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Atelopus onorei sp. nov. Atelopus bomolochos (non Peters): Lötters, 1996: Fig. 6; Coloma and Ron, 2001: Fig. 9. Holotype. QCAZ 1860, adult female, from Ecuador: Provincia Azuay: 10–15 Km E Luz María, Río Chipla (2 ° 47 ' 24 " S, 79 ° 21 ' 36 " W), approx. 2500 m above sea level, obtained on 9 April 1990, by Stella de la Torre- Salvador, John J. Wiens, Felipe Campos-Yánez, and Luis A. Coloma. Paratypes. KU 217448 – 57, QCAZ 1861 –63, 1865–67, 1868 (cleared and double stained preparation), 32258 – 59, same data as holotype; QCAZ 3440, 3442, same locality as holotype, obtained on 21 April 1990, by Fabián Toral, María Augusta Bravo, and Gustavo Morejón. Diagnosis. (1) A moderate-sized species with mean SVL in adult males 38.8 mm (35.2–41.3 mm, n = 9) and in adult females 44.8 mm (41.9–47.9, n = 4); (2) hind limbs short, mean tibia length/SVL 0.381 (0.341– 0.409, n = 13); (3) phalangeal formula of hand 2 - 2-3 - 3, basal webbing absent; (4) foot webbing formula I (0– 1)—(0–2) II (1 / 2 – 2)—(1–3) III (1–3)—(2–4) IV (2–4)—(1–3) V; (5) snout acuminate, slightly protruding beyond lower jaw; (6) tympanic membrane absent; (7) dorsal surfaces of body usually smooth, (one [out of 13] individuals bears a few scattered small spiculae); (8) yellow spiculae (creamy white in preservative) present mostly on flanks; (9) vertebral neural processes inconspicuous; (10) dorsum yellow-orange to light green (creamy yellow to gray in preservative); (11) venter uniform cream to pale yellow in preservative; (12) gular region without warts, spiculae or coni. By having in life an aqua blue iris, Atelopus onorei is distinct from all other species of Atelopus. By displaying in life yellow-orange, green, or a combination of yellow-orange and green colors, A. onorei is most similar to A. bomolochos. Descriptions of the type specimens of A. bomolochos did not include colors in life. Nonetheless, Peters (1973) described the color in alcohol as “may be yellowish, yellow-green, or light brown.” Specimens assignable to A. bomolochos, collected near the type and paratype localities reveal large color variation as described by Peters. All of them have a predominantly black iris as depicted in Figs. 3 B, C, D; although, specimens from 10 km S Cutchil show a fine green line around pupil (Field notes of John J. Wiens, 29 April 1990). Entirely yellow-orange individuals are most similar to A. bomolochos (from vicinities of Sigsig and Cutchil, Provincia Azuay; see Lötters 1996: Fig. 7), some specimens of Atelopus sp. from Departamento Nariño, Colombia (formerly referred to A. ignescens; for discussion see Coloma et. al., 2000), A. guanujo Coloma, 2002 from Provincia Bolívar, Ecuador, A. carbonerensis Rivero 1974, A. chrysocorallus La Marca, 1994, and A. sorianoi La Marca, 1983 all from the Andes of Venezuela (Estados Mérida and Trujillo). Atelopus onorei further differs from A. bomolochos by lacking minute black stippling regularly distributed on dorsum (Fig. 7), and by differences in hand and foot lengths of females. Herein, we compare A. onorei to populations described by Peters (1973) as A. bomolochos (specimens also used for morphometric analyses are indicated in the Appendix). The first two components with eigenvalues> 1.0 were extracted from the PCA of 7 morphometric variables of 15 males and 11 females from populations of A. bomolochos (Sevilla de Oro, Cuenca, and Juncal-General Morales) and A. onorei from Río Chipla. The axes accounted for 54.6 % of the total variation. There is no overlap among the morphometric space of females of A. onorei vs. A. bomolochos (Fig. 8), mostly because of hand and foot lengths, which are the highest loadings along PC II. Hand and foot lengths differ significantly between females of A. onorei vs. A. bomolochos (log comparisons of HAND of females t -test, t = 5.165044, df = 6.4, P> 0.0009; FOOT of females t -test, t = 4.578094, df = 8.9, P> 0.0007). Atelopus onorei (mean SVL of females 44.8; mean SVL of males 38.8) is significantly larger (SVL of females t -test, t = 2.992, df = 14, P t -test, t = 7.320, df = 36, P A. guanujo (mean SVL of females 39.6, 35.0–46.0, SD 3.0, n = 12; mean SVL of males 33.9, 29.9–37.1, SD 1.6, n = 29; after data in Coloma, 2002) and lacks white spiculae in life. Atelopus onorei differs from Atelopus sp. from Nariño by lacking a patch of black spiculae on females’ gular-chest region. Atelopus onorei differs from A. carbonerensis and A. sorianoi in having a less protuberant snout and lacking a noticeably postocular crest. Furthermore, it differs from A. carbonerensis and A. chrysocorallus by having vocal slits in males and lacking a row of warts on the dorsolateral surfaces of the body in males (after data from La Marca, 1996; Rivero, 1974; Coloma, 1997). Description of holotype. (Figs. 9 A–C). Head about as long as wide, HLSQ and HDWD less than one third SVL (HLSQ /SVL = 0.280, HDWD /SVL = 0.282); snout acuminate, its margin slightly rounded in dorsal view; profile of tip of snout in lateral view curved and slightly protuberant to the anterior margin of jaw; no swollen gland on tip of snout; nostrils slightly protuberant, directed laterally, situated posterior to level of apex of lower jaw; canthus rostralis distinct, weakly concave from eye to nostril; loreal region concave; lips not flared; interorbital and occipital regions flat, smooth; eyelid flared without tubercles; postorbital crest slightly raised, glandular; low pretympanic and post tympanic areas warty; tympanic membrane and tympanic annulus absent; temporal area smooth; choanae small, rounded, widely separated (28 % of HW); tongue about twice as long as wide, its posterior half not attached to mouth’s floor; ostia pharyngea absent. Forearm relatively short (RDUL /SVL = 0.278); palmar tubercle round; palmar, supernumerary palmar, thenar and subarticular tubercles nearly indistinct; digital tips with round pads; thumb relatively long (THBL / HAND = 0.717), apparently having two phalanges; webbing on hands absent, fingers lacking lateral fringes; relative length of fingers III 1 — 1 II 1— 2 III 2— 3 IV 3 — 2 V; relative length of toes I Dorsal surfaces smooth; flanks rugose with scattered spiculae; anterior and proximal upper surfaces of fore limbs lacking spiculae; throat rugose, contrasting to smooth chest, belly, and undersides of hind limbs; cloacal opening a tube at nearly midlevel of thighs, directed posteriorly; skin lateral to cloacal opening rugose. In preservative (~ 70 % ethanol), dorsum of head mostly cream with diffuse gray marks, loreal region gray; dorsum of body gray extending onto upper flanks; dorsal and ventral surfaces of limbs creamy yellow, except tibia, fingers III, V, toes IV and V, which have gray marks with diffuse borders; minute gray stippling absent on dorsum of body (viewed at 8 X magnification); spiculae white; throat, chest, belly, ventral surfaces of limbs, palms and plants yellowish cream; proximal end of tongue lacking black pigmentation, inner margin of mandible with a fine gray line. Measurements (in mm). SVL 47.9, TIBL 17.7, FOOT 18.1, HLSQ 13.4, HDWD 13.5, ITNR 3.7, EYDM 4.5, EYNO 3.3, RDUL 13.3, HAND 10.6, THBL 7.6, SW 14.6. Variation. Meristic variation is given in Table 1. The paratypes resemble the holotype with the following exceptions. Sexual dimorphism is evident in that females are larger than males (Table 1). Males have vocal slits and keratinized nuptial pads on the dorsal and inner surfaces of the thumb. Fore limbs are relatively long and slender in females only (Table 1). Spiculae are most abundant on the flanks in a female (QCAZ 3440) and some are scattered on the dorsum of the body in that specimen. Color variation in preservative (~ 70 % ethanol): Dorsal colors vary from uniform yellow (QCAZ 3440) to mostly gray (QCAZ 3442). In QCAZ 2798, colors are faded because of poor preservation. Color in life (Figs. 3 E–F). Data from color transparencies (no museum numbers associated) and the senior author’s field notes of specimens from the type locality. The dorsal ground color of body and limbs is orange yellow. The dorsum of one male is bright green, except the forearms, hands and feet, whereas other males and females show varying degrees of green coverage. In a few individuals the dorsum is pale yellowish brown with diffuse black blotches; the spiculae are yellow. The iris is conspicuously aqua blue with fine black reticulation and with a fine bluish-white stripe at the upper margin of pupil. Lötters (1996: Fig. 6) and Coloma and Ron (2001) published the same color photograph of an amplectant pair of A. onorei (as A. bomolochos). Nogales and Rueda-Almonacid (2005) also provided a painting and a color photo (under A. bomolochos). Distribution, ecology and current population status. Atelopus onorei is known only from the type locality in the Azuay Basin of the Cordillera Occidental of Ecuador, Provincia Azuay (Fig. 6) and a nearby creek (1–2 km West of Miguir, Morley Read pers. com. and his personal archive of photographs). At the latter locality, it occurs in sympatry with A. nanay Coloma 2002. The type locality is at about 2500 m above sea level in humid cloud forest on the Pacific versant of the Cordillera Occidental. Annual mean rainfall is 1000– 2000 mm and the annual mean temperature is 12–18 ºC. Frogs were collected at Cloud Montane Forest areas (sensu Val en c ia et al. 1999), at the margin of the Río Chipla and near a small tributary. At the time of collection, during day at the type locality, most individuals were active and numerous pairs were in amplexus, while drizzle occurred (Field notes of L. A. Coloma, 9 April 1990). A female (QCAZ 3440) contained 133 ovarian eggs that are 2.14 mm in diameter (range = 1.67–2.58, n = 20, SD = 0.219). Although this species previously was abundant at the type locality, no individuals have been found after 1990, despite at least four occasional search efforts by the senior author to locate this frog at the same site as before and surroundings on 28 November 1992, 20 February 1993, 21 April 1993, and 11 October 1993. Etymology. The specific name is a noun in the genitive case and it is a patronym for Giovanni Onore, former Curator of Entomology at Pontificia Universidad Católica del Ecuador (PUCE). We recognize his invaluable contribution to the collections of Ecuadorian frogs and pioneer efforts on forming collections for scientific research at PUCE. His collecting efforts, especially during the eighties, resulted in the discovery of many new amphibians from Ecuador, among them Atelopus. As well, his amphibian collections in the Andean region of Ecuador provided key data, later used to document the amphibian extinctions and to shed light on its causes.

Atelopus pachydermus (Schmidt, 1857) Phirix pachydermus Schmidt, 1857: 15. Holotype: KM 1013 / 1356 from ���Neu-Granada���; Schmidt, 1858: 257. Phryniscus pachydermus: Hoffmann, 1878: 635 (by implication). Phryniscus cruciger (non Lichtenstein and Martens): Boulenger, 1882: 154 (partim). Atelopus cruciger (non Lichtenstein and Martens): Nieden, 1926: 84 (partim). Phirix pachydermus: Dunn, 1931: 397. Atelopus subornatus (non Werner): Dunn, 1944: 81. A.[telopus] pachydermus: Rivero, 1963: 107. Atelopus pachydermus: Rivero, 1965: 138 (partim); Peters, 1973: 34 ���37 (partim); L��tters, 1996: 39 ���40 (partim), 2005: 97 (partim). Referred specimens. Peru: MUSM 6524 from Camporedondo, Departamento Amazonas; MUSM (field number 890081 a), MUSM (field number 890085), MUSM 13810 ��� 12, MUSM 5984, 6507 collected from nearby Grutas of Parque Nacional Cutervo, Departamento Cajamarca, obtained on March 1984 by Alfonso Miranda-Leiva; Ecuador: MHNG 2259.8 ��� 9 from Valladolid-Yangana, approximately 2600 m above sea level, Provincia Zamora Chinchipe, obtained on September 1985 by Giovanni Onore and Luis A. Coloma. Diagnosis. (1) A large species with mean SVL in adult males 51.3 mm (44.2���56.4, n = 7) and in adult females 55.9���61.6 (n = 2); (2) hind limbs short, tibiotarsal articulation reaching to temporal region, when leg carried forward along body (tibia length/SVL = 0.375���0.439, n = 9); (3) phalangeal formula of hand 2 ��� 2���3 ��� 3 (suspected from external examination), basal webbing absent; (4) foot webbing formula I (0+)���(1 - to 1 +) II (0+ to 1 -)���(1 to 2 +) III (1 to 2 -)���(2 + to 3) IV (2 to 3 +)���(0+ to 1 -) V; (5) snout acuminate, with tip gently rounded in dorsal view; upper jaw slightly protruding beyond lower; (6) tympanic membrane absent; (7) dorsum with widely scattered or dense well-defined, rounded warts; (8) scattered spiculae on posterior and ventrolateral surfaces of body or absent; (9) vertebral neural processes inconspicuous; (10) dorsum yellow with dark brown pattern of marks, blotches and/or lines (dorsum cream, tan, or olive with pale to dark brown pattern in preservative); (11) venter white, occasionally orange or red (venter tan to yellow with pale to dark brown blotches in preservative); (12) gular region without warts, spiculae, or coni. Atelopus pachydermus is distinguished from all Atelopus by its large size, except A. boulengeri Peracca, 1904, A. arsyecue Rueda-Almonacid, 1994, and A. laetissimus Ru��z-Carranza, Ardila-Robayo and Hern��ndez-Camacho, 1994. It differs from all of them by a different color pattern. Atelopus boulengeri usually has a uniform dorsum, and is larger than A. pachydermus (mean female SVL of A. boulengeri [after data in Coloma, 1997]) = 65.9 vs. 55.9���61.6 in A. pachydermus). Atelopus arsyecue has large white, cream, creamy-yellow, round or elongated marks (Rueda-Almonacid, 1994; Renjifo and Renjifo, 2005: 57). Atelopus laettisimus has a reticulated pattern usually bearing an X-mark on the anterior dorsum (Ru��z-Carranza et al., 1994; Galvis, 2005: 83). Description of holotype. Adult male (Fig. 1), well preserved; original colors apparently bleached; forearm twice as thick proximally as distally; keratinized areas on thumb. Head slightly longer than wide; head length and head width each less than one third SVL; snout acuminate, tip of snout with swollen gland, rounded in dorsal view, slightly protruding beyond anterior margin of the jaw in profile; nostrils not protuberant, barely visible from above, situated slightly behind level of apex of lower jaw; canthus rostralis distinct, convex from tip of snout to nostril, concave from nostril to eye, swollen posteriorly; loreal region concave, upper lip slightly flared, interorbital and occipital regions flat, smooth, eyelid flared without distinct tubercles; postorbital crest glandular; tympanic membrane absent; choanae minute, rounded; tongue twice as long as wide, broader anteriorly, less than its posterior half not attached to mouth���s floor. Forearm relatively short, length of forearm less than one third SVL; metacarpal tubercles distinct, round, outer about twice size of somewhat weaker inner; few weak supernumerary tubercles present; subarticular tubercles at joints of fingers IV and V; digital tips slightly broadened, with round pads; thumb relatively short, more than half length of hand, apparently having two phalanges; nuptial pads on inner or dorsal thumb absent; basal webbing and fringes on fingers absent; relative length of fingers III( 0+)���(1) II (0+)���(2) III( 2 -)���(3) IV (3 +)���(0+) V, webbing always becoming fringelike distally; relative length of toes IMeasurements (mm). SVL 52.9, TIBL 20.7, FOOT 23.5, HLSQ 14.9, HDWD 15.2, EYDM 4.6, EYNO 3.7, ITNA 4.5, SW 15.6, RDUL 16.7, HAND 14.8, THBL 9.3. Variation. Meristic variation of seven adult males (including holotype) and two females is given in Table 1. Two juveniles from Ecuador (MHNG 2259.8 ��� 9) have SVL 29.7 and 21.7, respectively. They are tentatively assigned to Atelopus pachydermus. All specimens resemble the holotype. The two available females (MUSM 5984, 6524) are larger than the males; the forearm of females is not swollen; nuptial pads, present in adult males (but absent in MUSM 6507), cover dorsum and inner margins of Finger II. The number of warts is more or fewer than in the holotype. Warts, and spiculae are highly visible. In MUSM 6524 spiculae are present behind the eye, on posterior part of dorsum and on the thighs, and in MUSM 13812 also ventrolaterally on the body. In some specimens warts are present in the interorbital region, but never anterior to it; laterally warts are present in the temporal region; warts also are present around the cloaca opening. In MUSM 5984 (female) some warts are present on the chest, and in MUSM (field number 890085, male) warts are present on the chest and the ventral surfaces of the thighs. Subarticular tubercles are present on joints of phalanges of all toes and fingers. Scattered supernumerary tubercles are present on the soles and palms of all individuals. Foot webbing ranges as follows: I (0+)���(1 - to 1 +) II (0+ to 1 -)���(1 to 2 +) III (1 to 2 -)���(2 + to 3) IV (2 to 3 +)���(0+ to 1 -) V. Atelopus petersi Atelopus pachydermus Provincia Napo Provincia Chimborazo Departamentos Cajamarca & Amazonas Color variation in preservative (~ 70 % ethanol): Dorsum and venter color pattern variation is depicted in Figure 2. The dorsum is tan or olive with a pale to dark brown pattern, or blackish brown with tan to pale yellow pattern. The dorsal pattern consists of irregular or semi-symmetrical markings, blotches and/or lines; the anterior head, or at least its periphery, and most parts of the feet and hands are tan or pale yellow. The arms and legs are tan or olive with pale to dark brown semi-symmetrical dots and blotches, or blackish brown with tan or pale yellow markings. The venter is tan or yellow with few irregular pale to dark brown markings, most evident on the posterior part of the belly (at least brown markings below the cloaca); legs and arms always with markings, sole and palm in part or without markings, tarsal tubercles usually uniform yellow. Color in life (Fig. 3 A): L��tters (2005: 97) provided a color photograph of Atelopus pachydermus from Cutervo. As well, he provided a color in life description. ���Dorsum very variable, mostly bright yellow with large black markings. The yellow color turns to white (occasionally orange or red) in ventral region.��� Juveniles from Ecuador (MHNG 2259.8 ��� 9, color transparencies (QCAZ transparencies 2292 ��� 94, available at AmphibiaWebEcuador: http://www.puce.edu.ec/zoologia/vertebrados/amphibiawebec/anfibiosecuador/ bufonidae.html) have dorsum, dorsal surfaces of limbs and flanks yellow-green with black marks. The iris has a yellow ring around the pupil. Distribution, ecology and current population status. Atelopus pachydermus is known from the Cordilleras de Tarros (at Cordillera Occidental) and Central of the Andes of northern Peru, departamentos Cajamar-ca and Amazonas, and from Cordillera Oriental of the Andes in southern Ecuador, Provincia Zamora Chinchipe at about 2600 m above sea level (Fig. 4). In Peru, the area in which A. pachydermus occurs includes Humid Montane Forest, following the classification of types of vegetation of the Peruvian Andes as modified by Duellman (2004), who followed the system proposed by Tosi (1960). At the Peruvian Parque Nacional Cutervo (6 �� 00'��� 6 �� 20 ' S 78 �� 40 '��� 78 �� 53 ' W, 2350���3350 m above sea level) annual mean temperature is 13.8 C. According to Duellman and Pramuk (1999), the vegetation in Peru of the humid montane forest of Cordillera Occidental is reduced to few trees, bushes (Baccharis) and bunch grasses; much of this forest has been cleared and cultivated, and at higher elevation Eucalyptus has been planted. In Ecuador, the locality is in Montane Cloud Forest (Valencia et al. 1999) in the Nudo de Sabanilla. In its proximities, 2000 hectares of forest and paramo habitats are protected within the Reserva Tapichalaca, which is located south of the Parque Nacional Podocarpus. Annual mean precipitation in this region is 1000���2000 mm and annual mean temperature is 12���18 C (Ca��adas-Cruz, 1983). In both adult females studied by us, eggs are visible through the skin of the venter. The current population status of Atelopus pachydermus in Peru is unknown. Fieldwork by The University of Kansas in February 1989 in the vicinity of Cutervo, Departamento de Cajamarca revealed no Atelopus. Nonetheless, the most recent record is from 1994 or 1995, close to San Andr��s caves in Parque Nacional Cutervo, when Heinz Plenge took the photograph shown in L��tters (2005). Searches for A. pachydermus were carried out by AML at Parque Nacional Cutervo in 1996 and nearby localities in Provincia Chota in 1997 and 1998; at north of San Andr��s de Cutervo by Pablo Venegas in January 2007; they revealed no Atelopus. Also, people at these localities mentioned their absence. The San Andr��s caves are located in Parque Nacional Cutervo, which has 8214 hectares of land protected by the Peruvian Government. In spite of its protection status, potato and artichoke field crops are located in the area of the Cutervo caves and there is extensive cattle ranching inside the park. The only record from Ecuador is from September 1985. Searches at the Ecuadorian locality were carried out on February and September 2001 and on April 2004; they revealed no Atelopus. Comments. Rodr��guez et al. (1993) and L��tters (2005) reported Atelopus pachydermus from the Andes of northern Per�� (departamentos Amazonas, Cajamarca, and La Libertad). They failed to provide museum and specimen numbers, and specific locality data. Listing Departamento La Libertad was an error and referred to an undescribed smaller, but otherwise similar species from Pataz (see Catenazzi and Venegas, 2005: 148). Translation of the original description of Phirix pachydermus Oscar Schmidt (1857: 14���15). " Phirix with robust fore- and hind limbs, head moderate; snout somewhat acute; skin thick, firm; color sulphur to whitish, light bluish pattern on dorsum and on femur as well as in cloacal region." For a translation from German into English of Schmidt��s 1858 description see Cochran and Goin (1970: 120). Etymology. The specific name pachydermus is Greek and means thick skin., Published as part of Coloma, Luis A., L��tters, Stefan, Duellman, William E. & Miranda-Leiva, Alfonso, 2007, A taxonomic revision of Atelopus pachydermus, and description of two new (extinct?) species of Atelopus from Ecuador (Anura: Bufonidae), pp. 1-32 in Zootaxa 1557 on pages 4-11, DOI: 10.5281/zenodo.178262, {"references":["Schmidt, O. (1857) Diagnosen neuer Frosche des Zoologischen Cabinets zu Krakau. Sitzungsberichte der Akademie der Wissenschaften in Wien. Mathematisch-Naturwissenschaftliche Klasse, 24, 10 - 15.","Schmidt, O. (1858) Deliciae herpetologicae musei zoologici cracoviensis. Denkschriften der Kaiserlichen Akademie der Wissenschaften in Wien. Mathematisch-Naturwissenschaftliche Klasse, 14, 237 - 258.","Hoffmann, C. K. (1878) In Bronn (ed.), Die Klassen und Ordnungen des Thier-Reichs, 6 (2), pp. 635.","Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. London: Taylor & Francis. xvi + 495 pp., 30 pl.","Nieden, F. (1926) Anura II. Das Tierreich. Walter de Gruyter & Co., Berlin and Leipzig, Germany, 110 pp.","Dunn, E. R. (1931) New frogs from Panama and Costa Rica. Occasional Papers of the Boston Society of Natural History, 5, 385 - 401.","Dunn, E. R. (1944) Herpetology of the Bogota area. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 6, 68 - 81.","Rivero, J. A. (1963) Five new species of Atelopus from Colombia, with notes on other forms from Colombia and Ecuador. Caribbean Journal of Science, 3, 103 - 124.","Rivero, J. A. (1965) Notes on the Andean salientian (Amphibia) Atelopus ignescens (Cornalia). Caribbean Journal of Science, 5, 137 - 140.","Peters, J. A. (1973) The frog genus Atelopus in Ecuador (Anura: Bufonidae). Smithsonian Contributions to Zoology, 145, 1 - 49.","Lotters, S. (1996) The Neotropical Toad Genus Atelopus. Checklist - Biology - Distribution. M. Vences & F. Glaw, Cologne, Germany, 143 pp.","Peracca, M. G. (1904) Viaggio del Dr. Enrico Festa nell'Ecuador e regioni vicine: rettili ed amfibii. Bolletino dei Musei di Zoologia ed Anatomia Comparata della Universita di Torino 19 (465): 1 - 41.","Rueda-Almonacid, J. V. (1994) Una nueva especie de Atelopus Dumeril & Bibron 1841 (Amphibia: Anura: Bufonidae) para la Sierra Nevada de Santa Marta, Colombia. Trianea, 5, 101 - 108.","Ruiz-Carranza, P. M., Ardila-Robayo, M. C. & Hernandez-Camacho, J. I. (1994) Tres nuevas especies de Atelopus A. M. C. Dumeril & Bibron 1841 (Amphibia: Bufonidae) de la Sierra Nevada de Santa Marta, Colombia. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 19, 153 - 163.","Coloma, L. A. (1997) Morphology, systematics and phylogenetic relationships among frogs of the genus Atelopus (Anura: Bufonidae). Ph. D. dissertation, Lawrence, University of Kansas, 287 pp.","Renjifo, J. M. & Renjifo, C. (2005) Atelopus arsyecue. In Rueda-Almonacid, J. V., Rodriguez-Mahecha, J. V., Lotters, S., La Marca, E., Kahn, T., & Angulo, A. (Eds.), Ranas arlequines. Conservacion Internacional. Panamericana Formas e Impresos S. A., Bogota, Colombia, pp. 57.","Galvis, P. A. (2005) Atelopus laetissimus. In Rueda-Almonacid, J. V., Rodriguez-Mahecha, J. V., Lotters, S., La Marca, E., Kahn, T. & Angulo, A. (Eds.), Ranas arlequines. Conservacion Internacional. Panamericana Formas e Impresos S. A., Bogota, Colombia, pp. 83. Gray, P. (1983) Morphometrics of the Atelopus ignescens Complex (Anura: Bufonidae). M. S. Thesis, University of Kansas, Lawrence, Kansas, U. S. A.","Duellman, W. E. (2004) Frogs of the genus Colostethus (Anura; Dendrobatidae) in the Andes of northern Peru. Scientific Papers Natural History Museum The University of Kansas, 35, 1 - 49.","Tosi, J. A., Jr. (1960) Zonas de vida natural en el Peru. Instituto Interamericano de Ciencias Agricolas, Zona Andina, Boletin Tecnico 5, Lima, Peru.","Duellman, W. E. & Pramuk, J. B. (1999) Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers Natural History Museum The University of Kansas, 13, 1 - 78.","Valencia, R., Ceron, C., Palacios, W. & Sierra, R. (1999) Las formaciones naturales de la Sierra del Ecuador. In Sierra, R. (Ed.), Propuesta preliminar de un sistema de clasificacion de vegetacion para el Ecuador continental. Proyecto INEFAN / GEF-BIRF y Ecociencia, Quito, Ecuador, pp. 79 - 108.","Canadas-Cruz, L. (1983) El mapa bioclimatico y ecologico del Ecuador. Ministerio de Agricultura y Ganaderia. Programa Nacional de Regionalizacion Agraria. Banco Central del Ecuador, Quito, Ecuador, 209 pp.","Rodriguez, L. O., Cordova, J. & Icochea, J. (1993) Lista preliminar de los anfibios del Peru. Publicaciones del Museo de Historia Natural, Universidad Nacional Mayor de San Marcos (A), 45, 1 - 22.","Catenazzi, A. & Venegas, P. J. (2005) Atelopus sp. 12. In Rueda-Almonacid, J. V., Rodriguez-Mahecha, J. V., Lotters, S., La Marca, E., Kahn, T., & Angulo, A. (Eds.), Ranas arlequines. Conservacion Internacional. Panamericana Formas e Impresos S. A., Bogota, Colombia, pp. 148.","Cochran, D. M. & Goin, C. J. (1970) Frogs of Colombia. United States National Museum Bulletin, 288, 1 - 655."]}

Atelopus petersi sp. nov. Atelopus pachydermus (non Schmidt): Rivero, 1963:107, 1968: 23; Harding, 1983: 58; Peters, 1973: 34; Frost, 1985: 31; Lötters, 1996: 39; Coloma and Ron, 2001: 38, La Marca et al. 2005: 195. Atelopus sp.: Almendáriz and Cisneros, 2005: 155. Holotype. QCAZ 300, from Ecuador: Provincia Napo: Cantón Quijos: near Oyacachi, (00° 12 ' 36 " S, 78 ° 42 '00" W; approximately 3200 m above seal level), obtained on 3 April 1988 by Stella de la Torre-Salvador. Paratypes. QCAZ 301 and 302 (cleared and double stained preparation), same data as holotype, collected by Manuel Pallares; QCAZ 298 (cleared and double stained preparation), 299, and 3154, same data as holotype, obtained on 30 March 1988 by Felipe Campos-Yánez and Stella de la Torre-Salvador; QCAZ 4555 from Río Oyacachi, 2 Km east from Oyacachi, obtained on 19 August 1993 by Elicio Tapia; QCAZ 7678 from 1 km E of Oyacachi, 3150 m above sea level, obtained by Felipe Campos-Yánez, Jorge Washington Izquierdo, and Diego Almeida-Reinoso, on 8 November 1996; USNM 193494, 193523, 193526 – 31 from Oyacachi, obtained by A. Proaño (no date); USNM 193521 from 2 Km E of Chalpi, 2755 m above sea level obtained on 7 July 1962 by James A. Peters; MHNG 2246.93 – 94 from 11 km (by road) east-southeast of Papallacta (00°03'S, 78 °08' W), 2660 m above sea level, obtained on November 1985 by Giovanni Onore and Luis A. Coloma; KU 117874 – 79 from 3 Km east of Papallacta, 2900 m above sea level, obtained on 7 March 1968 by John D. Lynch, KU 142950 from 7.8 Km west-north-west of Cuyuja, KU 142951 – 2 from 5.7 Km east of Papallacta, KU 142953 – 54 from Río Papallacta, 4.6 Km east of Papallacta, 2890 m above sea level, obtained on 28 October 1971 by William E. Duellman; USNM 193496 –502, 193506– 18 from 0.5 km E of Papallacta, 3150 m above sea level, obtained on 5 July 1962 by James A. Peters and P. Spoecker; USNM 193503, from 1 km W of Papallacta, 3150 m above sea level, obtained on 6 July 1962 by James A. Peters et al. Referred specimens. QCAZ 3870 and 3871 from Provincia Chimborazo: Lagunas de Atillo, obtained on 3 November 1990 by Anne Claire Desfossey; EPN 3307 from Laguna Cuyug (Cantón Guamote), 3318 m above sea level, 02° 10 ' 42 '' S, 78 ° 30 ' 30 '' W, obtained on 19 September 1986 by Ana Almendáriz. Diagnosis. (1) A moderate-sized species with mean SVL in adult males 38.9 mm (35.2–42.1 mm, n = 16) and in adult females 46.0mm (43.3–50.2, n = 11); (2) hind limbs short, mean tibia length/SVL 0.370 (0.327– 0.420, n = 26); (3) phalangeal formula of hand 2 - 2-3 - 3, basal webbing absent; (4) foot webbing formula I( 0+)—(0+– 1) II (0+– 1)—(0+– 1) III (0+– 1)—(2) IV (2 -– 2)—(0+– 1) V; (5) snout acuminate, slightly protruding beyond lower jaw; (6) tympanic membrane, tympanic annulus and stapes absent; (7) dorsal surfaces varying from smooth to warty; (8) white spiculae (cream in preservative) on forearms, flanks, and dorsal surfaces of thighs; (9) vertebral neural processes inconspicuous; (10) dorsum with yellow orange pattern to almost entirely black, flanks white (cream to cream and black in preservative); (11) venter yellow with or without black marks, or uniform orange to reddish orange (cream to pale yellow, occasionally with black marks in preservative); (12) gular region without warts, spiculae or coni. Atelopus petersi is distinguished from all other species of Atelopus (except an undescribed species from southern Ecuador, Provincia Loja) by its white digital pads that contrast with adjacent black phalanges. Atelopus petersi is most similar to A. bomolochos, A. pachydermus, and Atelopus sp. (Provincia Loja) by having digital pads contrasting in color to adjacent phalanges. It differs from A. bomolochos and Atelopus sp. by having few spiculae on flanks (i.e., almost lacking); numerous spiculae present on flanks of A. bomolochos (Figs. 3 B–D) and A. sp. It further differs from A. bomolochos by having white marks on flanks (absent in A. bomolochos). It differs from A. pachydermus by significantly smaller size (mean female SVL of A. petersi = 46.0 vs. 55.9–61.6 in A. pachydermus) and by lacking brown colors in life. Description of holotype. (Figs. 5 A–C). Head slightly longer than wide; head length and head width less than one third SVL (HLSQ /SVL = 0. 284; HDWD /SVL = 0.262; snout truncate in dorsal view; barely protruding beyond anterior margin of jaw; tip of snout lacking swollen gland; nostrils slightly protuberant, directed laterally, situated behind level of apex of lower jaw; canthus rostralis distinct, slightly swollen from eye to nostril; loreal region concave, lips not flared, interorbital and occipital regions flat, smooth, eyelid flared without distinct tubercles; postorbital crest glandular; temporal areas warty; tympanic membrane and tympanic annulus absent; choanae small, rounded, widely separated; tongue more than twice as long as wide, slightly broader anteriorly, its posterior half not attached to mouth’s floor; ostia pharyngea absent. Forearm relatively short (RDUL /SVL = 0.297); palmar tubercle round, supernumerary palmar tubercles absent; thenar and subarticular tubercles distinct; digital tips with round pads; thumb relatively long (THBL / HAND = 0.628) having two phalanges; webbing on hands absent, fingers having lateral fringes; relative length of fingers II Tibia relatively short (tibia/SVL = 0.360); no fold on distal half of inner edge of tarsus; outer metatarsal tubercle round, slightly raised, about two thirds length of oval inner metatarsal tubercle; supernumerary plantar tubercles absent; subarticular tubercles conspicuous, digital pads distinct; toes webbing formula I( 0+)—(0+– 1) II (0+– 1)—(0+– 1) III (0+– 1)—(2) IV (2 -– 2)—(0+– 1) V; relative length of toes I Dorsal surfaces varying from smooth to warty; scattered, irregularly distributed white spiculae and warts on flanks; warts on anterior and proximal dorsal surface of fore limb, throat, chest, and belly; undersides of hind limbs rugose, but free of warts; cloacal opening an inconspicuous tube at midlevel of thighs, directed posteriorly; low, elongated warts lateral to cloacal opening. In preservative (~ 70 % ethanol), cream with black, irregular marks on dorsum of body and limbs. Tips of fingers and toes cream dorsally and ventrally, contrasting to black adjacent phalanges. Dorsum of snout cream. Flanks cream and black, except infraorbital and canthal regions, which are cream. Venter cream anteriorly and cream with five scattered, irregular, black marks posteriorly. Limbs ventrally mostly cream, except at joints with irregular black marks. Palms and soles cream with black marks. Black mark below cloaca. Measurements (mm). SVL 45.8, TIBL 165, FOOT 18.7, HLSQ 13.2, HDWD 12.0, ITNR 4.6, EYDM 4.5, EYNO 3.3, RDUL 13.8, HAND 12.2, THBL 7.6, SW 14.4. Variation. Meristic variation is given in Table 1. Besides larger female size, the forearm in females is not swollen. Nuptial pads of adult males cover dorsum and inner margins of Fingers II and III. Nuptial pads vary from covering only Finger II (KU 117876) to Fingers II and III (KU 142950). When covering Finger III, nuptial excrescences can be more abundant proximally and at inner margin (KU 142954). Color variation in preservative (~ 70 % ethanol): Specimens from Napo, i.e., Oyacachi and Papallacta show large variation in dorsal and ventral patterns, from dorsum nearly entirely black (KU 117875) to mostly cream (QCAZ 299), and from venter white (KU 117875) to mostly marbled (KU 117879). Thus, they fit well descriptions and illustrations of color patterns in dorsal and ventral views provided by Peters (1973:Figs. 25– 27, under A. pachydermus) of specimens from the same region. A juvenile (QCAZ 3154, SVL = 17.8 mm) has a fine mid-dorsal line. Referred Specimens from Provincia Chimborazo (QCAZ 3870, 3871, EPN 3307) are dorsally predominantly black, but have some cream blotches. QCAZ 3871 has a black dorsum with cream blotches, one on the snout, one on the upper eyelid, a small one on the interorbital region, two on the scapular region, and three on the sacral region; the flanks are black with minute white tips of spiculae. The loreal and suborbital regions are cream stippled with brown; the lower lip, throat, and chest are entirely white, whereas the belly and undersides of the limbs are black, except for a large white mark on belly. The palmar surfaces are white, and gray digits contrast with white tips of fingers dorsally, and all of Finger II is white dorsally. The dorsal surfaces of the feet, and soles are black; the ventral surfaces of Toe I, tips of toes, and metatarsal and subarticular tubercles are white. QCAZ 3870 differs from the previous individual by having the dorsum posterior to the head entirely black, whereas the dorsum of the head is mostly black with diffuse gray on the snout; the belly has four cream blotches and palms are mostly cream. EPN 3307 differs from QCAZ 3871 by having the dorsum of head mostly cream and four asymmetrical cream round blotches on the dorsum of the anterior part of body, the throat with a small black blotch, and the belly with a large white mark Color in life. Peters (1973: 35) described specimens from Provincia Napo (Chalpi, Oyacachi and Papallacta) as follows: “The lighter dorsal areas are bright yellow. Many of the warts and pustules have white tops. The ventral surfaces are white in almost all individuals, but some may have orange areas or spots and some have red bellies.” USNM 193518 has yellowish green spotting on dorsum and a bright orange belly (Field Notes of James A. Peters, 5 July 1962). KU 117874 – 75 is black above, 117874 is red below, 117875 is orange; metatarsal tubercles are yellow; KU 117876 – 79 juveniles are black above with dark faintly yellow blotches; black and white below. All specimens except 117879 are white below with black markings; in 117879 there is pale red pigment in the white areas posteriorly and dorsum black with few to many yellow spots. KU 117879 has yellow-white spots on back; flanks black with white spots; metatarsal tubercles white; iris dark brown (Field Notes of John D. Lynch, 7 March 1968). KU 142950, dorsum yellow and black; venter white and black; iris reddish brown (Field Notes of William E. Duellman, 27 October 1971). KU 142951 – 52, dorsum black with greenish yellow markings; venter black and white (Field Notes of William E. Duellman, 17 October 1971). KU 142953, dorsum green and black; KU 142954, yellow and black (Field Notes of William E. Duellman, 28 October 1971). Lötters (1996: Fig. 11) and Coloma and Ron (2001:Fig. 29) provided color photographs of MHNG 2246.93 from 11 Km ESE of Papallacta. Almendáriz and Cisneros (2005: 155) provided a color description and photograph of this species. Referred specimen EPN 3307 from Provincia Chimborazo had a black dorsum, bearing scattered, small, greenish-yellow marks mostly on the cephalic region; venter with two yellow marks separated by a black stripe that extends toward dorsum (Field Notes of Ana Almendáriz, 19 September 1986). Morphometric comparisons. SVL is not significantly different between females from Provincia Napo vs. Provincia Chimborazo (t = 0.342, df = 9, P = 0.7402). No males from Chimborazo are available for comparisons. Two components with eigenvalues> 1.0 were extracted from the PCA of eight morphometric variables of 26 specimens from populations throughout the range of Atelopus petersi. The axes accounted for 57.5 % of the total variation. Along PC I, the highest loadings were for FL and HLSQ. Along PC II, the highest loadings were for SW and SVL. There is a wide overlap among the morphometric range of these populations. Comments. The Provincia Chimborazo population seems conspecific with Atelopus petersi. However, at this time, further analyses are precluded by the small sample size (three females) from Provincia Chimborazo. Because this small sample size, we preferred to list the Provincia Chimborazo specimens as referred material and not as paratypes. Peters (1973) discussed a series of specimens in the Gustavo Orcés collection (GOV 8698 – 700, currently USNM 236942 – 44) resembling Atelopus petersi (i.e., A. pachydermus sensu Peters, 1973), from the Andes of Provincia Loja in southern Ecuador. He stated that these specimens were identical to those from Oyacachi. However, these specimens are unicolor cream, in contrast to Oyacachi specimens. Because at that time A. petersi (A. pachydermus sensu Peters, 1973) was unknown from the area between Napo and Loja, Peters (1973: 37) suggested that the locality data were erroneous. However, we prefer to assume that the Loja population may refer to a distinct (unnamed) taxon, although they are morphologically similar. Our PCA analysis shows that morphometric variation between the Loja specimens and A. petersi widely overlaps. Our assumption is based on the large geographic gap between A. petersi and the Loja population. Nonetheless, resolution of this issue will require further analyses of Atelopus specimens and populations from intermediate areas and from Loja and Azuay. Distribution, ecology and current population status. According to our data, Atelopus petersi occurs in the Cordillera Oriental of the Andes in Ecuador, Provincias Napo and – according to specimens tentatively referred to the new species – Chimborazo (Fig. 6), at elevations of 2660–3300 m above sea level. The area in which A. petersi occurs includes Cloud Montane Forest and High Montane Evergreen Forest, following the classification of the natural plant formations of the Ecuadorian Andes (Valencia et al., 1999). Annual mean precipitation is about 1000–2000 mm and annual mean temperature is about 12–18 ºC (Cañadas-Cruz, 1983). According to Peters (1973), A. petersi (as A. pachydermus) occurs sympatrically with A. ignescens sensu stricto (Cornalia, 1849). Knowledge on the life history of Atelopus petersi is poor. KU 117874 – 79 were collected on 7 March 1968 under rocks at the edge of a stream. KU 142950 was collected in 27 October 1971 in streambed by day. KU 142951 – 54 were collected in 17, 27 and 28 October 1971 under logs on grassy hillside. The referred specimen EPN 3307 (female) was collected active on cushions plants in paramo habitat at 10: 59 hrs on 19 September 1986. QCAZ 298–302 (two males, three females) were collected on April 1988 during the day when active on a trail. A juvenile (QCAZ 3154) was collected on 30 March 1988. QCAZ 4555 (female) was collected on 19 August 1993 when walking along the border of the Río Oyacachi. QCAZ 7678 was found dead, at 1 Km E of Oyacachi, on 8 November 1996. This is the latest record for this species from Provincia Napo. Despite occasional efforts to locate A. petersi subsequently (one visit to Oyacachi on September 2003, two visits to 11 Km East of Papallacta on January and August 1999, four visits to Atillo on May 2001, on 15 and 30 June 2002, and on April 2006), no additional individuals have been found. According to Peters (1973: 36), in the past, A. petersi was “extremely common” in the vicinity of Papallacta, where it was found along the Río Papallacta, under rocks in moist, and spongy runoff areas. Interviewing local people at Oyacachi and Papallacta, they were familiar with these toads. Several different-aged persons revealed that these toads once were abundant but have not been seen for several years, although in Oyacachi some people claimed, on September 2003, that from time to time they still see single individuals. Remarks. The color pattern of specimen JAP 2267 of Atelopus petersi, and depicted on Figs. 26–27 in Peters (1973) from the environs of Papallacta conforms well to the color pattern of the holotype of A. bomolochos (CAS 91930; Peters, 1973: Fig. 15). This resemblance might be homoplastic, plesiomorphic, or indicate a close phylogenetic affinity between them. Etymology. The specific name is a noun in the genitive case and it is a patronym for James A. Peters, in recognition of his discovery of this species in 1962 and posterior detailed description in 1973. As well, we recognize his contribution to Atelopus taxonomy.