Anura: Hylidae) Author(S): Victor G
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The Taxonomic Status of Dendropsophus allenorum and Dendropsophus timbeba (Anura: Hylidae) Author(s): Victor G. D. Orrico , William E. Duellman , Moisés B. Souza , and Célio F. B. Haddad Source: Journal of Herpetology, 47(4):615-618. 2013. Published By: The Society for the Study of Amphibians and Reptiles DOI: http://dx.doi.org/10.1670/12-208 URL: http://www.bioone.org/doi/full/10.1670/12-208 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Journal of Herpetology, Vol. 47, No. 4, 615–618, 2013 Copyright 2013 Society for the Study of Amphibians and Reptiles The Taxonomic Status of Dendropsophus allenorum and Dendropsophus timbeba (Anura: Hylidae) 1,2 3 4 1 VICTOR G. D. ORRICO, WILLIAM E. DUELLMAN, MOISE´ S B. SOUZA, AND CE´ LIO F. B. HADDAD 1Departamento de Zoologia, I.B., Universidade Estadual Paulista Ju´lio de Mesquita Filho, Avenida 24-A, nu´mero 1515, Bela Vista, CEP: 13506-900, Claro, Sa˜o Paulo, Brazil 3Biodiversity Institute, University of Kansas, 1345 Jayhawk Boulevard, Lawrence, Kansas 66045 USA 4Centro de Cieˆncias Biolo´gicas e da Natureza, Universidade Federal do Acre, Rodovia BR-364, km 04, Distrito Industrial, CEP: 69915-900, Branco, Acre, Brazil ABSTRACT.—Careful examination of morphological characters, color patterns, and advertisement calls strongly supports the placement of Dendropsophus allenorum (Duellman and Trueb, 1989) (Cuzco Reserve Frog) into the synonymy of Dendropsophus timbeba (Martins and Cardoso, 1987) (Cardoso’s Treefrog). RESUMO.—Um exame cuidadoso dos caracteres morfolo´gicos, dos padro˜es de colorac¸a˜o e dos cantos de anu´ncio suportam fortemente que Dendropsophus allenorum (Duellman e Trueb, 1989) deva ser considerado um sinoˆnimo ju´nior de Dendropsophus timbeba (Martins e Cardoso, 1987). For more than 4 decades, small species of Hyla Laurenti (1768) 1987), and a species known only from British Guiana, from South and Central America, known or suspected to have a Dendropsophus grandisonae (Goin, 1966) (Mazaruni Treefrog) diploid number of 30 chromosomes, were believed to constitute (Lutz, 1973; Frost, 2013). Commonly, museum specimens placed a monophyletic group (Duellman and Cole, 1965; Bogart, 1973; in this species group are only tentatively assigned (e.g., Duellman and Trueb, 1983). Recently, Faivovich et al. (2005) Dendropsophus cf. brevifrons from French Guiana; Fouquet et resurrected the genus Dendropsophus Fitzinger, 1843, to accom- al., 2011). Therefore, the taxonomic status of many species modate these species. assigned to this group is uncertain. Dendropsophus is second only to Scinax Wagler, 1830 as the A recent taxonomic change in the D. parviceps group by Souza most speciose genus of Hylidae (Frost, 2013) and, because of the (2009) is the placement of D. allenorum (Duellman and Trueb, large amount of intra- and interspecific variation of its species, it 1989) as a junior synonym of D. timbeba (Martins and Cardoso, has been considered as ‘‘Probably the most taxonomically 1987). However, this author provided no evidence for this frustrating complex of hylid frogs in the American tropics ...’’ taxonomic change. Consequently, this action is little known and (Duellman, 1982:153). Species of this genus are highly variable has not yet been accepted widely (e.g., as of 24 January 2013, in morphology (e.g., Duellman and Fouquette, 1968; Franco- Frost, 2013 still recognizes D. allenorum as valid). Herein we Belussi et al., 2011), advertisement call (e.g., Martins and Jim, provide evidence for synonymizing these species. 2003, 2004), chromosome morphology (e.g., Gruber et al., 2005), and reproductive behavior (e.g., Touchon and Warkentin, 2008). MATERIALS AND METHODS Despite some advances in the understanding of intrageneric relationships (e.g., Faivovich et al., 2005; Wiens, 2006; Wiens et We compared the type series of D. allenorum and D. timbeba al., 2010), smaller-scale relationships are still unresolved. using the diagnostic characters employed by Martins and Faivovich et al. (2005) recognized nine species groups of Cardoso (1987) and Duellman and Trueb (1989). Museum Dendropsophus. One of these is the Dendropsophus parviceps abbreviations follow the Standard Symbolic Codes for Institu- (Sarayacu Treefrog) species group, first defined by Duellman tional Resource Collections in Herpetology and Ichthyology and Crump (1974). Currently its monophyly is regarded with (Sabaj Pe´rez, 2010). KUDA stands for the Digital Archives skepticism (Faivovich et al., 2005; Fouquet et al., 2011; Motta et Collection of the Division of Herpetology of the Biodiversity al., 2012) and relationships within the group are unclear. Institute of the University of Kansas. Recently,Fouquetetal.(2011)transferredDendropsophus gaucheri (Lescure and Marty, 2000), a species assigned to the RESULTS AND DISCUSSION D. parviceps group, to the Dendropsophus microcephalus (Cope, Duellman and Trueb (1989) stated, ‘‘The coloration immedi- 1886) (Small-headed Treefrog) group; their molecular evidence ately distinguishes Hyla allenorum from all other members of the emphasized the paraphyly of the D. parviceps group and the group, as well as from all other hylid frogs in the Amazon weakly supported relationships within the genus. Basin... H. timbeba, which also has blue flecks on the venter, The D. parviceps species group contains 15 species (Motta et differs by having dark transverse markings on the dorsum and al., 2012) and is distributed mainly throughout the Amazon lacking a lateral white stripe.’’ Nevertheless, Martins and Basin where 11 of the species occur. Exceptions are the Atlantic Cardoso (1987:550) stated that [translated from Portuguese] Forest endemics Dendropsophus giesleri (Mertens, 1950) (Giesler’s ‘‘...hind- and forelimbs with dark markings dorsally; lateral Treefrog), Dendropsophus microps (Peters, 1872) (Nova Friburgo longitudinal white stripes on the axillae and extending onto the Treefrog), and Dendropsophus ruschii (Weygoldt and Peixoto, sides of the body...’’ Some paragraphs later, Martins and 2 Cardoso (1987) noted that some individuals lack the lateral Corresponding Author. Present address: Departamento de longitudinal white stripes. Zoologia, I.B., Universidade de Sa˜o Paulo, Sa˜o Paulo, CEP: 05508- 090, Sa˜o Paulo, Brazil.. E-mail: [email protected] The absence of the lateral white stripe in D. timbeba was DOI: 10.1670/12-208 probably considered a significant diagnostic character because it 616 V. G. D. ORRICO ET AL. FIG. 1. (A) Lateral view of MZUSP 60560, the holotype of Hyla timbeba Martins and Cardoso, 1987; Bar = 4 mm. (B) A female paratype of Hyla timbeba Martins and Cardoso, 1987; (B1) dorsal, (B2) ventral, and (B3) lateral views of ZUEC 5723 (snout–vent length = 22.63 mm). (C) Dorsal views of two paratypes of Hyla timbeba Martins and Cardoso, 1987; (C1) CFBH 6220 (ex-ZUEC 5476) and (C2) ZUEC 5473; Bar = 4 mm. Figure lettering is located at left and above the corresponding figure. TAXONOMIC STATUS OF DENDROPSOPHUS ALLENORUM 617 FIG. 2. (A) Paratype of Hyla allenorum Duellman and Trueb, 1989 (KU 207607, male, SVL = 18.3 mm). (B) Topotype of Hyla allenorum (KU 215189, male, SVL = 18.3 mm). was used to distinguish two other species in the D. parviceps pulses per note (10 for D. timbeba whereas D. allenorum group. Dendropsophus microps has a broad lateral stripe, similar sometimes have 11; see Duellman, 2005) are similar. There is to that in D. allenorum; D. giesleri lacks such a lateral stripe some distinction in note repetition rate (50 to 60 notes/min in D. (Heyer, 1980). Upon examining the type series of D. timbeba,itis timbeba and about 40 notes/min in D. allenorum). However, clear that the description by Martins and Cardoso (1987) is Duellman (2005) commented that although a rate of 40 notes/ correct and is based on the complete type series; the presence of min was typical, higher rates did occur. In D. allenorum the note the lateral white stripe is variable. The lateral longitudinal white duration is 400–450 ms (Duellman, 2005) while in D. timbeba is stripe is not evident in the illustrations published by Martins 600 ms (Martins and Cardoso, 1987). Nevertheless, Martins and and Cardoso (1987). Nevertheless, the holotype (MZUSP 60560) Jim (2003) demonstrated that temporal parameters of the has a white lateral stripe (Fig. 1A). advertisement calls of some species of Dendropsophus are highly The holotype of D. allenorum (KU 207606) is a female with a influenced by social context. distinct lateral stripe that begins anteriorly at the lip and A Note on the Type Series of Dendropsophus timbeba.—Martins extends posteriorly to the groin. This stripe can be interpreted as and Cardoso (1987) reported that the museum number of the a long suborbital bar, similar to the bars cited by Duellman and holotype of D. timbeba was MZUSP 60550. However, the Crump (1974) for other species of the group. A female paratype specimen associated with this number is a Leptodactylus latrans of D. timbeba (ZUEC 5723; Fig. 1B), collected with the male (Steffen, 1815) and is part of another series of individuals holotype, is identical in terms of color and pattern to the collected elsewhere. The holotype is the only specimen of the holotype of D. allenorum. Male paratypes of D. allenorum have type series of D. timbeba housed in MZUSP; all others remained less-extensive lateral stripes.