Abstract
We reviewed the taxonomic status of populations of frogs in the genus Hemiphractus in Panama, which have all been referred to Hemiphractus fasciatus Peters, 1862 for over 40 years. Although relatively few specimens have been collected, mostly juveniles, it is clear that these frogs inhabit three separate upland regions of the country: The Cordillera de Talamanca in western Panama, the Chagres Highlands and Cordillera de San Blas in central Panama, and the Serranía de Pirre in the far eastern portion of the country. In accordance with previously published molecular data, we identified distinctive features of the skulls of frogs representing these three allopatric populations and herein revalidate H. panamensis (Stejneger, 1917), describe the new species Hemiphractus elioti sp. nov. from the Cordillera de Talamanca, and the new species Hemiphractus kaylockae sp. nov. from the Serranía de Pirre. We also propose that the taxon H. fasciatus is a South American species not occurring in Panama.
Keywords: Amphibia, Colombia, Ecuador, Hemiphractus elioti,Hemiphractus kaylockae, taxonomy
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FIGURE 3. Photographs of live specimens of Hemiphractus from Panama.
(A) Specimen of H. panamensis photographed in situ (T. Herman); sex and SVL unknown. (B) Specimen of H. panamensis displaying mouth gaping behavior while being handled (E. Baitchman); sex and SVL unknown. (C) Live specimen of H. panamensis photographed in situ (T. Herman); sex and SVL unknown. Specimens of H. panamensis all found at Cerro Brewster, Panama, Panama.
(D, E) Adults of H. elioti, from Altos de María, near El Valle de Antón, Panama (B. Wilson, courtesy of El Valle Amphibian Conservation Center); probably males, SVL unknown. (F) Captive adult female of H. elioti with eggs (B. Wilson, courtesy of El Valle Amphibian Conservation Center); SVL unknown. |
Hemiphractus elioti new species
Hemiphractus panamensis Duellman 1970 [in part; for reference to specimens from western Panama.
Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from western Panama]; Duellman 2001 [in part; for reference to specimens from western Panama]; Crawford et al. 2012 [in part, for samples from Río Blanco and El Copé, Prov. Coclé, and Altos de Maria, Prov. Panamá, Panamá]; Castroviejo-Fisher et al. 2015 [in part; for sample from El Copé Prov. Coclé, Panamá]; Köhler 2011 [fig. 467; in part, for map showing isolated populations in western Panama].
Diagnosis.Hemiphractus elioti may be distinguished from all species of Hemiphractus except H. fasciatus, H. kaylockae, H. panamensis, and H. scutatus by not having a postorbital indentation. Hemiphractus elioti may be distinguished from H. scutatus, an Amazonian species not occurring in Panama, by its smaller size (maximum SVL in males 52.5 mm vs. 57.4 mm in H. scutatus; females 64.7 mm vs. 80.5 mm in H. scutatus; data for H. scutatus from Trueb 1974), and by having small expanded pads on the fingers and toes (absent in H. scutatus). Hemiphractus elioti differs from H. fasciatus, H. kaylockae, and H. panamensis by having neopalatine odontoids in contact with vomerine teeth at the center of the palatal region (narrowly separated in H. kaylockae vs. widely separated in both H. panamensis and H. fasciatus; condition unknown in H. scutatus; Fig. 6). Hemiphractus elioti further differs from H. panamensis and H. fasciatus by having approximately seven vomerine odontoids (vs. approximately two; condition unknown in H. scutatus; Fig. 6). Hemiphractus elioti may be distinguished from H. kaylockae by having straight margin of the lateral margins of the paraoccipital horns (with distinct indentation, forming an angular margin in H. kaylockae; Fig. 2); supraorbital ridges absent (evident, distinct in H. fasciatus, weakly developed in H. panamensis; Figs. 2, 9); canthal ridges usually absent or indistinct (evident, distinct in H. fasciatus, H. kaylockae, and slightly developed in H. panamensis; Figs. 2, 9). Hemiphractus elioti has a subtemporal fenestra very small in males, approximately one-half of orbit in females, whereas the fenestrae are approximately one-third diameter of the orbit in both males and females of H. kaylockae (Fig. 2), approximately ½ diameter of the orbit of females in H. fasciatus (Fig. 9; males unknown), very large, approximately 1.5 times size of orbit, in females of H. panamensis (Fig. 9; males unknown); lateral margins of the quadratojugals extend to level of the lateral tips of, or very slightly beyond, the paraoccipital horns in dorsal view (extending far beyond the lateral tips of the paraoccipital horns in H. panamensis; Figs. 2, 9).
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Etymology. Edgardo Griffith and Heidi Ross have greatly contributed to the knowledge base of the natural history, reproductive biology, and conservation of Panamanian amphibians, including Hemiphractus. We are proud to acknowledge the efforts of our friends and colleagues by naming this new species of Hemiphractus in honor of their son, Eliot. The specific epithet is the singular genitive case of the name Eliot.
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| FIGURE 8. Live specimens of Hemiphractus kaylockae. (A) Adult specimens from vicinity of Cana, Darién, Panama (B. Wilson, courtesy of El Valle Amphibian Conservation Center): probable males, SVL unknown. (C) Photograph (KUDA 003746) of specimen KU 93506 (C.W. Myers) from Cerro Pirre, Darién, Panama; adult female, SVL = 54.2 mm. (D) Photograph (KUDU 003744) of specimen KU 93505 (C.W. Myers) from Cerro Pirre, Darién, Panama; adult female, SVL = 40. 5 mm. (E) Adult specimen from vicinity of Cana, Darién, Panama (B. Wilson, courtesy of El Valle Amphibian Conservation Center): probably female, SVL unknown. |
Hemiphractus kaylockae new species
Hemiphractus panamensis Duellman 1970 [in part; for reference to specimens from eastern Prov. Darién, Panamá.
Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from eastern Prov. Darién, Panamá]; Duellman 2001 [in part; for reference to specimens from eastern Prov. Darién, Panamá]; Crawford et al. 2012 and Castroviejo-Fisher et al. 2015 [in part, for samples from Cana, Prov. Darién, Panamá]; Köhler 2011 [in part; for map showing isolated populations in extreme eastern Panama].
Diagnosis. Hemiphractus kaylockae may be distinguished from all species of Hemiphractus except H. elioti, H. fasciatus, H. panamensis and H. scutatus by not having a postorbital indentation (Fig. 2). Hemiphractus kaylockae may be distinguished from H. scutatus, an Amazonian species not occurring in Panama, by its smaller size (maximum SVL in males 52.5 mm vs. 57.4 mm in H. scutatus; females 64.7 mm vs. 80.5 mm in H. scutatus; data for H. scutatus from Trueb, 1974), and by having small expanded pads on the fingers and toes (absent in H. scutatus). Hemiphractus kaylockae differs from H. elioti by having neopalatine and vomerine odontoids narrowly separated (in contact in H. elioti; condition unknown in H. scutatus), whereas these rows of odontoids are widely separated in H. panamensis and H. fasciatus (Fig. 6). Hemiphractus kaylockae further differs from H. panamensis and H. fasciatus by having approximately seven vomerine odontoids (vs. approximately two in both H. panamensis and H. fasciatus). Hemiphractus kaylockae may be distinguished from H. elioti, H. fasciatus, and H. panamensis by having a distinct indentation on the paraoccipital horms, forming an angular margin of the paraoccipital horns (Figs. 2, 9); supraorbital ridges absent (evident, distinct in H. fasciatus and weakly developed in H. panamensis; Figs. 2, 9); canthal ridges evident, distinct (absent or weakly developed in H. elioti; Fig. 2). Hemiphractus kaylockae has a subtemporal fenestra approximately one-third diameter of the orbit in both males and females, approximately one-half diameter of orbit in females in H. elioti (very small in males; Fig. 2), approximately onehalf diameter of the orbit in females of H. fasciatus (Fig. 9; males unknown), very large, approximately 1.5 x size of orbit, in females of H. panamensis (Fig. 9; males unknown); lateral margins of the quadratojugals extend to level of lateral tips of, or slightly less than, beyond the paraoccipital horns in dorsal view (extending far beyond the profile of the paraoccipital horns in H. panamensis; Figs. 2, 9).
Etymology. This species is named in honor and memory of Julia Kaylock. She was a colleague and friend, beloved by many, who was passionate about the conservation of the world’s biodiversity, especially that of amphibians. She spent the latter part of her life working tirelessly to improve and document husbandry techniques for threatened Panamanian amphibians (e.g., Hill et al. 2012) maintained in ex situ facilities. She lost a lifelong battle with Type I Diabetes at the age of 28 on 12 June 2009. We graciously acknowledge her passion for amphibian conservation in the naming of this species of Hemiphractus. The specific epithet is the singular genitive case of the name Kaylock.
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FIGURE 3. Photographs of live specimens of Hemiphractus from Panama. (A) Specimen of H. panamensis photographed in situ (T. Herman); sex and SVL unknown. (B) Specimen of H. panamensis displaying mouth gaping behavior while being handled (E. Baitchman); sex and SVL unknown. (C) Live specimen of H. panamensis photographed in situ (T. Herman); sex and SVL unknown. Specimens of H. panamensis all found at Cerro Brewster, Panama, Panama.
(D, E) Adults of Hemiphractus elioti, from Altos de María, near El Valle de Antón, Panama (B. Wilson, courtesy of El Valle Amphibian Conservation Center); probably males, SVL unknown. (F) Captive adult female of H. elioti with eggs (B. Wilson, courtesy of El Valle Amphibian Conservation Center); SVL unknown.
FIGURE 8. Live specimens of Hemiphractus kaylockae. (A, B) Adult specimens from vicinity of Cana, Darién, Panama (B. Wilson, courtesy of El Valle Amphibian Conservation Center): probable males, SVL unknown. (C) Photograph (KUDA 003746) of specimen KU 93506 (C.W. Myers) from Cerro Pirre, Darién, Panama; adult female, SVL = 54.2 mm.
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| FIGURE 5. A map of the known distributions of the three species of Hemiphractus recognized herein, from Panama. Type localities are white symbols, black symbols represent localities for museum vouchers and photographic records (see Fig. 3). Symbols represent H. elioti (triangles); H. panamensis (circles), and H. kaylockae (square). At this time, we know of no confirmed records of these species from neighboring Costa Rica or Colombia, and we posit that H. fasciatus Peters, 1862, does not occur in Panama, but more likely is restricted to Ecuador and perhaps part of Colombia. |
Hemiphractus panamensis
Cerathyla panamensis Stejneger 1917.
Holotype: USNM 55320. Type locality: Signal Loma (Loma Peak) on the north coast of Panama, three miles [4.8 km] south of Santa Isabel, Prov. Colón, Panama.
Hemiphractus fasciatus Trueb 1974 [in part; for reference to specimens from Chagres Highlands, Panama]; Duellman, 2001[in part; for reference to specimens from Chagres Highlands, Panama]; Crawford et al. 2012 [in part, for samples from Cerro Brewster, Prov. Panama, Panama]; Castroviejo-Fisher et al. 2015 [in part, for samples from Cerro Bruja and Cerro Brewster, Prov. Panama, Panama]; Köhler, 2011 [figs. 472, 473; in part; for map showing isolated populations in the HEMIPHRACTUS OF PANAMA Zootaxa 4429 (3) © 2018 Magnolia Press · 507 Chagres Highlands, Panama].
Robert L. Hill, Kathryn G. Martin, Edward L. Stanley and Joseph R. I. Mendelson. 2018. A Taxonomic Review of the Genus
Hemiphractus (Anura: Hemiphractidae) in Panama: Description of Two New Species, Resurrection of
Hemiphractus panamensis (Stejneger, 1917), and Discussion of Hemiphractus fasciatus Peters, 1862.
Zootaxa. 4429(3); 495–512. DOI:
10.11646/zootaxa.4429.3.3
Resumen: Revisamos la situación taxonómica de las poblaciones de ranas del género Hemiphractus en Panamá, las cuales han sido referidas a Hemiphractus fasciatus Peters, 1862 durante más de 40 años. Aunque se conocen relativamente pocos ejemplares, la mayoría jóvenes, está claro que estas ranas habitan tres regiones separadas de elevaciones medias: La Cordillera de Talamanca al Oeste de Panamá, las Tierras Altas de Chagres y la Cordillera de San Blas en el centro de Panamá, y la Serranía de Pirre en el extremo oriental del país. En concordancia con datos moleculares previamente publicados, identificamos características distintivas de los cráneos de estas ranas que representan estas tres poblaciones alopátridas. De este modo, revalidamos H. panamensis (Stejneger, 1917) y describimos las especies nuevas Hemiphractus elioti sp. nov., de La Cordillera de Talamanca, y Hemiphractus kaylockae sp. nov., de la Serranía de Pirre. Proponemos que el taxón H. fasciatus es una especie sudamericana que no se encuentra en Panamá.
Palabras clave: Colombia, Ecuador, Hemiphractus elioti, Hemiphractus kaylockae, taxonomía
