (Anura: Hylidae) from southeastern Colombia. – Herpetologica, 40: 366–372.Pyron, R. A. & J. J. Wiens (2011): A large-scale phylogeny of Am-phibia including over 2,800 species, and a revised classifica-tion of extant frogs, salamanders, and caecilians. – Molecular Phylogenetics and Evolution, 61: 543–583.Schwartz, A. & R. W. Henderson (1991): Amphibians and Rep-tiles of the West Indies: Descriptions, Distributions and Natu-ral History. – Florida: The University of Florida Press.Telles, D.O.C., S. A. F. Vaz & M. Menin (2013): Reproductive bio logy, size and diet of Hypsiboas cinerascens (Anura: Hyli-dae) in two urban forest fragments in Central Amazonia, Bra-zil. – Phyllomedusa, 12: 69–76.Wells, K. D. (2007): The ecology and behavior of amphibians. – Chicago: The University of Chicago Press.Wiens, J. J., C. H. Graham, D. S. Moen, S. A. Smith & T. W. Reeder (2006): Evolutionary and ecological causes of the lati-tudinal diversity gradient in hylid frogs: treefrog tree unearth the roots of high tropical diversity. – The American Natural-ist, 168: 579–596.Wiens, J. J., C. A. Kuczynski, X. Hua & D. S. Moen (2010): An expanded phylogeny of treefrogs (Hylidae) based on nuclear and mitochondrial sequence data. – Molecular Phylogenetics and Evolution, 55: 871–882.Zina, J. & C. F. B. Haddad (2007): Courtship behavior of two treefrog species, Aplastodiscus arildae and A. leucopygius (An-ura: Hylidae), from the Atlantic rainforest, southeastern Bra-zil. – Herpetological Review, 38: 282–285.

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53CorrespondenceAll articles available online at http://www.salamandra-journal.com© 2014 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, GermanyCorrespondenceSALAMANDRA 50(1) 53–56 30 April 2014 ISSN 0036–3375The occurrence of unpigmented mature oocytes in Hypsiboas (Anura: Hylidae) Renato C. Nali¹, Julián Faivovich² & Cynthia P. A. Prado³¹) Programa de Pós-graduação em Ciências Biológicas (Zoologia), Universidade Estadual Paulista “Júlio de Mesquita Filho”, Rio Claro, São Paulo, Brazil ²) División Herpetología, Museo Argentino de Ciencias Naturales-CONICET, Ángel Gallardo 470, C1405DJR, Buenos Aires, Argentina & Departamento de Biodiversidad y Biología Experimental, Facultad de Ciencias Exactas y Naturales,  Universidad de Buenos Aires³) Departamento de Morfologia e Fisiologia Animal, FCAV, Universidade Estadual Paulista “Júlio de Mesquita Filho”, Jaboticabal, São Paulo, BrazilCorresponding author: Renato C. Nali, e-mail: r_nali@yahoo.com.brManuscript received: 25 October 2013Accepted: 10 December 2013 by Jörn KöhlerThe genus Hypsiboas is composed of 90 species distribut-ed in Central and South America from Nicaragua to Ar-gentina, as well as on Trinidad and Tobago (Frost 2014). Previous studies on the reproductive biology of species in this genus have reported that female mature oocytes or eggs generally have pigmented animal poles (Tab. 1), which has been generally suggested as a protection of the exposed embryos from solar radiation (Wells 2007). The Hispaniolan green treefrog, Hypsiboas heilprini (Noble, 1923) (Fig. 1A) is a vulnerable species assigned to the Hypsiboas albopunctatus group (Hedges et al. 2004, Faivovich et al. 2005), and occurs in the Dominican Re-public and Haiti (Hispaniola). While its tadpole was de-scribed by Noble (1927), its reproductive biology remains largely unknown even after 90 years since its original de-scription (Schwartz & Henderson 1991, Landestoy 2013). Herein, we report on the analysis of the ovaries of a gravid female of Hypsiboas heilprini, collected in Haiti in October 2007 and deposited as number CUMV 15991 at the Cornell University Museum of Vertebrates, Ithaca, NY, USA. The ovaries contained 332 unpigmented mature oocytes (Fig. 1B) with a mean diameter of 2.38 ± 0.10 mm (range = 2.19 – 2.51 mm; N = 25 oocytes), as measured un-der a Leica stereomicroscope with photo-capturing soft-ware (Leica Application Suite).The presence of unpigmented oocytes is a rare charac-ter state for the genus Hypsiboas. Currently, the only oth-er species reported to have them are H. lemai (Duellman 1997), H. nympha, and H. roraima (Faivovich et al. 2006; Tab. 1) from the Hypsiboas benitezi group (Faivovich et al., 2005), with stream-associated reproduction. Hypsiboas is included in the well-supported clade Copho mantini (Faivovich et al. 2005). In this clade un-pigmented oocytes or eggs are known to occur in Aplasto­discus, Hyloscirtus, and most Myersiohyla (Faivovich et al. 2013). However, with the exception of Aplasto discus (Hart-mann et al. 2004, Haddad et al. 2005, Zina & Haddad 2007) and a species of Hyloscirtus (H. platy dacty lus, La Marca 1985), little is known about their reproductive modes. In Aplastodiscus, the eggs are deposited inside sub-terranean nests constructed by the males in the muddy side of streams and ponds, where larvae spend the early stages of their development (Hartmann et al. 2004, Haddad et al. 2005, Zina & Haddad 2007). In Hylo scirtus platydacty­lus, the unpigmented eggs are deposited on leaves and ma-ternal care might be present (La Marca 1985). Among the few Hypsiboas known to have unpigmented eggs, observations for H. lemai suggest that eggs are nor-mally deposited on the vegetation overhanging streams, from where the tadpoles would drop and develop in the water (Duellman 1997). For H. heilprini, previous stud-ies and our observations indicate that eggs are deposit-ed in concealed and protected places, such as the wall of a streamside cave or other natural caves (Schwartz & Henderson 1991, Landestoy 2013). Considering the lim-ited knowledge on the diversity of the reproductive bio-lo gy of other Cophomantini with unpigmented eggs, the record of this species depositing its eggs in hidden places is 54Correspondencereminiscent of the reproductive strategy of Aplasto discus, although in the latter nests are generally constructed by males (e.g. Haddad et al. 2005). Future observations will hopefully provide more details. Hypsiboas heilprini has been consistently recovered as the sister taxon of all other species of the H. albopuncta­tus group (Faivovich et al. 2005, 2013, Wiens et al. 2006, 2010, Pyron & Wiens 2011) although usually with low sup-port values. Its phylogenetic relationships indicate that un-pigmented eggs evolved independently from those present in species of the H. benitezi group.Table 1. Occurrence of egg/oocyte pigmentation in species of the genus Hypsiboas. Species Pigmented animal poleReferenceH. heilprini – This studyH. lemai – Duellman 1997H. nympha – Faivovich et al. 2006H. roraima – Faivovich et al. 2006H. aguilari + Lehr et al. 2010H. albomarginatus + L. O. M. Giasson, pers. comm.H. albopunctatus + Muniz et al. 2008H. atlanticus + Camurugi & Juncá 2013H. bischoffi + Haddad 1991H. boans + Duellman 1978, Burger et al. 2002H. caipora + Antunes 2007H. calcaratus + Duellman 1978H. cinerascens + Duellman 1978, Telles et al. 2013H. cipoensis + Eterovick et al. 2002H. cordobae + Fernandez 1927H. crepitans + Hoogmoed & Gorzula 1979H. curupi + Garcia et al. 2007H. faber + Martins et al. 1998H. fasciatus + Duellman 1978H. geographicus + Kenny 1969, Duellman 1978, Lima et al. 2006H. goianus + Menin et al. 2004H. hutchinsi + Pyburn & Hall 1984H. joaquini + Kwet et al. 2010H. lanciformis + Duellman 1978H. latistriatus + Orrico et al. 2007H. leptolineatus + Kwet et al. 2010H. lundii + C. S. Ribeiro, pers. comm.H. marginatus + Kwet et al. 2010H. marianitae + Duellman et al. 1997H. picturatus + E. E. Tapia & L. A. Coloma, pers. comm.; Hoogmoed 1979H. poaju + Garcia et al. 2008H. polytaenius + Lutz 1973H. prasinus + Haddad 1991H. pulchellus + Barrio 1962H. punctatus + Kenny 1969, Duellman 1978H. raniceps + C. P. A. Prado, pers. obs.H. riojanus + Caruso 1949H. rosenbergi + Kluge 1981H. rufitelus + Duellman 1970H. sibleszi + Hoogmoed 1979H. wavrini + Martins & Moreira 199155CorrespondenceAcknowledgementsThe authors are grateful to L.O.M. Giasson, C.S. Ribeiro, E.E. Tapia and L.A. Coloma for their personal communications, and P.D.P. Pinheiro for his comments.. R.C. Nali thanks the São Paulo Research Foundation (FAPESP proc. 2012/06228-0) for a doctoral fellowship and C.P.A. Prado acknowledges the CNPq (proc. 471106/2010-0) and São Paulo Research Founda-tion (FAPESP proc. 2009/12013-4) for their financial support. J. Faivovich thanks ANPCyT 2007-2202, 2011-1895, UBACyT 20020090200727, PIP CONICET 11220110100889, and São Paulo Research Foundation (FAPESP proc. 2012/10000-5). The authors are also grateful to H.W. Greene for the photograph of Hypsiboas heilprini.ReferencesAntunes, A. P. (2007): Descrição taxonômica e história natural de uma nova espécie de Hypsiboas do alto da Serra de Para-napiacaba, estado de São Paulo. – Unpublished Master’s Dis-sertation (in Portuguese), Universidade Estadual Paulista, SP, Brazil.Barrio, A. (1962): Los Hylidae de Punta Lara, provincia de Bue-nos Aires. – Physis, 23: 129–142.Burger, J., W. Arizabal & M. Gochfeld (2002): Nesting be-havior of a gladiator frog Hyla boans in Peru. – Journal of Her-petology, 36: 640–648. Camurugi, F. & F. Juncá (2013): Reproductive biology of Hypsi­boas atlanticus (Anura: Hylidae). – Herpetology Notes, 6: 489–495.Caruso, M. A. (1949): Sobre el ciclo sexual anual de algunos “Hylidae” del norte Argentino (“Phyllomedusa sauvagii” e “Hyla raddiana”). – Acta Zoologica Lilloana, 8: 83–103.Duellman, W. E. (1970): Hylid frogs of Middle America. – Mono graphs of the Museum of Natural History, University of Kansas, 1–2: 1–753.Duellman, W. E. (1978): The biology of an equatorial herpeto-fauna in Amazonian Ecuador. – Miscellaneous Publications of the Museum of Natural History of the University of Kansas, 65: 1–352.Duellman, W. E. (1997): Amphibians of La Escalera region, southeastern Venezuela: taxonomy, ecology, and biogeogra-phy. – Scientific Papers, Natural History Museum, University of Kansas, 2: 1–52.Duellman, W. E., I. De La Riva & E. R. Wild (1997): Frogs of the Hyla armata and Hyla pulchella groups in the Andes of South America, with definitions and analyses of phylogenetic rela-tionships of Andean groups of Hyla. – Scientific Papers of the Natural History Museum, The University of Kansas, 3: 1–41.Eterovick, P. C., I. S. Barros & I. 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The occurrence of unpigmented mature oocytes in Hypsiboas (Anura: Hylidae)

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