Anura: Hylidae) Author(S): Ulisses Caramaschi and Marcelo F
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Nomenclatural Status of the Synonyms of Hyla pardalis , and Taxonomic Position of Hyla biobeba (Anura: Hylidae) Author(s): Ulisses Caramaschi and Marcelo F. Napoli Source: Journal of Herpetology, 38(4):501-509. 2004. Published By: The Society for the Study of Amphibians and Reptiles DOI: http://dx.doi.org/10.1670/211-02A URL: http://www.bioone.org/doi/full/10.1670/211-02A 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. 38, No. 4, pp. 487–494, 2004 Copyright 2004 Society for the Study of Amphibians and Reptiles Taxonomy of Costa Rican Toads Referred to Bufo melanochlorus Cope, with the Description of a New Species 1 ERIC M. O’NEILL AND JOSEPH R. MENDELSON III Department of Biology, Utah State University, Logan, Utah 84322-5305, USA ABSTRACT.—We review the taxonomic status of populations of toads referred to Bufo melanochlorus Cope, 1877, that occur in the wet forests on both Atlantic and Pacific versants of Costa Rica. Populations from Pacific versant of Costa Rica and adjacent Panama are qualitatively diagnosable from all other populations and are described herein as a new species. The taxon B. melanochlorus is restricted to populations in the Atlantic and Montane Slopes and Cordillera Central faunal areas of Costa Rica. RESUMEN.—Revisamos el estado taxono´mico de las poblaciones de los sapos referidos a Bufo melanochlorus Cope, 1887. Estos ocurren en los bosques me´sicos de la vertiente del Atla´ntico y la vertiente del Pacı´fico de Costa Rica y Panama´. Las poblaciones de la vertiente del Pacı´fico de Costa Rica y de Panama´ se pueden diagnosticar de todas las dema´s poblaciones y aquı´ se describen como una nueva especie. El taxo´n B. melanochlorus esta´ restringido a las poblaciones de las a´reas faunisticas de Costa Rica como el Atla´ntico, Vertientes Montan˜ osos y la Cordillera Central. Populations of toads referred to Bufo melano- The following morphological measurements chlorus Cope, 1877 are found in wet forests on were taken from adult specimens: snout–vent both Atlantic and Pacific versants of Costa Rica length (SVL), head length (HL), head width (Savage, 2002). Although there are ample series of (HW), tibia length (TIB), tarsus length (TAR) foot specimens from the Southwest faunal area (sensu length (FL), width of tympanum (TYM), length of Savage, 2002:fig 15), populations in Atlantic, parotoid gland (PARL), maximum width of Montane Slopes and Cordillera Central, and parotoid gland (PARW), length of supratym- Pacific Northwest faunal areas are documented panic crest (SPTYM), and length of postorbital by relatively small series of specimens that crest (PORB). These variables, which represent consist mostly of juveniles and subadults. As repeatable morphological landmarks, were mea- part of continued efforts to revise the taxonomy sured with digital calipers, rounded to nearest 0.1 of Mesoamerican species of Bufo—so as to better mm. Sex of individuals was determined by reflect actual biodiversity in this region—we have presence/absence of male secondary sexual examined series of specimens from throughout characters (nuptial excresences and/or vocal the geographic range of B. melanochlorus. slits), or by direct examination of gonads. We compared means between sexes and MATERIALS AND METHODS among species of 66 adult males and 39 adult General terminology and format for diagnoses females using ANCOVA, for all variables with and description follow that of Mendelson (1997). SVL as the covariate. We compared mean SVL Foot-webbing formulae follow the system sum- using a t-test. Statistical analyses were performed marized by Savage (2002). Museum abbrevia- using SAS (SAS Institute, vers. 8). Because of tions are those proposed by Leviton et al. (1985), limited availability of museum specimens of with the following addition: UCR for the Museo adult B. melanochlorus, our samples for this study de Zoologı´a, Escuela de Biologı´a, Universidad de are heavily skewed (especially for males). Nev- Costa Rica, Ciudad Universitaria Rodrigo Facio, ertheless, we used morphometric analyses to San Jose´, Costa Rica. We follow Tyler et al. (2001) assess variation in body shape. These analyses in our usage of the term ‘‘parotoid gland.’’ should be taken as preliminary; more thorough We examined specimens for fixed morpholog- sampling will be required before robust conclu- ical and color pattern differences. Species recog- sions may be drawn. nition follows the Phylogenetic Species Concept (Cracraft, 1983). Because we adhere to this con- cept, we considered groups, which are geograph- SPECIES ACCOUNTS ically isolated and diagnosable, different species. Bufo aucoinae sp. nov. Figures 1–3 1 Corresponding Author. E-mail: [email protected]. Bufo melanochloris.—Savage and Villa, 1986 [In edu part]. 488 E. M. O’NEILL AND J. R. MENDELSON III FIG. 1. Dorsal (left) and ventral (right) pattern variation among adult male Bufo aucoinae and Bufo melanochlorus. Bufo aucoinae: top left, UCR 8717 (holotype, SVL 5 61.4 mm); top right, KU 91674 (SVL 5 50.3 mm). Bufo melanochlorus: bottom left, UCR 10431 (SVL 5 72.3 mm); bottom right, UCR 6989 (SVL 5 73.6 mm). Bufo melanochlorus.—Savage, 2002 [In part]; Airport, 3 km west of Rinco´n, 25 m; LACM-CRE Frost, 2002 [In part]. 9357 from 6 km southwest of Rinco´n de Osa; Holotype.—UCR 8717, an adult male from the Savage Woods, 10 m; LACM-CRE 9647 from 7.5 airport at the town of Aguabuena, Distrito km southwest of Rinco´n de Osa, Quebrado Rayo, Sierpe, Osa Peninsula, Puntarenas Province, 20 m. Costa Rica (approximately 88269N, 838389W). Referred Specimens.—See Appendix 1. Paratypes.—All from Puntarenas Province, Diagnosis.—A large species of Bufo (males to Costa Rica. Males: UCR 2082, from Eucalipto, 67.2 mm SVL; females to 104.5 mm SVL) having Rinco´n de Osa, Sierpe, Osa; UCR 12240 and the following combination of characters: (1) 12996 from bridge over Quebrada Can˜aza, tympanum small, 12.3–18.4% head length in Golfito, Golfito; KU 91663 and 91665 from Rı´o males, 11.2–15.5% in females; (2) preorbital and La Viejo, 30 km east of Palmar Norte, 100 m; KU pretympanic crests poorly developed, or absent, 91668 from 2 km northwest of Dominical, 10 M; in both sexes; (3) tibia short 42–50% of SVL; (4) LCAM-CRE 6538 from 7 km south-southwest of feet relatively short 40–47% of SVL; (5) dorsal Rinco´n Carretera al Pacifico; LACM-CRE 7083 skin evenly covered with tiny spiculae, spiculae from Rio La Vieja, 18.7 mi [30.1 km] by road east in males smaller and more concentrated than in of Palmar Norte, 110 m; 8031 from 4.2 km females; (6) lateral row of tubercles present as northwest by road from Villa Neily, 20 m. a series of low, rounded tubercles in males, in Females: UCR 0894 from Boscosa, Aguabuena, females as a series of large sharply pointed Sierpe, Osa; UCR12433 from Parque Nacional spiculae; (7) vocal slits present, bilateral, small; Esquinas, Sendero La Trocha, Golfito; UCR 12239 (8) m. interhyoideus forming a small, unilobed and 12997 from bridge over Quebrada Can˜aza, unpigmented vocal sac; (9) snout sharply pointed Golfito; UCR 14092 from La Gamba, Golfito, in dorsal view, rounded in profile; (10) cranial Golfito; KU 116978 from 8 km east-northeast of crests low, thin; (11) parotoid glands small, Palmar Norte, 90M; KU 65535 from Quebrada distinctly triangular; (12) tips of digits same color Boruca, 22 km east of Palmar Norte, 45M; KU as rest of digit. 65536–65538 from 12.3 km west-northwest of Bufo aucoinae (Figs. 1–3) is similar to B. Villa Neily, 25 m; LACM-CRE 6550 from Rinco´n melanochlorus (sensu stricto) but differs by lacking NEW SPECIES OF BUFO FROM COSTA RICA 489 transverse folds between parietal crests, lacking or having poorly developed pretympanic and preorbital crests, having a relatively unmarked venter, and by having the m. interhyoideus forming a small, unpigmented vocal sac. Bufo melanochlorus has transverse folds between the parietal crests (Fig. 2), cranial crests that are greatly elevated vertically, distinct pretympanic and preorbital crests, a black throat and chest with mottling on the flanks, and m. interhyoideus forms a larger, heavily pigmented vocal sac. Morphometrically, males of B. aucoinae differ from B. melanochlorus by having relatively smaller HL, HW, TYM, and PARW; females of FIG. 2. Skin texture of the dorsal surface of the area B. aucoinae have a larger TYM and PORB. Males between the cranial crests of Bufo aucoinae (left, of B. aucoinae are smaller (SVL) than are males of holotype, UCR 8717) and Bufo melanochlorus (right, B. melanochlorus. Morphometric statistics are UCR 8506) showing the distinctive folds in B. melano- presented in Table 1. chlorus. Description of Holotype.—Body robust; head wider than long, width 34.5% SVL, length 32.6% SVL; snout sharply pointed in dorsal view, subarticular tubercles distinct, elevated, triangu- rounded in profile, rostral keel absent; canthal, lar in profile, single; supernumerary tubercles supraorbital, supratympanic, postorbital, and unequal in size, distinct, distributed evenly over parietal crests present, low, thin; preorbital and ventral surfaces of foot and toes.