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Guayasamin, Juan M., Krynak, Tim, Krynak, Katherine, Culebras, Jaime, Hutter, Carl R. (2015): Phenotypic plasticity raises questions for taxonomically important traits: a remarkable new Andean rainfrog (Pristimantis) with the ability to change skin texture. Zoological Journal of the Linnean Society 173 (4): 913-928, DOI: 10.1111/zoj.12222, URL: http://dx.doi.org/10.1111/zoj.12222

Pristimantis munozi sp. nov. (Figures 1���3, Table 1) English name: Mu��oz��� Rainfrog Spanish name: Cut��n de Mu��oz Holotype. Adult male, MZUTI 1782 (field number ANF 1129), from Reserva Verdecocha, Province of Pichincha, Ecuador (0�� 5 ' 46.9 "S, 78 �� 36 ' 15.3 "W; elevation 2851 m), collected on 21 of July 2012, by F.J.M. Rojas-Runjaic and J.A. Delgado. Paratopotypes. Seven adult males, MZUTI 1777 ���1781, 1783��� 1784 (field numbers ANF 1124 ���1128, 1130��� 1131 respectively), with the same collection data as the holotype. Diagnosis. A species of the genus Pristimantis, assigned to the Pristimantis myersi species group (sensu Lynch & Duellman 1997, and Hedges et al. 2008), by its small size (SVL canthus rostralis weakly concave in dorsal view, nearly angular in cross-section; (7) upper eyelid with one large and few low tubercles; (8) choanae large, round, not concealed by palatal shelf of maxillary arch; (9) dentigerous processes of vomers low, triangular, slightly smaller than choanae, arranged in V-shaped, posterior and medial to choanae, each bearing one to four teeth; (10) tongue oval, longer than wide, posterior 1 / 4 free, posterior edge feebly notched; (11) males with prominent vocal slits; (12) males without nuptial pads on thumbs; (13) FI shorter than FII; (14) preaxial keels only on FII and FIII, very weak and barely visible; (15) two to four large ulnar tubercles longitudinally aligned on the outer ventrolateral surface of forearm; (16) finger disc and pad absent on FI, pads present and disc slightly expanded on FII to FIV; (17) calcar tubercles absent; heel with few, low tubercles; (18) four to five fold-like tarsal tubercles aligned on the outer ventrolateral surface of tarsus; (19) inner tarsal fold short, low and straight, extending on the distal fourth of the tarsus; (20) outer metatarsal tubercle large, round, swollen; inner metatarsal tubercle oval, elongate, about two times the size of the outer; (21) toes without lateral fringes or keels; (21) toes not webbed; (22) all toes with slightly expanded discs and digital pads; (23) TV slightly longer than TIII; (24) throat white reticulate of black, rest of ventral surfaces pale orange finely punctuated of black. Comparison with similar species. Pristimantis munozi has a small but well-defined tympanum and slightly expanded discs on FII to IV, these traits differentiate P. munozi from P. scopaeus, which lacks tympanum and disc on FII-IV. In P. munozi the tips of finger and toe discs are rounded and present an inner metatarsal tubercle two times the size of the outer and distally not free, which differs notably from the acuminate tips of the discs and the larger (ca. four times the size of the outer) and distally free inner metatarsal tubercle of P. hectus, P. lucidosignatus, and P. onorei. Adult males of P. munozi have prominent vocal slits while there are absent in P. xeniolum and P. floridus; additionally, the new species differs from P. xeniolum by having odontophores and vomerine teeth (absent in the latter), and from P. floridus by having prominent dorsolateral folds, and discs only slightly expanded on fingers and toes (P. f l o r i d u s lacks dorsolateral folds and presents wide discs, about twice the width of the last phalange). Prominent dorsolateral folds, weak preaxial keels on FII and III, and four to five fold-like tarsal tubercles aligned on the outer ventrolateral surface of tarsus distinguishes to P. munozi from P. repens, which lacks dorsolateral folds the dorsum, keels on fingers and tubercles on the tarsus. Pristimantis munozi differs from P. ocreatus by having odontophores on vomer, preaxial keels on FII���III, pads on FII���IV, fold-like tubercles on tarsus, and males with prominent vocal slits (P. ocreatus lacking odontophores on vomers, keels and pads on fingers, tubercles on tarsus, and only with short vocal slits). From P. pyrrhomerus, the new species differs in having prominent dorsolateral folds, preaxial keels on FII���III, weak discs and pads on FII���IV, fold-like tubercles on outer ventrolateral edge of tarsus, dentigerous process of vomers low, males with prominent vocal slits and without red color on groin and concealed parts of thighs (P. pyrrhomerus lacks dorsolateral folds and lateral keels on fingers, discs and pads absent on FI���II, outer tarsal tubercles small and sub-conical, prominent dentigerous process, and males whit short vocal slits and red colored groins and hidden parts of thighs). Pristimantis munozi differs from P. myersi by having dentigerous process on vomers, white ventral color finely spotted of black, with translucent gray groins, and by lacking papilla at tip of snout, lateral keels on toes, and by have a large tubercle on the upper eyelid (P. myersi presents a fleshy papilla at tip of snout, lateral keels on toes, upper eyelid with numerous small tubercles, black ventral color spotted with white, groins with red spots, and lacks dentigerous process of vomers). Pristimantis munozi distinguished from P. f e s t a e by having the dorsal skin shagreen with middorsal raphe, scapular and dorsolateral folds, small tympanum, snout acuminate in dorsal view, canthus nearly angular in cross-section, enlarged tubercle on the upper eyelid, pads on FII���IV, white venter finely spotted with black, and lacking of lateral keels on toes (P. f e s t a e have dorsal skin smooth without middorsal raphe nor dorsolateral folds, present a prominent tympanum, snout rounded in dorsal view, canthus rounded, upper eyelid without enlarged tubercles, fingers lacking pads, toes whit lateral keels, and black venter spotted with white and pale cream). Pristimantis gladiator differs from the new species by lacking middorsal raphe and dorsolateral folds, and having only small tubercles on the upper eyelid and a prominent tympanum; additionally, P. gladiator shows slightly pointed and asymmetrical discs (discs with tip rounded in P. munozi). From P. bicantus the new species is distinguished by having prominent dorsolateral folds, an enlarged tubercle on the upper eyelid, snout acuminate in dorsal view, prolateral keels on FII��� III, ulnar tubercles and outer tarsal tubercles present (P. bicantus usually lacks dorsolateral folds, lateral keels on fingers, ulnar and outer tarsal tubercles, and has small tubercles on the upper eyelid, and rounded snout in dorsal view). At the type locality, P. m u n o z i is sympatric with P. leoni and P. sirnigeli; however, it is distinguished from the former by having one large and rounded tubercle on the upper eyelid, preaxial keels on FII���III, toes without lateral keels, tips of the discs on toes rounded, and fold-like outer tarsal tubercles (P. leoni with numerous enlarged tubercles on the upper eyelid, lateral keels absent on the fingers but present on the toes, tips of the discs on toes slightly pointed, and outer tarsal tubercles small and conical). Finally, P. m u n o z i is distinguished from P. sirnigeli by having a smaller body size in adult males (SVL = 14.9���17.9 mm), prominent dorsolateral folds, relatively short fingers, and by lacking lateral keels on toes (P. sirnigeli with lateral keels on toes, relatively long and thin fingers, and male SVL = 18.6���20.6 mm). Description of the holotype. Adult male of 17.6 mm SVL. Body short and robust (Figs. 1 a���b, 2 a���b). Head slightly narrower than body, slightly longer than wide (Figs. 2 a���b, 3 a���b); HeL 39.5 % of SVL; greatest head width between angles of jaws 38 % of SVL; cranial crests absent. Snout short, acuminate in dorsal view (Fig. 3 a), and rounded in profile (Fig. 3 c); larger than eye (ETS/ED = 1.5); distance eye-nostril slightly shorter than eye diameter (EN/ED = 0.8). Canthus rostralis weakly concave in dorsal view; nearly angular in cross-section; loreal region slightly concave, sloping outward to lip. Not protruding lips. Nares not protuberant, with dorsolateral orientation, barely visible from the front and completely observable in dorsal view. Upper eyelid wide, narrower than interocular region (UEW/IO = 0.8), with one enlarged and rounded tubercle on each eyelid and few additional low tubercles. Tympanum small (TD/ED = 0.4) but well-defined, slightly higher than long; tympanic membrane differentiated; tympanic annulus prominent anteroventrally, posterodorsally concealed beneath skin, barely visible in dorsal view. Supratympanic fold absent, with several large and conical postrictal tubercles below tympanum. Choanae large, round, not concealed by palatal shelf of maxillary arch; dentigerous processes of vomers low, triangular, slightly smaller than choanae, arranged in V-shape, posterior and medial to choanae, each bearing threeto-four teeth. Tongue oval, longer than wide, posterior 1 / 4 free, posterior edge feebly notched. Vocal slits present and prominent. Skin of dorsum and upper eyelids shagreen; middorsal raphe present, but low and barely distinguishable; scapular folds present, sinusoidal and prominent, ending posteriorly in a large conical tubercle; dorsolateral folds straight and prominent, extending from posterior end of scapular folds to level of groins; no paravertebral folds (Fig. 2 a); cloacal sheath absent, small cloacal tubercles present but not conspicuous; flanks and posterior third of dorsum slightly tuberculate; dorsal surface of limbs finely tuberculate. Throat smooth, chest and belly areolate; ventral surface of limbs smooth to slightly areolate (Fig. 1 b). Two-to-three large ulnar tubercles along outer ventrolateral surface of forearm. Hand length 28.6 % of SVL. Relative lengths of adpressed fingers III> IV> II> I; adpressed FI does not reach proximal edge of disc pad of FII; nuptial pads absent; adpresed FIV reaches past distal subarticular tubercle of FIII. Preaxial keels only present on FII and III, very weak and barely visible. Finger disc absent on FI, slightly expanded on FII to IV; digital pad absent on FI, present in all other fingers, rounded, almost longer than wide; circumferential groove of digital pads poorly defined laterally and incomplete proximally; distal edge of discs rounded; disc of FIII about 1.4 times wider than distal end of adjacent phalanx. Subarticular tubercles large, longer than wide, and slightly swollen; palmar tubercle large, slightly swollen, deeply bifid, V-shaped; thenar tubercle large, slightly swollen, elongate; supernumerary tubercles present, relative large (between 1 / 3 ��� 1 / 2 size of subarticular tubercles) (Fig. 3 d). Hind limbs relatively long (thigh length 50.5 % SVL; shank length 56.3 % SVL; foot length 50.5 % SVL); heels overlap when shanks are held perpendicular to sagittal plane; relative length of adpressed toes IV> V> III> II> I; several small, low, and rounded tubercles on heel, calcar tubercle absent; four to five fold-like tarsal tubercles aligned on the outer ventrolateral surface of tarsus; inner tarsal fold short, low and straight, extending on the distal fourth of the tarsus. Tip of TV barely extends past distal edge of mid subarticular tubercle of TIV; tip of TIII reaches distal half of mid subarticular tubercle of TIV. Disc of TIV wider than disc of FIII (F 3 D/T 4 D = 0.8); all toes with slightly expanded discs and digital pads; circumferential grooves of discs absent. Toes without lateral fringes or keels; webbing absent. Outer metatarsal tubercle large, round, swollen; inner metatarsal tubercle oval, about two times the size of outer metatarsal tubercle. Supernumerary plantar tubercles scarce, small, low, and round. Subarticular tubercles large, round and slightly swollen (Fig. 3 e). Color of the holotype in life (based on field notes and color digital photographs; Figs. 1 a���b). Dorsal background color grayish brown, with small and irregular black spots scattered on the dorsal surface of head, upper eyelids, and dorsum; several conspicuous black spots flanking externally the scapular fold. All tubercles and folds on dorsum of head and body with reddish brown coloration. Canthus rostralis with a black canthal stripe, extending from tip of snout to anterior corner of eye. Lips with vertical black and brown bars, except for cream bar just below eye. A conspicuous dark supratympanic stripe extends from upper and posterior parts of the tympanum and fails to reach the shoulder; the inferior border of this stripe is also delineated of creamish white. Flank grayish brown, slightly lighter than dorsum, with scattered small black spots and with one or two diffuse diagonal cream stripes. Groin background translucent gray, with several small immaculate white dots. Upper arm light brown; forearm grayish brown with two-to-three irregular blackish brown cross-bands in the distal half, more intensely pigmented the proximal one. Hand and fingers light brown with three-to-four ill-defined pale brown cross-bands; tips of finger discs reddish brown. Hind limbs background grayish brown, becoming paler distally (on feet), with three to four blackish brown cross-bands on thighs, shanks, tarsi and feet; blackish cross-bands on thighs laterally bordered with light lines; background color of the inner surface of thighs translucent gray, densely spotted of black and with scattered small white dots; transversal rows of tubercles on shanks pigmented of reddish brown. Toes pale orange; toe discs dorsally grayish, bordered by reddish brown coloration. Throat white, finely reticulated with black; chest, belly, undersurface forelimbs of thighs and shanks pale orange, finely punctuated of black; tarsi ventrally grayish brown; palms and soles pale orange; subarticular tubercles of feet bright orange. Iris golden with fine black venation and a broad horizontal copper band; pupil ring golden. Color of the holotype in preservative (after sixteen months, June 2013; Figs. 2 a���b, 3 a���e). Dorsal background almost uniformly gray; reddish brown coloration of dorsal tubercles and folds is replaced by gray and light gray; dorsal and lateral black spots turns more dark and conspicuous. Flanks turn grayish, lighter towards lower portion; diagonal light bands of flanks disappear. Groins and ventral surfaces turn white, with black reticulation. Palms and soles white, with diffuse pink tone and several small black blotches. Fingers and toes with dorsal dark gray, transversal bars, and ventrally colored of white, densely spotted of black. Iris grayish, finely punctuated with black; horizontal copper band turns black. Measurements of the holotype (in mm). SVL: 17.6; ThL: 8.9; SL: 9.9; FL: 8.9; HaL: 5.0; HeL: 6.9; HW: 6.7; InD: 1.9; IOD: 2.1; UEW: 1.7; EN: 1.7; ED: 2.2; ETS: 3.1; TD: 0.9; F 3 D: 0.6; T 4 D: 0.8; F 1 L: 2.6; F 2 L: 3.3. Variation. Females of Pristimantis munozi are unknown; then, sexual dimorphism is not evaluated. We mention, however, that females of all other species of Pristimantis myersi group are larger than males, and several of them have red blotches on the groins (gender-specific coloration). Thus, we suppose that adult females of P. munozi may be are larger than males and may have red flash coloration on the groins, as other species of the group. Adult males range from 14.9���19.7 (16.7 �� 1.6; n = 8). Variation of morphometric characters is shown in Table 1. Head length ranges from 39.1 %��� 42.3 % SVL (40.6 % �� 1.1; n = 8), and maximum head width from 35.4 %��� 42.6 % SVL (38.3 % �� 2.2; n = 8). Head can be as long as wide, or slightly longer than wide (HeL/HW: 1.0 ��� 1.1 [1.1 �� 0.1; n = 8]). The EN/ED ratio is somewhat variable but every eye-naris distance is smaller than eye diameter (0.7���0.9 [0.8 �� 0.1; n = 8]). The width of the upper eyelid is narrower than the interorbital distance, and its ratio ranges from 0.7���0.8 (0.78 �� 0.1; n = 8). The snout length also is slightly variable but always longer than eye diameter (1.2���1.6 [1.4 �� 0.1; n = 8]). Hand length ranges from 26.4���29.7 % SVL (28.2 % �� 1.2; n = 8). The tympanum is small and its ratio respect to eye has little variation (TD/ED: 0.4���0.5; n = 8). Thigh length ranges from 49.3 %���55.0% SVL (51.9 % �� 2.2; n = 8); shank length from 53.6���62.1 % SVL (58.0% �� 2.8; n = 8), and foot length from 48.5 %��� 54.4 % (52.1 �� 2.0; n = 8). In almost all specimens dorsal skin texture is shagreen with scattered tubercles of variable size, except MZUTI 1781, which have dorsal skin nearly smooth, non-tubercular, and with scattered granules on flanks and posterior third of dorsum. Flanks and posterior third of dorsum densely tubercular in MZUTI 1777. Upper eyelid typically with one tubercle; MZUTI 1777 has two tubercles on each upper eyelid. Middorsal raphe absent in MZUTI 1778, but prominent in MZUTI 1779. Scapular folds extending anteriorly and reaching tubercles of upper eyelids in MZUTI 1783, nearly straight and continuous with dorsolateral folds in MZUTI 1779 and 1781. Dorsolateral folds extending to sacral region (MZUTI 1777 ���1778, 1784), to groins (MZUTI 1780���1781) or inclusively converging above the vent level (MZUTI 1779, 1783). Ulnar tubercles on forearms range from two-to-four; tarsal tubercles in number of four or five. Throat typically smooth, but slightly areolate in some specimens (MZUTI 1777, 1780, 1784). Number of vomerine teeth somewhat variable, from one or two (MZUTI 1777���1778) to four (MZUTI 1783), but typically three-to-four (MZUTI 1779���1782). In almost all specimens examined, the color pattern is similar to that of the holotype (Figs. 1 a���b), except specimens MZUTI 1779 and 1781 (Figs. 1 c���d, 2 c���d), which show striking differences. These specimens have a dorsal background pale ocher (pale gray in preservative), extended from the tip of snout to the vent, and medially to the scapular and dorsolateral folds; this area is bordered by a sinuous black stripe that runs from the tip of snout, through nares, canthus, external border of upper eyelid, external flanks of the scapular an dorsolateral folds to upper vent; the middorsal raphe has a light cream coloration and is subtly bordered with pale brown; some small black spots flanking the middorsal raphe in the interorbital and, Published as part of Rojas-Runjaic, Fernando J. M., Delgado, J. Amanda & Guayasamin, Juan M., 2014, A new rainfrog of the Pristimantis myersi Group (Amphibia, Craugastoridae) from Volc��n Pichincha, Ecuador in Zootaxa 3780 (1), DOI: 10.11646/zootaxa.3780.1.2, http://zenodo.org/record/225615, {"references":["Lynch, J. D. & Duellman, W. E. (1997) Frogs of the genus Eleutherodactylus (Leptodactylidae) in western Ecuador: systematics, ecology, and biogeography. The University of Kansas Natural History Museum, Special Publications, 23, 1 - 236. http: // dx. doi. org / 10.5962 / bhl. title. 16289","Hedges, S. B., Duellman, W. E. & Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182.","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 INEFA / GEF- BIRF y EcoCiencia, Quito, pp. 79 - 108.","Neill, D. A. & Jorgensen, P. M. (1999) Climates. In: Jorgensen, P. M. & Leon-Yanez, S. (Eds.), Catalogue of the vascular plants of Ecuador. Monographies in Systematic Botany from the Missouri Botanical Garden, 75, 8 - 13."]}

Rojas-Runjaic, Fernando J. M., Delgado, J. Amanda, Guayasamin, Juan M. (2014): A new rainfrog of the Pristimantis myersi Group (Amphibia, Craugastoridae) from Volcán Pichincha, Ecuador. Zootaxa 3780 (1), DOI: http://dx.doi.org/10.11646/zootaxa.3780.1.2

Pristimantis mazar Guayasamin & Arteaga, new species Holotype. QCAZ 27556 (SC 12660), an adult female (Figs. 1, 2) obtained by Joe Mendelson, Michelle Cummer, and Mart��n Bustamante on February 2004, at La Libertad, Reserva Mazar (02.546589�� S, 78.698375 �� W; 2895 m.a.s.l.), Cant��n Azogues, Provincia Ca��ar, Ecuador; airline distance from Rivera is 6.6 km. Paratypes. Paratypes QCAZ 27553 ��� 54 have the same collecting data as the holotype. Paratypes QCAZ 27493, 27503���05, 27507 ���08, 27511, 27514, 27519, 27536, 27555, 27560 were collected by Joe Mendelson, Michelle Cummer, and Mart��n Bustamante on February 2004, at Rumiloma, Reserva Mazar (02.574616�� S, 78.745462 �� W; 3400 m.a.s.l.), Cant��n Azogues, Provincia Ca��ar, Ecuador. Paratypes QCAZ 29563, 29565, 32619, 49750, 49764 were obtained either by Alejandro Arteaga or Mart��n Bustamante at the type locality in following years. From all these, ten are adult males (QCAZ 27493, 27504���05, 27507, 27519, 27553 ���54, 29563, 29565, 49764) and nine adult females (QCAZ 27503, 27508, 27511, 27514, 27536, 27555, 27560, 32619, 49750). Diagnosis. The new species is placed in the genus Pristimatis, as diagnosed by Hedges et al. (2008), because of the presence of the following traits: cranial crests absent, dentigerous process of the vomers present, and Tshaped terminal phalanges. Pristimantis mazar is diagnosed by having: (1) skin texture of dorsum shagreen with few widely interspersed low tubercles, which became more abundant on flanks and posterior end of dorsum; occipital and dorsolateral folds low or absent; venter areolate; (2) tympanic membrane and tympanic annulus present, with upper rim obscured by supratympanic fold; (3) snout short, pointed in dorsal view and rounded in profile; (4) upper eyelid lacking enlarged tubercles; tubercles are low and almost undistinguishable from surrounding skin texture; cranial crests absent; (5) dentigerous process of vomers well developed, oblique in outline, positioned posterior to level of choanae and separated medially by distance less than width of odontophore, each process bearing 0���7 teeth in females and 0���3 teeth in males; (6) males with a small subgular vocal sac; vocal slits present; nuptial pads absent; (7) first finger shorter than the second; discs on fingers rounded, slightly expanded (Fig. 2); (8) fingers bearing narrow lateral fringes; palmar tubercle bifurcated distally; supernumerary tubercles round, fleshy (Fig. 2); (9) ulnar tubercles low or absent in females, present and low in males; inner tarsal fold present, short; (10) heel and tarsal tubercles low or absent in females, present and low in males; (11) toes bearing narrow fringes; webbing absent; Toe V slightly longer than Toe III; toe discs rounded and slightly expanded (Fig. 2); (12) inner metatarsal tubercle elliptical, about 1.4���2 times the size of outer, rounded metatarsal tubercle; supernumerary plantar tubercles round and fleshy (Fig. 2); (13) in ethanol, dorsum creamish gray to dark brownish gray, with or without darker marks and small white spots; lower flanks with a black-and-white pattern; venter from uniformly whitish cream to cream with darker marking that may form a reticulated pattern. In life, dorsum brown to reddish brown, with or without small white dots and dark brown marks; black spots on groin; flanks and venter cream to gray, usually with blackish mottling that may form a reticulated pattern; iris cream to pale brown, with reddish brown to orange horizontal streak; (14) small SVL in females 19.9���23.7 (mean = 21.5 �� 1.18, n = 10), in males 14.9���18.1 mm (mean = 16.6 �� 1.03, n = 10). Similar species. The smaller, non-overlapping SVL of adult males of Pristimantis mazar readily distinguishes them from most species of the P. o re s te s group (P. atrabracus, P. chimu, P. cordovae, P. corrugatus, P. melanogaster, P. orestes, P. pataikos, P. pinguis, P. simonsii, P. stictoboubonus; Table 1). Similarly, adult females of P. m a z a r are smaller than females of P. bambu, P. corrugatus, P. mariaelenae, P. melanogaster, P. pinguis, P. simonsii, P. stictoboubonus, P. stipe, and P. ventriguttatus (Table 1). The new species differs from P. v i d u a by its ventral color pattern (usually with a reticulated pattern in P. mazar; with minute brown flecking in P. v i d u a), skin texture (dorsolateral folds present in P. v i d u a and usually absent in P. m a z a r), and tympanum (visible in P. m a z a r, concealed beneath skin in P. v i d u a). The call of the new species has a higher dominant frequency (3122���3171 Hz in P. mazar; 2500���2560 Hz in P. bambu) and is longer (0.14 ��� 0.022 s in P. mazar; 0.005��� 0.006 s in P. b a m b u) than the call of Pristimantis bambu. Finally, P. m a z a r differs from its sister species mostly by having a reticulated venter (Fig. 3; uniform in males and females of P. simonbolivari), notorious black stripes on flanks (Fig. 3; uniform dark coloration in P. simonbolivari), and by inhabiting on the Cordillera Oriental of the Ecuadorian Andes (P. simonbolivari is restricted to one locality in the Cordillera Occidental). The two sister species are reciprocally monophyletic (Fig. 7). Description of the holotype. Adult female (QCAZ 27556; Figs. 1, 2). Head as long as wide, slightly wider than body; upper eyelid bearing few low tubercles; head width 38.5 % of SVL; head length 39 % of SVL; snout relatively short (snout to eye distance 15 % of SVL), rounded in dorsal and lateral views; tongue longer than wide, posterior half notched and not adherent to floor of mouth; eye diameter larger than eye���nostril distance; nostrils not protuberant, directed anterolaterally; canthus rostralis weakly concave in profile; loreal region slightly concave; upper eyelid width 68 % of interorbital distance; cranial crests absent; tympanic annulus distinct, except for upper border, which is obscured by supratympanic fold; tympanic membrane distinct; several low postrictal tubercles situated posteroventrally to tympanic annulus; choanae round, not concealed by palatal shelf of maxillary; vomerine odontophores posteromedial to choanae, oblique in outline, about the same size of choana, separated medially by distance less than width of odontophore, each bearing 7���10 teeth; skin on dorsum finely shagreen with interspersed low tubercles; dorsal folds absence, except for thin middorsal fold starting at tip of snout and ending at cloaca; skin of lower flanks and venter areolate; no discoidal or thoracic folds; cloacal sheath absent; two low ulnar tubercles barely evident; outer palmar tubercle oval, large (Fig. 2); subarticular tubercles prominent, round; supernumerary palmar tubercles low and rounded, much lower than subarticular tubercles; fingers bearing narrow lateral fringes; Finger I shorter than Finger II; disc of Finger I not expanded; all other discs slightly expanded, rounded to slightly truncate (Fig. 2); ventral pads defined by circumferential grooves. Tibia length 40.8 % of SVL; foot length 39 % of SVL; tarsal and heel tubercles absent; inner metatarsal tubercle oval, about twice the size of the outer, rounded tubercle; subarticular tubercles round; plantar supernumerary tubercles indistinct; toes bearing narrow lateral fringes; webbing absent; all other toe discs expanded, rounded; toes with ventral pads well defined by circumferential grooves; relative length of toes: I Measurements of holotype (in mm). SVL 21.3; tibia length 8.7; foot length 8.3; head length 8.3; head width 8.2; interorbital distance 2.5; upper eyelid width 1.7; eye diameter 2.3; tympanum diameter 1.0; radioulna length 4.6; hand length 5.1; Finger I length 3.2; Finger II length 3.6; Disc of Finger III width 0.7. Coloration of holotype in preservative. Dorsal surfaces pale brown with dark gray marks, including interorbital bar, W-shaped occipital mark, sacral chevron, as well as thin middorsal line. Arms faintly barred. Thighs cream with black, oblique stripes. Flanks cream with contrasting black marking; throat cream with dark brown flecks. Venter cream with black reticulum (Fig. 1). Coloration in life (based on field notes by Alejandro Arteaga and photographs; Fig. 3). Frogs have a dorsal brown to reddish brown coloration, with or without small white dots and dark brown marks (i.e., interorbital bar, chevrons). Black spots on groin area, indistinct when the animal is at rest. Flanks and ventral surfaces cream to gray, with fine blackish mottling that in most individuals forms a reticulated pattern; ventral portion of fingers light reddish brown. Iris cream to pale brown with reddish brown to orange horizontal streak. Variation. Males present vocal slits and median, subgular vocal sac, and lack nuptial pads. Ulnar and tarsal tubercles are more conspicuous in males than in females. Some males present a thin vertebral fold (QCAZ 27553, 27505) and/or dorsolateral folds (QCAZ 27553, 12602, 27554). Dorsal and ventral color variation in preservative is shown in Figure 4. Vocalization. (QCAZ 49764; adult male recorded on Reserva Mazar on February 2010, temperature not recorded). Pristimantis mazar emits calls at an approximate rate of 9.3 calls per minute (n = 1). Each call is composed by a pulsed, non-frequency modulated note (Fig. 5). Call duration is 0.14 ��� 0.022 s (mean = 0.0185 �� 0.0032; n = 12); intervals between calls vary from 4.89 to 12.24 s (mean = 6.36 �� 2.097; n = 11). The fundamental dominant is at 3122���3171 Hz (mean = 3148.9 �� 12.551; n = 12); the 1 st harmonic is at 6169���6317 Hz (mean = 6244.4 �� 37.792; n = 12) and the 2 nd harmonic is at 9294���9415 Hz (mean = 9328.2 �� 37.358; n = 12). Etymology. This specific epithet mazar refers to the type locality of the species, Reserva Mazar. The epithet is a noun in apposition. Distribution. Pristimantis mazar occurs at elevations between 2895 and 3415 m in three localities within Reserva Mazar (Provincia Ca��ar, Ecuador; Fig. 6): La Libertad (S02.54804, W 78.69741), Rumiloma (S02.57120, S 78.74563), and Gasualpampa (S02.55558, W 78.70150). The area is a patchwork herbaceous paramo, native evergreen montane forest, shrub, secondary forest and pastures. Natural history. At Reserva Mazar, two distinct seasons are recognizable: a dry season (Oct/Nov - Jan/Feb) and a wet season (FCT 2008). Pristimantis mazar is a nocturnal, cryophilic and hygrophilic specialist of herbaceous/shrubby paramos and adjacent high evergreen montane forests. This dominant, and over-abundant rainfrog is active whenever the ambient temperature and humidity levels are favorable (mostly at night), at ground level and on low vegetation (0���67 cm from ground). Diurnal retreats for Pristimantis mazar include grass tussocks, leaf-litter and bromeliads. The species is highly vocal and ubiquitous in open paramo, but considerably less vocal and dominant within unbroken forest. Sympatric species of Pristimantis mazar include P. bambu, P. pycnodermis, P. gagliardoi, and P. r i v e t i. Discussion. The complexity of the genus Pristimantis is exacerbated by its outstanding diversity and recurrent morphological similarities. At this moment, the most recent topologies of Pristimantis (Heges et al. 2008; Pinto- Sanch��z et al. 2012), which contains about 20 % of the described diversity of the genus, shows that the P. o re s t e s group is not monophyletic. Under this hypothesis, similar morphologies (those that diagnose the group) have evolved multiple times, probably because of comparable environments and selective forces acting on species of different clades. Furthermore, at least one additional species groups within Pristimantis (i.e., P. myersi group) shares multiple morphological features that characterize the P. o re s t e s group, also suggesting convergence. Within the P. o re s t e s group, the most closely related species to P. m a z a r is P. simonbolivari (Fig. 7), a species restricted to Caschca-Totoras, on the Ecuadorian Andes. However, our interpretation on the species relationships and evolution of morphological traits in Pristimantis may change as taxon sampling increase and more species are discovered. Currently, we hope that contributions like ours help in having a better understanding of this frog genus, as well as identifying areas of high endemism for conservation proposes. Given the available information on Pristimantis mazar, and following IUCN (2001) criteria, we place this species in the Data Deficient category. As the English common name for this species, we suggest Mazar Rainfrog. We suggest the name Cut��n de Mazar as the common name in Spanish., Published as part of Guayasamin, Juan M. & Arteaga, Alejandro F., 2013, A new species of the Pristimantis orestes group (Amphibia: Strabomantidae) from the high Andes of Ecuador, Reserva Mazar, pp. 345-356 in Zootaxa 3616 (4) on pages 346-353, DOI: 10.11646/zootaxa.3616.4.3, http://zenodo.org/record/218752

Guayasamin, Juan M. (2013): A new yellow species of glassfrog (Centrolenidae: Nymphargus) from the Amazonian slopes of the Ecuadorian Andes. Zootaxa 3651 (1): 193-200, DOI: http://dx.doi.org/10.11646/zootaxa.3652.1.8

Guayasamin, Juan M., Arteaga, Alejandro F. (2013): A new species of the Pristimantis orestes group (Amphibia: Strabomantidae) from the high Andes of Ecuador, Reserva Mazar. Zootaxa 3616 (4): 345-356, DOI: http://dx.doi.org/10.11646/zootaxa.3616.4.3

The Linnaean classification system provides the universal reference system for communicating about the diversity of life and its hierarchic history. Several limitations that challenge the stability of this system have been identified and, as a result, alternative systems have been proposed since its early inception. The revolution caused by molecular phylogenetics has, more than ever, exemplified that Linnaean classification schemes are subject to a degree of instability that may hamper their significance and communication power. Our analysis of recent changes in the classification of several groups of organisms, with a focus on amphibians and reptiles, reveals two main sources of instability: (i) revisionary, objective (empirical) changes based on the discovery of unambiguous instances of non-monophyly and on progress in the Globe's species inventory, and (ii) subjective changes based on author preferences or on a poor analysis of the advantages and limitations of new classification schemes. To avoid subjective taxonomic instability, we review and elaborate proposals for the assignment of Linnaean rank to clades, and thereby for the naming of these clades as Linnaean taxa (Taxon Naming Criteria: TNCs). These are drafted from the perspective of practicing taxonomists and can help choosing among alternative monophyly-based classifications under a premise of economy of change. We provide a rationale for each TNC along with real and theoretical examples to illustrate their practical advantages and disadvantages. We conclude that not all TNCs lead to equally informative and stable taxonomies. Therefore, we order the various TNCs by the generality of their implications and provide a workflow scheme to guide the procedure of taxonomic decisions concerning the creation or modification of supraspecific classifications. The following criteria are considered primary when naming taxa: (i) Monophyly of the taxon in an inferred species tree; (ii) Clade Stability, i.e., the monophyly of a clade to be named as taxon should be as strongly supported as possible by various methods of tree inference, tests of clade robustness, and different data sets; and (iii) Phenotypic Diagnosability, i.e., ranked supraspecific taxa should be those that are phenotypically most conspicuous although in phenotypically cryptic groups of organisms it can be warranted to name taxa based on molecular differences alone. We consider various other criteria as secondary (i.e., the Time Banding, Biogeography, Adaptive Zone, and Hybrid Viability TNCs) and refute using them as sole arguments for the modification of established classifications or proposal of new ones. Taxonomists are encouraged to be explicit and consistent when applying TNCs for creating or modifying classifications. We emphasize that, except for monophyly, the priority TNCs are not proposed as mandatory requisites of a Linnaean taxon but as yardsticks to allow for an informed choice among various clades in a tree that could alternatively be named as Linnaean taxa. Despite a need for plurality, classifications should avoid deliberately violating any of the three primary TNCs because taxa of unstable monophyly or poor diagnosability reduce the information content and hence the utility of the Linnaean system., IDlR worked on this manuscript partially supported by a grant from the Programa Nacional de Movilidad de Recursos Humanos del Plan Nacional de I+D+i 2008-2011 of the Spanish Ministry of Education, and by projects CLG2008-04164 and CLG2011-30393.

Nymphargus lasgralarias Hutter & Guayasamin, new species Figures 4 A��� 4 C; 5 B; 6 A��� 6 D; 7 A��� 7 D; 13 D Holotype. MZUTI 0 96, adult male collected by Carl R. Hutter on 0 5 April 2011 from ���Five Frog Creek��� (0��01.870��� S, 78 �� 42.358 ��� W; 2150 m) at Reserva Las Gralarias, Pichincha province, Ecuador. Figure 5 B. Paratypes. MZUTI 091���095, and 0 97, adult males obtained from Reserva Las Gralarias by Carl R. Hutter. MZUTI 093���095 were collected on 0 5 April 2011; MZUTI 0 92 on 17 April 2011; and MZUTI 0 91 on 18 April 2011 from ���Kathy���s Creek��� (0��01.398��� S, 78 �� 43.772 ��� W; 2000 m). MZUTI 0 97 was collected on 0 1 July 2011 from ���Hercules Giant Tree Frog Creek��� (0��01.529��� S, 78 �� 42.243 ��� W; 2175 m). Generic placement. All species in Nymphargus share an absence of webbing among Fingers I���III and absence or reduced webbing between Fingers III and IV. Additionally, males lack humeral spines (except N. grandisonae). The new species presents the aforementioned traits and, therefore, is placed in Nymphargus (sensu Guayasamin et al. 2009). Diagnosis. The new species can be distinguished from most species of Nymphargus by having a uniformly green dorsum (see Guayasamin et al. 2009). Within Nymphargus, the only species with a green dorsum that lacks spots are: N. cristinae (Ruiz-Carranza & Lynch 1995), N. prasinus (Duellman 1981), and N. wileyi (Guayasamin et al. 2006). Nymphargus lasgralarias sp. nov. is distinguished from N. cristinae by being smaller (male SVL in N. lasgralarias = 24.6���26.5 mm [mean = 25.3 mm; n = 7]; male SVL in N. cristinae = 26.0��� 31.1 mm [mean = 28.0 mm; n = 12]), having a snout that is truncate in dorsal view and protruding in lateral view (subacuminate in dorsal view, truncate in lateral view in N. cristinae; see Ruiz-Carranza & Lynch 1995: Fig. 4), lacking vomerine teeth (present or absent in N. cristinae), and lacking palmar supernumerary tubercles (supernumerary small, abundant in N. cristinae). Nymphargus prasinus differs from N. lasgralarias sp. nov. by having a round snout in dorsal view (truncate N. lasgralarias sp. nov.), 5���7 teeth on each process of the vomer (vomerine teeth absent in N. lasgralarias sp. nov.), and being considerably larger (male SVL 33.0��� 34.5 mm; n = 3; see Duellman 1981). Nymphargus wileyi (an endemic of the Amazonian slopes of the Ecuadorian Andes) is distinguished from N. lasgralarias sp. nov. by having its kidneys covered by a white peritoneum with small, unpigmented spots (see Guayasamin et al. 2006: Fig. 12), whereas in the new species, the kidneys are covered by a homogenously white layer. Additionally, among Nymphargus species found on the Pacific versant of the Andes of Ecuador, Nymphargus lasgralarias sp. nov. could only be confused with N. buenaventura (Cisneros-Heredia & Y��nez-Mu��oz 2007) and N. griffithsi (Goin 1961). Dorsal texture and color pattern readily separates N. buenaventura, which, in life, has a light green dorsum with warts corresponding to pale yellow spots ��� whereas the dorsum of N. lasgralarias sp. nov. is shagreen (lacking warts) and homogenously green (lacking yellow spots). Additionally, N. buenaventura is smaller, although sample size is low (male SVL in N. lasgralarias sp. nov. = 24.6���26.5 mm [mean = 25.3 mm; n = 7]; male SVL in N. buenaventura = 20.9���22.4 mm [mean = 21.8; n = 4]). Nymphargus lasgralarias sp. nov. and N. buenaventura are not known to occur sympatrically. Nymphargus lasgralarias sp. nov. is most similar to N. griffithsi. However, the two species have differences in terms of dorsal color pattern (homogenously green with minute dark melanophores in N. lasgralarias sp. nov. [Figs. 4 B, 6 A��� 6 B]); green with small black spots and/or both minute and small dark melanophores in N. griffithsi [Figs. 4 E, 6 E��� 6 F]), body size (male SVL in N. lasgralarias sp. nov. = 24.6���26.5 mm [mean = 25.3 mm; SD = 0.73 mm; n = 7]; male SVL in N. griffithsi = 22.5���24.2 mm [mean = 23.0 mm; SD = 0.70 mm; n = 5]; T-test: p Advertisement call section). Additionally, in life, N. griffithsi has an iris background coloration of white-silver with larger and less abundant spotting with some medium-dark reticulation (Fig. 7 E��� 7 H), whereas N. lasgralarias sp. nov. has a yellow-golden iris background color with lighter reticulation and more numerous, smaller spots (Fig. 7 A��� 7 D). Characterization. (1) Vomerine teeth absent; (2) snout truncate in dorsal profile, protruding in lateral profile; (3) tympanum small; supratympanic fold present; tympanic membrane translucent, pigmented only on its upper half; (4) skin texture finely shagreen, with microspiculations; (5) ventral skin areolate, with pair of large, round warts on ventral surfaces of thighs below vent; cloaca surrounded by low warts, non-enameled; (6) upper half of ventral parietal peritoneum covered by iridophores (= white), all other peritonea translucent, except for thin layer of iridophores covering heart and renal capsules; (7) liver tetralobed; (8) humeral spines absent; (9) webbing absent between fingers; (10) foot about half webbed; webbing formula: I (2 ��� 2 ���) ��� (2 +��� 2 1 / 2) II (2 ��� 2 ���) ��� (3 ��� ��� 3) III (2 ��� ��� 2) ��� (3 ��� ��� 3) IV (3 ��� 3 +) ��� 2 V; (11) ulnar and tarsal folds low, barely evident, non-enameled; (12) nuptial pad Type I; prepollex not separated from Finger I; (13) first finger slightly shorter than second; (14) eye diameter larger that width of disc on Finger III; (15) in life, green dorsum, with minute dark melanophores; (16) in preservative, dorsum pale lavender; (17) iris golden-yellow, with numerous small black spots; weakly reticulated; (18) hands and feet yellowish green; melanophores absent from fingers and toes or, when present, restricted to dorsal surfaces of Finger IV and Toes IV and V; (19) males call from the upper side of leaves along streams; (20) calls emitted in series of 1���4 calls; each call sounding like a ���tick��� or ���click���; pulsed; duration of 0.0160��� 0.0440 s (mean = 0.0257 �� 0.0058; n = 119); call non-modulated to weakly modulated; dominant frequency at 3445.3���3962.2 Hz (mean = 3691.4 �� 131.9 Hz); (21) fighting behavior unknown; (22) egg clutches deposited on upper surface of leaves at terminal margin, transitioning to hanging as eggs develop; (23) tadpoles unknown; (24) SVL in adult males 24.6���26.5 mm (mean = 25.3 �� 0.737; n = 7); females unknown. Description of holotype. MZUTI 0 96, adult male, SVL 25.5 mm. Head wider than long; head length 32 % SVL; snout truncate in dorsal profile, protruding in lateral view; canthus rostralis indistinct, straight; loreal region slightly concave; lips slightly flared; nostrils protuberant, closer to tip of snout than to eye, directed dorsolaterally; internarial area barely depressed. Eye large, directed anterolaterally at an angle of 45 ��; transverse diameter of disc of Finger III 57.6 % eye diameter. Supratympanic fold conspicuous, obscuring dorsal portion of tympanic annulus; tympanum small (3 % of SVL), oriented mostly vertically, but with slight posterolateral inclination; tympanic membrane transparent, partially pigmented and differentiated from surrounding skin. Dentigerous processes of vomer low, situated transversely between choanae, lacking teeth; choanae large, longitudinally rectangular; tongue ovoid, with ventral posterior third not attached to mouth floor and posterior margin notched; vocal slits extending posterolaterally from the lateral edge of tongue to angle of jaws. Humeral spine absent; ulnar fold low, barely evident, nonenameled; relative lengths of fingers: III> IV> II> I; webbing between fingers absent; discs expanded, nearly round; disc pads triangular; subarticular tubercles small, round, simple; few palmar supernumerary tubercles evident, low; palmar tubercle elliptical, simple; nuptial pad Type I (sensu Flores 1985), ovoid, granular, extending from ventrolateral base to dorsal surface of Finger I, covering the proximal half of Finger I. Length of tibia 56 % SVL; low inner tarsal fold barely evident; outer tarsal fold absent; feet about half webbed; webbing formula of foot: I 2 ��� ��� 2 1 / 2 II 2 ��� ��� 3 III 2 ��� ������ 3 ��� IV 3 ��� 2 V; discs on toes round; disc on Toe IV narrower that disc on Finger III; disc pads triangular; inner metatarsal tubercle large, ovoid; outer metatarsal tubercle round, barely evident; subarticular tubercles small, round; supernumerary tubercles absent. Skin on dorsal surfaces of head, body, and lateral surface of head and flanks shagreen with numerous minute spinules; throat smooth; belly and lower flanks areolate; cloacal opening directed posteriorly at upper level of thighs; cloacal warts small, fleshy, located immediately posterior to cloacal slit, non-enameled. Pair of large subcloacal tubercles evident in ventral aspect. Measurements of holotype. Morphometrics of the holotype and paratypes are summarized in Table 1. Nymphargus lasgralarias sp. nov. Color in life. Dorsum light green, with minute melanophores; flanks yellowish white; bones green; fingers and toes yellow with a faint green tint. Venter white anteriorly and translucent posteriorly. Iris background golden with numerous dark spots and very light reticulation. Color in preservative. Dorsal surfaces of head and body are cream; fingers and toes cream. Upper half of ventral parietal peritoneum covered by iridophores (= white), all other peritonea translucent, except for thin layer of iridophores covering heart and renal capsules. Variation. In life, dorsal coloration varies from very light green to light green. Coloration of dorsum in preservation varies from cream to medium-dark lavender. Females are unknown. Advertisement Call. The call of Nymphargus lasgralarias sp. nov. is reminiscent of a short ���ticking��� or ���clicking��� noise and is easily distinguishable from the significantly longer ���whistle��� produced by Nymphargus griffithsi (Figs. 8���10). The call consists of a short, pulsed note lasting 0.016��� 0.044 s (mean = 0.026 �� 0.006 s) with 1���3 pulses (mean = 1.5 �� 0.6 pulses) (Figs. 9, 11 A��� 11 C). Calls emitted in a series, which typically includes 1���4 calls (mean = 2.7 �� 0.7 calls) (Fig. 12 A��� 12 D). Five-call series had been observed, but were not recorded. Each series has duration of 0.033��� 2.541 s (mean = 1.529 �� 0.597 s) and an interval of 8.6��� 78.6 s (mean 33.8 �� 18.4 s) between series with an interval of 0.088��� 1.513 s (mean = 0.873 �� 0.205 s) between calls within a series. The call repetition rate is 2.0��� 9.9 (5.5 �� 2.7) calls per minute (n = 6 individuals). The dominant frequency is measured at 3445.3���3962.2 Hz (mean = 3691.4 �� 131.9 Hz); contained within the fundamental frequency. The individual call begins at an initial fundamental frequency of 2561.0���3441.0 Hz (mean 3063.6 �� 162.5 Hz). The fundamental frequency is bound between the lower frequency of 2939.4���4145.2 Hz (mean = 3236.3 �� 168.7) and the upper frequency of 3887.7���4473.4 Hz (mean = 4139.8 �� 139.7 Hz). The call has three harmonic frequencies at 6546.1��� 8096.5 Hz (mean = 7298.9 �� 305.8 Hz), 9991.4���12058.6 Hz (mean = 11034.2 �� 478.3 Hz), and 13781.2 ���14928.0 Hz (mean 14317.1 �� 245.5 Hz). A quantitative comparison between the calls of Nymphargus lasgralarias sp. nov. and Nymphargus griffithsi is shown in Table 2. Structurally, the calls of the two species are quite different (Fig. 8). The call of N. griffithsi is a single tonal or multi-pulsed (i.e., 2 or more pulses) call (Figs. 8 B, 10, 11D��� 11 F), whereas the calls of N. lasgralarias sp. nov. are always pulsed (Figs. 8 A, 9, 11 A��� 11 C). Nymphargus griffithsi emits its advertisement call as a single call (absent from a series) while N. lasgralarias sp. nov. emits its calls as a single call or in a series, demonstrating a highly variable calling pattern in contrast to N. griffithsi (Fig. 12 A��� 12 D). In addition, N. lasgralarias sp. nov. has a significantly shorter call duration than N. griffithsi (call duration in N. lasgralarias sp. nov. = 0.016��� 0.044 s [mean = 0.026 s; SD = 0.006 s; n = 119]; call duration in N. griffithsi = 0.103��� 0.148 s [mean = 0.122 s; SD = 0.009 s; n = 48]; T-test: p N. lasgralarias sp. nov. than N. griffithsi (dominant frequency in N. lasgralarias sp. nov. = 3445.3���3962.2 Hz [mean = 3691.4 Hz; SD = 131.9 Hz; n = 119]; dominant frequency in N. griffithsi = 3789.8���4306.6 Hz [mean = 4107.4 Hz; SD = 105.5 Hz; n = 48]; T-test: p N. lasgralarias sp. nov. and N. griffithsi do not show a conspicuous change in dominant frequency, the two species show a slight increase in the dominant frequency, an increase that is more pronounced it N. griffithsi. Furthermore, N. lasgralarias sp. nov. has a significantly lower initialization frequency (Hz) than N. griffithsi (initial frequency in N. lasgralarias sp. nov. = 2561.0���3441.0 Hz [mean = 3063.6 Hz; SD = 162.5 Hz; n = 119]; initial frequency in N. griffithsi = 2821.0���3776.0 Hz [mean = 3328.6 Hz; SD = 300.9 Hz; n = 48]; T-test: p N. lasgralarias sp. nov. and N. griffithsi is shown in Table 2. Additional detailed acoustic measurements can be found in APPENDIX II and APPENDIX III. Distribution. Nymphargus lasgralarias sp. nov. is known only from its type locality at Reserva Las Gralarias (Fig. 1) in Pichincha province, Ecuador, between an elevation of 1850���2200 m. Within the reserve, N. lasgralarias sp. nov. is known from the Chalguayacu Grande River (0��01.868��� S, 78 �� 44.057 ��� W; 1925���2000 m), ���Five Frog Creek���, ��� Heloderma Creek��� (0��01.245��� S, 78 �� 42.370 ��� W; 2175���2225 m), ���Hercules Giant Tree Frog Creek���, ���Kathy���s Creek���, and ���Lucy���s Creek��� (0��00.585��� S, 78 �� 43.901 ��� W; 1850���1875 m). Nymphargus lasgralarias sp. nov. is quite ubiquitous throughout Reserva Las Gralarias, with observations only absent from the Santa Rosa River (0��01.192��� S, 78 �� 43.212 ��� W; 1825���1850 m) (Table 3; Fig. 2). Species N. lasgralarias sp. nov. N. griffithsi Parameter Range Mean �� SD Range Mean �� SD Species Centrolene ballux Centrolene lynchi Centrolene peristictum Centrolene heloderma Nymphargus grandisonae Nymphargus griffithsi Nymphargus lasgralarias Ballux Creek (2150���2200 m) Five Frog Creek (2100��� 2150 m) Heloderma Creek (2175��� 2225 m) Hercules Creek (2150���2200 m) Chalguayacu River (1900��� 1950 m) Kathy��s Creek (1950���2050 m) Lucy��s Creek (1825���1875 m) Santa Rosa River (1800��� 1875 m) Ecology and natural history. Nymphargus lasgralarias sp. nov. inhabits small sized permanent streams (ca. 3 m width) within primary montane forest with minimal disturbance. The species is active during the night and emits advertisement calls from the tops of small sized ferns, small leaves, and long palm leaves 1���6 m above the stream (Fig. 13 D). Nymphargus lasgralarias sp. nov. occurs sympatrically with the following members of Centrolenidae: Centrolene ballux, Centrolene heloderma, Centrolene lynchi (Duellman 1980), Centrolene peristictum (Lynch & Duellman 1973), Nymphargus grandisonae, and Nymphargus griffithsi (Table 3). Other anuran species sympatric along the creeks include: Hyloscirtus alytolylax (Duellman 1972), Pristimantis eugeniae (Lynch & Duellman 1997), Pristimantis calcarulatus (Lynch 1976), Pristimantis parvillus (Lynch 1976), and Pristimantis wnigrum (Boettger 1892). Eggs are deposited on the tips of leaves over the stream and later expand into a hanging gelatinous mass upon absorption of water. We observed 12��� 36 eggs per mass (mean = 25.4 �� 6.0 eggs; n = 23) for N. lasgralarias sp. nov. (Fig. 13 A��� 13 B); we observed a single mass of N. griffithsi eggs containing 14 eggs (Fig. 13 C). The quantity of eggs per mass appears to be highly variable. The egg masses were distinguished by continual monitoring of calling male activity and observed close proximity of calling males. Nymphargus lasgralarias sp. nov. marks the third species of glassfrog in Ecuador with this egg habit type (i.e., eggs dangling from the tips of leaves) ��� after N. griffithsi and N. wileyi (Guayasamin et al. 2006). The Saloya River basin is the type locality for N. griffithsi (Goin 1961), a locality that is about 11 km from the population of N. griffithsi at Reserva Las Gralarias (Fig. 2). These populations are nearly connected through the regional river system ��� the Canchupi River and Saloya River both flow into the Mindo River connecting the two systems, with ca. a 1 km gap between the start of the Canchupi River and ���Five Frog Creek���. It is unknown whether N. griffithsi and N. lasgralarias sp. nov. populations occur in between these two observed localities. At Reserva Las Gralarias, N. griffithsi was observed to occur sympatrically with N. lasgralarias sp. nov. at ���Five Frog Creek���, ���Hercules Giant Tree Frog Creek���, and ��� Heloderma Creek���. Another conspicuous spatial element is the absence of N. griffithsi and presence of N. lasgralarias sp. nov. at ���Kathy���s Creek���, ���Lucy���s Creek���, and the Chalguayacu Grande River, separated from the Santa Rosa River and associated creeks by less than 1 km (Fig. 2). See Figure 2 for a map of the localities and Table 3 for a detailed species account for each creek. Nymphargus griffithsi a, Published as part of Hutter, Carl R. & Guayasamin, Juan M., 2012, A new cryptic species of glassfrog (Centrolenidae: Nymphargus) from Reserva Las Gralarias, Ecuador, pp. 1-21 in Zootaxa 3257 on pages 5-16, DOI: 10.5281/zenodo.280678, {"references":["Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Ruiz-Carranza, P. M. & Lynch, J. D. (1995) Ranas Centrolenidae de Colombia VI. Cuatro nuevas especies de Cochranella de la Cordillera Occidental. Lozania, 63, 1 - 15.","Duellman, W. E. (1981) Three new species of centrolenid frogs from the Pacific versant of Ecuador and Colombia. Occasional Papers of the Museum of Natural History, University of Kansas, 88, 1 - 9.","Guayasamin, J. M., Bustamante, M. R., Almeida-Reinoso, D. & Funk, C. W. (2006) Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics. Zoological Journal of the Linnaean Society of London, 147, 489 - 513.","Cisneros-Heredia, D. F. & Yanez-Munoz, M. H. (2007) A new species of glassfrog (Centrolenidae) from the Southern Andean foothills on the west Ecuadorian region. South American Journal of Herpetology, 2, 1 - 10.","Goin, C. J. (1961) Three new centrolenid frogs from Ecuador. Zoologischer Anzeiger, 166, 95 - 104.","Flores, G. (1985) A New Centrolenella (Anura) from Ecuador, with comments on nuptial pads and prepollical spines in Centrolenella. Journal of Herpetology, 19, 313 - 320.","Duellman, W. E. (1980) The identity of Centrolenella grandisonae Cochran and Goin (Anura: Centrolenidae). Transactions of the Kansas Academy of Science, 83, 26 - 32.","Lynch, J. D. & Duellman, W. E. (1973) A review of the Centrolenid frogs of Ecuador, with descriptions of new species. The University of Kansas Museum of Natural History Occasional Papers, 16, 1 - 66.","Duellman W. E. (1972) A review of the neotropical frogs of the Hyla bogotensis group. Occasional Papers of the Museum of Natural History of the University of Kansas, 11, 1 - 31.","Lynch, J. D. & Duellman, W. E. (1997) Frogs of the genus Eleutherodactylus in western Ecuador: Systematics, ecology, and biogeography. The University of Kansas Museum of Natural History Special Publication, 23, 1 - 236.","Lynch, J. D. (1976) New species of frogs (Leptodactylidae: Eleutherodactylus) from the Pacific versant of Ecuador. Occasional Papers of the Museum of Natural History, University of Kansas, 55, 1 - 33.","Boettger, O. (1892) Katalog der Batrachier-Sammlung im Museum der Senckenbergischen Naturforschenden Gesellshaft in Frankfurt am Main. Frankfurt a. M.: Gebruder Knauer."]}

Centrolene sabini new species (Figures 1���4) Centrolene sp.: Catenazzi et al. 2009: 84���87. Holotype. MUSM 27941, an adult male from Esperanza (13 �� 10 ��� 37 ��� S, 71 �� 36 ��� 31 ��� W, 2800 m), Distrito Kos��ipata, Provincia Paucartambo, Regi��n (Departamento) Cusco, Per��, collected on 9 February 2009 by A. Catenazzi, J. C. Jahuanchi and R. Sotelo. Paratopotypes. MUSM 28017, an adult male from the type locality, collected on 4 January 2010 by A. Catenazzi. MUSM 28018���28019, two adult males ~ 100 m S of the type locality (13 �� 10 ��� 41 ��� S, 71 �� 36 ��� 31 ��� W, 2750 m), collected on 6 January 2010 by A. Catenazzi and J. C. Jahuanchi. Diagnosis. Centrolene sabini is distinguished from species in other centrolenid genera in Peru by having a white ventral parietal peritoneum (transparent in Hyalinobatrachium) and by having humeral spines in males (Fig 1; absent in Cochranella, Hyalinobatrachium, Nymphargus and Rulyrana). Morphologically, the new species is most similar to C. lemniscatum from the eastern slopes of the Andes in Regi��n (Departamento) San Mart��n, 1050 km north of the Kos��ipata Valley (Duellman & Schulte 1993). Both species have humeral spines that are evident and lack vomerine teeth. Centrolene sabini differs from C. lemniscatum (Figs. 1���2; characters of the latter in parentheses) by having distinct yellowish white labial stripes that extend into continuous lateral stripes (white labial stripes not continuous with thin lateral stripes), larger SVL ranging between 29.6���31.2 mm (n = 5) with the smallest male being larger than an adult female of C. lemniscatum (27.2 mm in a female; 27.0 mm in the male holotype), strongly protruding nostrils (directed anterolaterally, but not strongly protruding; Fig. 2), internarial area depressed (barely depressed), snout profile obtuse or inclined anteroventrally (bluntly rounded; Fig. 2), and by its long advertisement call composed of 8���14 peaked notes (short call with a single tonal note in C. lemniscatum, G. Gagliardi- Urrutia, pers. obs.). Additionally, C. sabini lay eggs on the upper side of leaves (Type A of Guayasamin et al. 2009), whereas C. lemniscatum lay eggs on the tip of upper sides of leaves (Type B; G. Gagliardi-Urrutia, pers. obs.). Moreover, the new species has green-yellowish coloration (bright green in C. lemniscatum), large dorsal spicules with yellowish-green tips (smaller, white spicules), and iris silvery-bronze with fine black reticulations (iris white with fewer reticulations). Centrolene sabini could also be confused with the morphologically similar C. buckleyi, which has a call with 1���5 notes at fundamental frequencies of 3520���4139 Hz in Yanayacu, Amazonian slopes of the Andes in Ecuador (Guayasamin et al. 2006); in Colombia, advertisement calls of C. buckleyi are short and composed of a single note (Bernal et al. 2004) with a fundamental frequency of up to 5200 Hz (Bol��var et al. 1999). In contrast, the advertisement call of C. sabini is long and includes 8���14 peaked notes with fundamental frequencies ranging between 2597 and 3919 Hz (Catenazzi et al. 2009; as Centrolene sp.). Morphological characters found in species of Peruvian Centrolene but not in C. sabini include: Finger I longer than Finger II in C. fernandoi, and C. muelleri; dermal fringes on limbs in C. hersperium and C. muelleri; and vomerine teeth in C. azulae and C. fernandoi. Additionally, analysis of the mitochondrial genes 16 S and ND 1 clearly differentiate C. sabini from morphologically similar species found in the Andes (i.e., C. altitudinale, C. buckleyi, C. heloderma, C. hesperium, C. notostictum, C. muelleri, and C. venezuelense; no sequences of C. lemniscatum are available). Genetically, with the current taxon sampling, the most closely related species to C. sabini is C. lynchi (Duellman 1980), an endemic to Colombia and Ecuador; C. lynchi differs from C. sabini by having both dark and light spots on the dorsum and an advertisement call that consists of a tonal note followed by 1���3 peaked notes with higher dominant frequencies (Dautel et al. 2011). Centrolene lynchi further differs from C. sabini in lacking a yellowish white lateral stripe (Duellman 1980), and in having smaller SVL averaging 24.7 mm in males and 24.9 mm in a single female, snout truncate in dorsal view and rounded in profile, dorsal skin shagreened with fine spicules, and different webbing formulas between finger and toes (Duellman 1980). Centrolenids not found in Peru that are morphologically similar to C. sabini include C. altitudinale (endemic to Cordillera de M��rida, Venezuela), C. bacatum, C. ballux (endemic to the Andes of Colombia and Ecuador), C. notostictum, C. quindianum (endemic to the Colombian Andes), and C. venezuelense (endemic to Cordillera de M��rida, Venezuela). Centrolene altitudinale differs from C. sabini by lacking a yellowish white line dividing the upper and lower flanks. Centrolene bacatum is distinguished from C. sabini mainly by having white warts below the eye and being considerable smaller (SVL Centrolene ballux is recognized by having, in life, small greenish white to yellowish white warts and an SVL C. sabini). Centrolene notostictum is also considerable smaller than C. sabini (male SVL C. nostostictum) and lacks the line between the upper and lower flanks that characterizes C. sabini. Centrolene quindianum is easily differentiated by having small black spots on the dorsum (absent in C. sabini). Finally, C. venezuelense is distinguished by its call consisting of 2���4 notes and a dominant frequency at 3900���4400 Hz (Se��aris & Ayarzag��ena, 2005), whereas C. sabini has a call with 8���14 notes and a frequency of 2597���3919 Hz (Catenazzi et al. 2009). Characterization. (1) Males with humeral spines; (2) liver tetralobed, covered by clear peritoneum; (3) parietal peritoneum white; visceral peritoneum translucent; (4) in life, dorsum green with yellowish-green spots and patches, iris silvery-cream with fine black reticulations, yellowish white labial stripe continuous with distinct stripe on flanks, except for area between the edge of the mouth and point of arm insertion, where stripe becomes thin (Fig. 1 A���C); bones green; (5) in preservative, pale lavender with white lateral stripe; (6) dorsal surfaces of males spiculate, with spicules more numerous near orbits; (7) snout obtuse or inclined anteroventrally in lateral profile and rounded in dorsal profile (Fig. 2 A); (8) tympanum small (tympanum diameter 16���23 % of eye diameter), oriented posterolaterally; tympanic annulus indistinct; tympanic membrane pigmented, similarly to surrounding skin; (9) no webbing between Fingers I and II, webbing between Fingers II and III vestigial (Fig. 3 A), webbing formula III 2 �������� 2 ��IV; (10) prepollex not separated from Finger I; in males, nuptial pad Type I (Fig. 1 B, 3 A); (11) Finger I shorter than Finger II (Finger II 110���117 % of length of Finger I); (12) ulnar and inner tarsal folds low; outer tarsal fold absent; (13) webbing on foot moderate (Fig. 3 B), webbing formula: webbing on foot I 1 ����� 2 II 1��� 2 III 2 ��� 2 �� IV 3 ��� 2 ��V; (14) one pair of enlarged tubercles below vent; (15) disc of Finger III of large size, about 56���65 % of eye diameter; (16) vomerine teeth absent; (17) males call from upper sides of leaves; females deposit eggs on upper sides of leaves (Fig. 4); fighting behavior unknown; (18) in adult males, SVL 30.5 �� 0.3 (range 29.6���31.2 mm; n = 5); females unknown. Description of holotype. Adult male, SVL 31.2 mm. Head width 118 % of head length, 32 % of SVL; head length 27 % of SVL; snout rounded in dorsal view and obtuse or inclined anteroventrally in lateral profile; canthus rostralis indistinct, straight; loreal region slightly concave; nostril ovoid, directed dorsolaterally on protuberance, closer to tip of snout than to eye; internarial area slightly depressed. Eye moderately large, ED 40 % of HL, directed anterolaterally; greatest diagonal EW 82 % of IOD; transverse diameter of disc of Finger III 65 % eye diameter. Tympanum small, oriented laterally with slight posterior inclination; tympanic annulus not evident; supratympanic fold moderate, obscuring dorsal portion of tympanic annulus; tympanic membrane pigmented similarly to surrounding skin. Vomerine teeth absent; choanae widely separated and quadrangular; tongue ovoid, with ventral posterior third not attached to mouth floor and posterior margin notched. Vocal sac single, median, subgular; vocal slits extending posterolaterally from the posterolateral base of tongue to angle of jaws. Humeral spine present and visible externally (Fig. 1 A). Hand length 132 % of head length; digits broad, bearing narrow lateral fringes, including outer edge of Finger III and inner edge of Finger I; low ulnar fold visible. Relative lengths of fingers: III> IV> II> I; webbing absent between Fingers I and II and vestigial between fingers II and III; webbing formula for outer fingers: III 2 ����� 2 ��IV; terminal discs subtruncate; width of disc on Finger III 61.5 % of eye diameter; width of disc on Finger I 56.9 % of that of Finger II. Subarticular tubercles broad, nearly as wide as digits, round, elevated; supernumerary tubercles prominent on proximal segments; palmar tubercle moderately large, ovoid with diagonal orientation, elevated, its length 12 % of HnL. Prepollex slightly enlarged; nuptial pad present (Type I of Flores 1985), covering the proximal third of Finger I; prepollical spine absent. Hind limbs long, slender, length of tibia 55 % of SVL, length of foot 49 % of SVL; dermal fringes and tarsal tubercles absent; inner tarsal fold low, outer tarsal fold absent; inner metatarsal tubercle elongate and elliptical, low, measuring 10 % of foot length; outer metatarsal tubercle absent. Toes moderately slender, bearing lateral fringes, including outer edge of Toe V and inner edge of Toe I; length of toes IV> V> III> II> I; toes about three-fifths webbed; webbing formula 1 ����� 2 II 1��� 2 III 2 ��� 2 �� IV 3 ��� 2 ��V. Terminal discs subtruncate; width of digit on Toe I 55 % of that on Toe IV, which is 116 % of that on Finger III; width of disc on Toe IV 86 % of width of disc on Finger IV. Subarticular tubercles as wide as digits, round, elevated. Skin on dorsal surfaces of head and body spiculate; skin on dorsal surfaces of limbs smooth. Flanks smooth dorsally, coarsely granular ventrally. Skin on belly and ventral surfaces of thighs coarsely granular; other ventral surfaces (including throat) smooth. Cloacal opening directed posteroventrally at upper level of thighs; cloacal sheath short and unmodified; cloacal tubercles small, fleshy, located immediately posterior to cloacal slit; para- and subclocal folds absent; one pair of moderately large, round, elevated tubercles on ventral surfaces of thighs below vent. MUSM MUSM MUSM MUSM Character 27941 28017 28018 28019 Color in life. Dorsum green with posterior elongate patches of yellowish-green; tips of spicules yellowishgreen; yellowish-white labial stripe continuous with distinct lateral stripe above insertion of arm extending to groin; flanks below lateral stripe yellow; narrow white stripe on outer edge of forearm and foot; digits pale yellowish-green; iridophores covering between half to two-thirds of the parietal (condition P 3 of Cisneros-Heredia & McDiarmid 2007); heart not visible, iridophores covering the pericardium (condition V 1 of Cisneros-Heredia & McDiarmid 2007); bones green; iris silvery-bronze with fine black reticulations. Color in preservative. Dorsal surfaces of head and body pale lavender, limbs and venter cream; edge of upper lip and white lateral stripe distinct. Dense melanophores in skin of dorsal surfaces of head (including upper eyelid) and body; tips of spicules unpigmented; less dense melanophores in loreal and tympanic regions, on posterior side of dorsum and dorsal surfaces of forelimbs, thighs, shanks, tarsi, and fourth and fifth toes. Flanks, anterior and posterior surfaces of thighs, fingers and inert three toes, and all ventral surfaces are cream and lack melanophores. Measurements (in mm) of the holotype. SVL = 31.2; tibia length = 17.5; foot length = 15.3; head length = 8.4; head width = 9.9; interorbital distance = 3.0; upper eyelid width = 2.5; internarial distance = 2.2; eye-to-nostril distance = 1.8; eye-to-nostril distance = 1.8; eye diameter = 3.4; tympanum diameter = 0.8; eye-tympanum distance = 1.6; radioulna length = 7.0; hand length = 11.0; Finger-I length = 4.7; Finger-II length = 5.5; and disc of Finger III = 2.2 (Table 1). Variation. Measurements of the paratypes are given in Table 1. Only males are known. There is little variation in coloration among these specimens. In MUSM 28017 and MUSM 28018 the spicules are green and cannot easily be distinguished from the background dorsal coloration. Furthermore, in MUSM 28019 the lateral stripe is thin between the edge of the mouth and point of arm insertion. Reproduction and advertisement call. The advertisement call consists of 8���14 peaked notes with emphasized frequencies between 2597���3919 Hz (Catenazzi et al. 2009; as Centrolene sp.). All recorded calls of C. sabini displayed consistent patterns of amplitude modulation that are unusual among centrolenid frogs. Length of calls recorded at 12 ��C varied between 3089���7563 ms when the call included only a series of 6���11 notes, and 8537���10111 ms when the first series of notes was followed by a second series of 3 notes. These two series of notes were separated by intervals of 1459���1652 ms. Advertisement calls were separated by intervals of 258 ms on average. A complete description of the advertisement call on the basis of vocalizations produced by two specimens (MUSM 27941, holotype and MUSM 17966) was provided by Catenazzi et al. (2009). Epiphyllous egg clutches contained between 35 and 45 eggs (n = 5; Fig. 4). Phylogenetic relationships. The phylogenetic analysis of both genes and the combined dataset resulted in similar topologies (Fig. 5). The combined mitochondrial tree and the 16 S topology show that Centrolene sabini is closely related to C. lynchi and C. muelleri (Fig. 5 A, C). The phylogeny based on ND 1 places C. lynchi as the sister taxon of C. sabini (Fig. 5 B; ND 1 sequences were not available for C. muelleri). Our phylogenetic analyses confirm that C. buckleyi is not closely related to the new species, despite similarities in morphology (Bol��var et al. 1999; Bernal et al. 2004; Guayasamin et al. 2006; Catenazzi et al. 2009). Additionally, six species that until now were placed in the genus Centrolene (sensu Guayasamin et al., 2009) based on morphological traits (i.e., ballux, condor, heloderma, huilense, lynchi, and muelleri) show a close phylogenetic affinity with Centrolene species, although taxon sampling only included species of one additional genus (Nymphargus). Therefore, it is likely that placement of these six species in the genus Centrolene is correct. Distribution and ecology. Centrolene sabini is only known from elevations between 2750 and 2800 m in the Kos��ipata valley, upper Manu National Park. The species was first collected and recorded on 28 January 1999 (Catenazzi & Rodr��guez 2001). Males were active and calling at night temperatures ranging between 11.0��� 12.7 ��C. Sympatric frog species include Bryophryne nubilosus, Gastrotheca antoniiochoai, G. nebulanastes, Noblella pygmaea, Oreobates lehri, Pristimantis pharangobates, Psychrophrynella usurpator, Telmatobius sp. (Lehr & Catenazzi 2008, 2009; Duellman et al. 2011). Catenazzi et al. (2011) reported the presence of the fungus Batrachochytrium dendrobatidis in several species of frogs from the Kos��ipata Valley, but in none of the specimens of C. sabini collected in 2009 (n = 1) and 2010 (n = 4). Etymology. The specific name is dedicated to Andrew Sabin, in recognition of his support for herpetology. We propose the following common names: Sabin's glassfrog in English and Rana de cristal de Sabin in Spanish. Discussion. Seven species of Centrolene have been reported from Peru (Fig. 6), but the generic placement of half of these taxa is uncertain. Guayasamin et al. (2009) listed C. azulae and C. fernandoi as ��� incertae sedis within Centroleninae because molecular data are not available, and morphological and behavioral characters do not provide unambiguous evidence on their generic placement���. These two species differ from most other species of Centrolene by having vomerine teeth. A third species, C. lemniscatum, was tentatively assigned to Centrolene by Guayasamin et al. (2009) on the basis of morphology. The new species presents a combination of morphological and behavioral characteristics that is typical of the genus Centrolene (Guayasamin et al. 2009). These traits include the presence of humeral spines and nuptial pads in males, a tetralobed liver covered by a translucent hepatic peritoneum, a white ventral peritoneum anteriorly (concealing the heart) and translucent posteriorly, green bones, melanophores giving a lavender coloration in preservative, males calling from the upper side of leaves, and females depositing eggs on the upper side of leaves along forest streams. On the basis of these characteristics, and the phylogenetic trees presented in Fig. 5, we assign the new species to Centrolene. The discovery of Centrolene sabini considerably extends the geographic range of the genus from the type locality of C. azulae in the Cordillera Central in northern Peru (Flores & McDiarmid 1989) to the Cordillera de Paucartambo in southern Peru. However, another species of Centrolene is known from photographic records taken in a stream at 1750 m in Santuario Nacional Megantoni (12 �� 11 ��� 12 ��� S, 72 �� 28 ��� 12 ��� W; 145 km NW of the Kos��ipata Valley; Rodr��guez & Catenazzi 2005). We are unable to assess whether the geographic ranges of C. sabini and the species from Santuario Nacional Megantoni are disjunct from that of other congeneric species, because the montane forests on the eastern slopes of the Peruvian Andes have been poorly explored. However, along the elevational gradient of the Kos��ipata Valley, intensive sampling of amphibian communities over the past decade revealed the presence of, Published as part of Catenazzi, Alessandro, May, Rudolf Von, Lehr, Edgar, Gagliardi-Urrutia, Giussepe & Guayasamin, Juan M., 2012, A new, high-elevation glassfrog (Anura: Centrolenidae) from Manu National Park, southern Peru, pp. 56-68 in Zootaxa 3388 on pages 58-66, DOI: 10.5281/zenodo.215385, {"references":["Duellman, W. E. & Schulte, R. (1993) New species of centrolenid frogs from northern Peru. Occasional Papers Museum of Natural History University of Kansas, 155, 1 - 33.","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of Glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Guayasamin, J. M., Bustamante, M. R., Almeida-Reinoso, D. & Funk, W. C. (2006) Glassfrogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics. Zoological Journal of the Linnean Society, 147, 489 - 513.","Bernal, M. H., Montealegre, D. P. & Paez, C. A. (2004) Estudio de la vocalizacion de trece especies de anuros del Municipio de Ibague, Colombia. Revista de la Academia Colombiana de Ciencias, 28, 385 - 390.","Bolivar, W., Grant, T. & Osorio, L. A. (1999) Combat behavior in Centrolene buckleyi and other centrolenid frogs. Alytes, 16, 77 - 83.","Dautel, N., Salgado Maldonado, A. L., Abuza, R., Imba, H., Griffin, K. & Guayasamin, J. M. (2011) Advertisement and combat calls of the glassfrog Centrolene lynchi (Anura: Centrolenidae), with notes on combat and reproductive behaviors. Phyllomedusa, 10, 31 - 43.","Senaris, J. C. & Ayarzaguena, J. (2005) Revision taxonomica de la familia Centrolenidae (Amphibia: Anura) de Venezuela. Publicaciones del Comite Espanol del Programa Hombre y Biosfera-Red IberoMaB de la UNESCO.","Flores G. 1985. A new Centrolenella (Anura) from Ecuador, with comments on nuptial pads and prepollical spines in Centrolenella. Journal of Herpetology, 13, 313 - 320.","Cisneros-Heredia, D. F. & McDiarmid, R. W. (2007) Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. Zootaxa, 1572, 1 - 82.","Catenazzi, A. & Rodriguez, L. (2001) Diversidad, distribucion y abundancia de anuros en la parte alta de la Reserva de Biosfera del Manu. In: Rodriguez, L. (ed.). El Manu y otras experiencias de investigacion y manejo de bosques neotropicales. Pro-Manu, Cusco (Peru), pp. 53 - 57.","Lehr, E. & Catenazzi, A. (2008) A new species of Bryophryne (Amphibia: Strabomantidae) from southern Peru. Zootaxa, 1784, 1 - 10.","Duellman, W. E., Catenazzi, A. & Blackburn, D. (2011) A new species of marsupial frog (Anura: Hemiphractidae: Gastrotheca) from the Andes of southern Peru. Zootaxa, 3095, 1 - 14.","Flores, G. & McDiarmid, R. W. (1989) Two new species of South American Centrolenella (Anura: Centrolenidae) related to C. mariae. Herpetologica, 45, 401 - 411.","Rodriguez, L. & Catenazzi, A. (2005) Anfibios y reptiles / Amphibians and reptiles. In: Peru: Megantoni. Rapid Biological Inventories 15, pp. 199 - 204. The Field Museum, Chicago.","Duellman, W. E. (1976) Centrolenid frogs from Peru. Occasional Papers Museum of Natural History University of Kansas, 52, 1 - 11.","Cannatella, D. C. & Duellman, W. E. (1982) Two new species of Centrolenella, with a brief review of the genus in Peru and Bolivia. Herpetologica, 38, 380 - 388.","Duellman, W. E. & Wild, E. R. (1993) Anuran amphibians from the Cordillera de Huancabamba, northern Peru: systematics, ecology, and biogeography. Occasional Papers Museum of Natural History University of Kansas, 157, 1 - 53.","Cadle, J. E. & McDiarmid, R. W. (1990) Two new species of Centrolenella (Anura: Centrolenidae) from northwestern Peru. Proceedings of the Biological Society of Washington, 103, 746 - 768."]}

Oreobates lehri (Padial, Chaparro and De la Riva, 2007) Figures 5C–D, 13B Eleutherodactylus lehri Padial et al., 2007: 115. Oreobates lehri: Padial et al., 2008a: 759. DIAGNOSIS: A slender Oreobates (SVL of adult females 31.0–32.9) characterized as follows: (1) skin on dorsum, flanks, and dorsal surfaces of hind limbs coarsely shagreen, granules regular in size, small, round or conical, few of them slightly enlarged; venter smooth; posterior surfaces of limbs areolate; discoidal fold present; dorsolateral folds irregular, formed by granules, or absent; postrictal glands present, large, prominent; (2) tympanic membrane and annulus distinct, their length about half length of eye; supratympanic fold present, conspicuous; (3) head large, as long as wide; snout round in dorsal and lateral views; canthus rostralis straight in dorsal view, round in section; (4) cranial crests absent; upper eyelid bearing by small granules; (5) dentigerous process of vomers large, situated posteromedial to choanae; (6) males with vocal slits but no nuptial pads; (7) hands with long, slender fingers, finger I shorter than finger II; subarticular tubercles round to conical, well developed; supernumerary tubercles round to conical, smaller than subarticular tubercles; terminal discs of fingers round, not enlarged, lacking circumferential grooves and ungual flap; lateral fringes and keels on fingers absent; (8) ulnar tubercles absent, round, low; (9) no tubercles on heel and tarsus; (10) inner metatarsal tubercle ovate to round, prominent; outer metatarsal tubercle smaller, round, prominent; supernumerary tubercles absent; (11) toes long and slender (foot length 60–70% SVL), lacking lateral fringes or keels (or very weak at base of toe III), webbing absent; fifth and third toes reaching midpoint of second subarticular tubercle of toe IV; tips of toes moderately enlarged, rounded, with ungual flap not indented; (12) axillary glands present; (13) dorsal coloration in life olive green with dark brown to black marks, outlined with pale green, in occipital region and middorsum; belly yellowish white, throat gray with brown mottling or reticulations on anterior margin, hidden surfaces of hind limbs and groin orange. COMMENTS: This species is now restricted to Cordillera Vilcabamba, where it is known from two localities in Departamento Cusco (map 2), the type locality (Apurimac River Valley, Camisea Natural Gas Pipeline, Wayrapata Camp, 2445 m, 12° 50′ 10″ S, 73° 29′ 43″ W) and from 40 km from Vilcabamba, ca. 2850 m, (13°02′51.1″ S, 072°55′29.4″ W) (MUSM 27616). The localities are separated by an airline distance of ca. 60 km. Originally this species was discovered from a series of specimens collected at the first locality and deposited at USNM in Washington. One of the authors (EL), collected a specimen at the second locality, which has been sequenced (fig. 1), and which permitted the description of the coloration in life (fig. 5C–D)., Published as part of Padial, Jose M., Chaparro, Juan C., Castroviejo-Fisher, Santiago, Guayasamin, Juan M., Lehr, Edgar, Delgado, Amanda J., Vaira, Marcos, Teixeira, Mauro, Aguayo, Rodrigo & Riva, Ignacio De La, 2012, A revision of species diversity in the Neotropical genus Oreobates (Anura: Strabomantidae), with the description of three new species from the Amazonian slopes of the Andes, pp. 1-56 in American Museum Novitates 2012 (3752) on pages 38-39, DOI: 10.1206/3752.2, http://zenodo.org/record/5360637, {"references":["Padial, J. M., J. C. Chaparro, and I. De la Riva. 2007. A new species of the Eleutherodactylus discoidalis group (Anura: Brachycephalidae) from cloud forests of Peru. Herpetologica 63: 114 - 122.","Padial, J. M., J. C. Chaparro, and I. De la Riva. 2008 a. Systematics of Oreobates and the Eleutherodactylus discoidalis species group (Amphibia, Anura) based on two mtDNA genes and external morphology. Zoological Journal of the Linnean Society 152: 737 - 773."]}

We revisit species diversity within Oreobates (Anura: Strabomantidae) by combining molecular phylogenetic analyses of the 16S rRNA amphibian barcode fragment with the study of the external morphology of living and preserved specimens. Molecular and morphological evidence support the existence of 23 species within Oreobates, and three additional candidate species (Oreobates sp. [Ca JF809995], Oreobates sp. [Ca EU368903], Oreobates cruralis [Ca EU192295]). We describe and name three new species from the Andean humid montane forests of Departamento Cusco, southern Peru: O. amarakaeri New Species from Río Nusinuscato and Río Mabe, at elevations ranging from 670 to 1000 m in the Andean foothills; O. machiguenga, new species, from Río Kimbiri (1350 m), a small tributary of the Apurimac River, in the western versant of Cordillera Vilcabamba; and O. gemcare, new species, from the Kosñipata Valley at elevations ranging from 2400 to 2800 m. The three new species are readily distinguished from all other Oreobates by at least one qualitative morphological character. Three species are transferred to Oreobates from three genera of Strabomantidae: Hypodactylus lundbergi, Pristimantis crepitans, and Phrynopus ayacucho (for which the advertisement call, coloration in life, and male characteristics are described for first time). Oreobates simmonsi is transferred to the genus Lynchius. Hylodes verrucosus is considered a junior synonym of Hylodes philippi. In addition, H. philippi is removed from the synonymy of O. quixensis and considered a nomem dubium within Hypodactylus. The inclusion of Phrynopus ayacucho in Oreobates extends the ecological range of the genus to the cold Andean puna. Oreobates is thus distributed from the Amazonian lowlands in southern Colombia to northern Argentina, reaching the Brazilian Atlantic dry forests in eastern Brazil, across an altitudinal range from ca. 100 to 3850 m., This work was partially funded by projects CGL2005-03156 and CGL2008-04164 of the Spanish Ministry of Science and Innovation (I. De la Riva, Principal Investigator).

Padial, Jose M., Chaparro, Juan C., Castroviejo-Fisher, Santiago, Guayasamin, Juan M., Lehr, Edgar, Delgado, Amanda J., Vaira, Marcos, Teixeira, Mauro, Aguayo, Rodrigo, Riva, Ignacio De La (2012): A revision of species diversity in the Neotropical genus Oreobates (Anura: Strabomantidae), with the description of three new species from the Amazonian slopes of the Andes. American Museum Novitates 2012 (3752): 1-56, DOI: 10.1206/3752.2, URL: http://www.bioone.org/doi/abs/10.1206/3752.2

Hyloscirtus princecharlesi sp. nov. Figs. 1F, 11A, 12A–C Holotype. CJ 308 an adult male from a cascading stream on the border between Reserva Cotacachi-Cayapas and a private land owned by Manuel Quinchiguango, at Recinto San Antonio (ca. Cuellaje), Provincia Imbabura, Ecuador (0º 28' 23.772" N, 78º 34' 12.756" W; 2794 m); obtained on 22 July 2011 by Elicio E. Tapia, Carl R. Hutter, and Carlos Quinchiguango. Paratypes. QCAZ 43654, 44893 adult males with same locality as holotype (0º 28' 20.8194" N, 78º 33' 57.6" W; 2720–2794 m); the former obtained on 23 June 2009 by Elicio E. Tapia and William Quinchiguango, and the latter on 24 June 2009 by Luis A. Coloma, Manuel A. Morales-Mite, and Elicio E. Tapia. Diagnosis. A member of the Hyloscirtus larinopygion group as diagnosed by Faivovich et al. (2005). The new species differs from other members of that group by having a unique pattern of dorsal coloration (series of well defined orange blotches or spots densely and uniformly distributed on a black background), a genetic distance from other members of the group of at least 1.31% (for fragments of ~ 2.3 kb of aligned mitochondrial DNA sequences), poorly expanded digital discs, and a glandular nuptial pad on thumb. The most similar dorsal coloration is seen in H. pantostictus (Fig. 1E) and H. ptychodactylus (Fig. 1H), both of which have orange spots or blotches. Hyloscirtus princecharlesi differs from H. pantostictus from the slopes of northeastern Ecuador by having gray digital discs (yellow in H. pantostictus); also, the two species are not sister taxa (Fig. 3). The new species is distinguished from its sister species, H. ptychodactylus from the slopes of western Ecuador at Cotopaxi province by having well defined spots or blotches on dorsum (scattered, diffuse, spots or blotches and dense stipling among blotches in H. ptychodactylus), and by having a gray iris color (sky-blue in H. ptychodactylus). Description of holotype. SVL 70.5 mm. Body and limbs robust. Snout nearly truncate in dorsal and lateral view. Head about as broad as long. Head width at level of eyes (21.8 mm), 30.9% of SVL. Canthus rostralis rounded. Loreal region slightly concave. Lips rounded, not flared. Dorsal surface of internarial region nearly flat. Nostrils barely protuberant, directed anterolaterally, slightly posterior to anterior margin of lower jaw. Top of head nearly flat. Tympanum vertically ovoid, tympanic annulus distinct. Supratympanic fold thick, curved, covering dorsal edge of tympanum, extending from eye to posterior end of mandible and to shoulder. Forearms robust compared to upper arms. Axillary membrane absent. Ulnar tubercles absent. Fingers broad. Disks round, barely expanded or Finger I about same width as finger. Relative lengths of fingers 1 Skin on throat and anterior portion of chest bearing irregular scattered folds on a weakly areolate skin that extends to abdomen. Pelvic patch areolate. Transverse supracloacal flap long. Margins of vent with numerous small folds and two large lateral swollen glandular areas at proximal posterior thighs. Cloacal opening directed posteriorly. Vocal slits present at posterior lingual margins of mandibles. Dentigerous processes of vomers long, transverse, abutting medially, behind level of large, ovoid choana, bearing 24 teeth evident in the buccal mucosa. Tongue broad, cordiform, shallowly notched posteriorly, fully attached to mouth floor, rugose on its anterior portion. Vocal sac single, median, subgular. Color in preservative (~70% ethanol). Dorsum with a pattern of dense, brown-orange, round-oval marks, and blotches on a black background, forming a reticulated pattern at mid-dorsum and more isolated blotches towards the flanks. Dorsum of limbs with larger marks and blotches than body. Anterior and posterior surfaces of thighs nearly black with two large faint gray blotches. Dorsum of inner fingers and toes gray with faint creamy-gray blotches that become brighter toward Fingers IV and Toes IV–V. Glandular area lateral to vent brown-orange. Throat, chest, abdomen with diffuse black-gray marbling. Inner margin of lower lip cream. Anterior flanks with three large round marks, posterior flanks barred. Tympanum mostly black. Color in life (Fig. 6B). Same as above except in that dorsal marks and blotches are orange, varying to pale orange. Blotches on hidden surfaces of thighs are creamy white. Ventral surfaces are black with yellow-cream marbling, more yellow at gular region. Tips of dorsal surfaces of fingers and toes are pale creamy-orange with gray. Palmar and plantar surfaces are gray to black. Ventral pads of digital discs on fingers and toes are gray. Iris is dark gray. Measurements of holotype (mm). SVL 70.5, TIBL 32.8, FEL 31.5, FOL 31.5, RDUL 17.9, HANDL 24.7, THBL 16.7, HLSQ 23.9, HDW 23.4, ITN 6.0, EYD 7.2, EYNO 4.7, TYD 3.6, DFW 3.9, DTW 3.1. Variation. Measurements variation of two paratypes (QCAZ 44893, 43654) and the holotype (CJ 308) are indicated in Table 7. The adult male paratypes overall are very similar to the holotype except for differences in color strength and patterns (Fig. 14), by bearing 29 vomerine teeth in QCAZ 44893 and 23 in QCAZ 43654, and by having slightly smaller finger discs in QCAZ 44893. Tadpoles. See under Tadpoles and ontogeny section. Distribution, natural history, and conservation status. Hyloscirtus princecharlesi is known only from its type locality (San Antonio, ca. border of Reserva Ecológica Cotacachi-Cayapas), a cascading stream in Montane Cloud Forest (Fig. 11C, D) (according to the classification proposed by Valencia et al. 1999). The locality is in northwestern Ecuador in the Cordillera de Toisán, a mountain range that is part of the Cordillera Occidental of the Ecuadorian Andes, in Provincia de Imbabura, at an elevation of 2720–2794 m (Fig. 9). At the type locality, the annual mean precipitation is 1671 mm and the annual mean temperature is 14.1 ºC (based on the WorldClim database; Hijmans et al. 2005). Toral et al. (2002) provide a detailed description of the type locality and a record of a female of this species (under the name Hyla sp 1.). The holotype CJ 308 was active during the night (21:50h) 2 m above the ground, on bushy vegetation, containing decomposing leaves mixed with mosses, and epiphytes, and surrounded by dense natural vegetation. It was located at the upper part of a cascade that was about 10 m wide and 8 m high. The headwater springs are about 20 m higher than the collecting site and about 80 m from the nearest divide. Paratypes QCAZ 44893 and 43654 were found active at 19:50h and 21:30h, respectively, at about the same site as the holotype. QCAZ 44893 was 1 m above ground on a leaf (approximately 30 x 40 cm) of an Anthurium sp., at the margin of a stream with natural vegetation. QCAZ 43654 was climbing on a branch 2 m above ground, close to a natural wall of stones, with abundant earth and epiphytic vegetation. There was no rain, the forest was cloudy. QCAZ 43654 was found at about 10 m from an individual of Hyloscirtus criptico . Hyloscirtus princecharlesi herein is considered as Endangered (A3ce IUCN criteria) due to a suspected population size reduction of ≥ 50% suspected to be met within the next 10 years, where the reduction or its causes may not have ceased. The single locality currently known for this species is being modified by human activities and is being severely affected by growing habitat destruction. Threats are logging, burning, unregulated use of land for agriculture, cattle raising, pesticide use, and invasive trout in the regional streams. Besides, it is likely that climate change and emerging pathogens are affecting its populations as has been documented for numerous other Andean frogs (Pounds et al. 2010). Rapid and integrative conservation measures are urgently needed, among which the protection and restoration of its habitat are a priority, as well as the establishment of an ex-situ assurance colony. Etymology. The specific name princecharlesi is a patronym that honors His Royal Highness Charles, Prince of Wales (Charles, Philip, Arthur, George, Windsor). In his call to halt tropical deforestation, Prince Charles uses frogs as symbols, and his Rainforests SOS Campaign includes a video with a frog as a rainforest ambassador. For this reason he is affectionately known by the media as the ‘frog prince’. Prince Charles is contributing significantly to the growth of awareness in the battle against tropical deforestation, climate change, and the catastrophic extinction of rainforest amphibians. His work is leading to increased awareness of these issues, and this increased awareness benefits biodiversity conservation, sustainability, alleviation of poverty, and ensures ecosystem services for present and future generations.

Pristimantis bambu Arteaga-Navarro & Guayasamin, new species Holotype. QCAZ 46740, an adult male (Figs. 1���3) obtained by Alejandro F. Arteaga-Navarro on January 0 7, 2010, at La Libertad, Reserva Mazar (02.54804 S, 78.69741 W; 2976 m.a.s.l.), Cant��n Azogues, Provincia Ca��ar, Ecuador; airline distance from Rivera is 6.6 km. Paratypes. All 24 paratypes (Figs. 4 A, B) were collected by Alejandro F. Arteaga-Navarro on January and March 2010 at the Reserva Mazar, southern Sangay National Park, east of Pallcayacu river, within the Cant��n Azogues, Provincia Ca��ar, Ecuador. From these, 16 are adult males (QCAZ 46704, 46706���7, 46710���12, 46724��� 27, 46741���42, 47555, 47558, 47556��� 57); 6 are adult females (QCAZ 46705, 46708, 46739, 46743���44, 7559); and 2 are juvenile females (QCAZ 46713 ��� 14). Diagnosis. The new species is placed in the genus Pristimatis, as diagnosed by Hedges et al. (2008), because of the presence of the following traits: cranial crests absent, dentigerous process of the vomers present, and Tshaped terminal phalanges. Pristimantis bambu is diagnosed by having: (1) skin texture of dorsum shagreen with few widely interspersed low tubercles; middorsal fold barely noticeable and fragmented when present (Fig. 1 A); dorsolateral folds ill-defined and incomplete, sometimes absent; venter areolate, lacking discoidal or thoracic folds (Fig. 1 B); (2) tympanic membrane absent; tympanic annulus present, 40���71 % of eye length in females and 42��� 65 % in males, with upper rim obscured by supratympanic fold; (3) snout short, acuminate in dorsal view and rounded in profile; (4) upper eyelid lacking enlarged tubercles; when present, tubercles are low and almost undistinguishable from surrounding skin texture; upper eyelid width 49���83 % of interorbital distance; cranial crests absent; (5) dentigerous process of vomers well developed, oblique in outline, positioned posterior to level of choanae and separated medially by distance less than width of odontophore, each process bearing 4���10 teeth; (6) males with a small subgular vocal sac; vocal slits present; nuptial pads absent; (7) first finger shorter than the second; discs on fingers expanded and rounded to slightly truncated (Fig. 2 A), except for Finger I that is only barely expanded; (8) fingers bearing narrow lateral fringes; outer palmar tubercle divided at least distally, inner portion larger and elliptical and outer portion smaller and round; supernumerary tubercles round (Fig. 2 A); (9) ulnar tubercles coalesced into fold; (10) no heel or tarsal tubercles; (11) toes bearing narrow fringes; webbing absent; Toe V slightly longer than Toe III; toe discs expanded, rounded to slightly truncated (Fig. 2 B); (12) inner metatarsal tubercle elliptical, about 1.2���2 times the size of outer, rounded and enlarged metatarsal tubercle; supernumerary plantar tubercles round and weakly developed (Fig. 2 B); (13) in ethanol, dorsum pale brownish gray in males (blackish or slate gray in females), with minute dark spots on the scapular and sacral region (Fig. 3), and faint barring on the dorsal surfaces of limbs (absent in females); dark postocular stripe and faint supralabial bars present; flanks with some localized rusty tinge on the axilla, groin and hidden surfaces of hind limbs; pale cream spots enclosed by blackish pigment may be present on the groin and posterior surfaces of the thigh (bolder and invariably present in females); ventral surfaces uniformly grayish tan with very fine, dark, localized mottling on throat. In life (Fig. 3), dorsal surfaces of males tan, with or without dark symmetrical markings overall (uniformly blackish olive drab in females); dark transverse bars on the hind limbs poorly marked if present (absent in females); black postocular blotch present; in males, background color of throat, belly and ventrolateral surfaces khaki, with or without a yellowish tint on the throat that may shift to orange-red on the belly; rest of underparts, grayish brown to rosy brown; in females, all ventral surfaces uniform gray or deep purplish brown; a rusty tinge on the hidden surfaces of hind limbs is present in both males and females, almost always surrounding one or several amber to orange blotches in the axilla, groin and posterior surfaces of the thigh; these marks may be enclosed by black pigment; upper half of the iris pale goldenrod yellow; lower half, deep coppery brown; (14) SVL in females 24.6���26.4 (mean = 25.3 �� 0.6, n = 5), in males 17.4���20.1 (mean = 19.2 �� 0.7, n = 13); (15) body mass in living females 0.9���1.6 g, in males 0.5��� 0.8 g. Similar species. Pristimantis bambu differs from all other members of the P. o re s t e s group (i.e., P. a t r a b r a - chus, P. chimu, P. cordovae, P. corrugatus, P. melanogaster, P. orestes, P. pataikos, P. pinguis, P. seorsus, P. simonbolivari, P. simonsii, P. stictoboubonus, P. vetriguttatus, P. vidua; sensu Hedges et al. 2008) by having slightly expanded, rounded to slightly truncate finger pads. Pristimantis corrugatus (Duellman & Lehr 2009) may as well have expanded discs, but these are elliptical and wider than those of P. bambu. The new species may be further distinguished from all other species in the group (Table 1), except from P. c h i m u, P. pinguis, P. seorsus and P. s i m o n s i i (Duellman & Lehr 2009), by having ulnar tubercles coalesced into a low fold. Pristimantis chimu, P. seorsus and P. simonsii differ from P. b a m b u by lacking vocal slits; P. pinguis differs by having a differentiated tympanic membrane. From those species in the group bearing small eyelid tubercles (P. c h i m u, P. cordovae, P. o re s t e s, and P. v e n - triguttatus: Duellman & Lehr 2009; Lynch 1979), it differs by lacking a distinct discoidal fold. Finally, in life, P. bambu is unique among the species assigned to the P. ore s te s group on the basis of the rusty pigment of the hidden surfaces of the hind limbs and the bold, black-bordered amber blotches on the groin and posterior surface of the thighs. Description of the holotype. Adult male (QCAZ 46740; Figs. 1���3). Head slightly narrower than body, slightly longer than wide; upper eyelid bearing several, feebly visible tubercles; head width 35 % of SVL; head length 36 % of SVL; snout relatively short (snout to eye distance 17 % of SVL), acuminate in dorsal view (Fig. 1 A) and rounded in profile; tongue longer than wide, posterior half notched and not adherent to floor of mouth; eye diameter slightly larger than eye���nostril distance; nostrils not protuberant, directed anterolaterally; canthus rostralis weakly concave in profile; loreal region slightly concave; upper eyelid width 58 % of interorbital distance; cranial crests absent; tympanic annulus distinct, round; tympanic membrane absent; two enlarged postrictal tubercles, situated posteroventrally to tympanic annulus; choanae round, not concealed by palatal shelf of maxillary; vomerine odontophores posteromedial to choanae, oblique in outline, about the same size of choana, separated medially by distance less than width of odontophore, each bearing 5 teeth; vocal slits and median, subgular vocal sac present; skin on dorsum shagreen with widely interspersed low tubercles; dorsolateral fold visible on either side but only on anterior half of the body; middorsal fold fragmented, barely noticeable (Fig. 1 A); skin on venter areolate with no discoidal or thoracic folds (Fig. 1 B); cloacal sheath absent; cloacal region bordered ventrally by small, closely packed warts; ulnar tubercles coalesced into fold, continuous throughout forearm; outer palmar tubercle divided, inner portion larger and elliptical, outer portion smaller and round (Fig. 2 A); subarticular tubercles round in section; supernumerary palmar tubercles low and rounded, much lower than subarticular tubercles; fingers bearing narrow lateral fringes; Finger I shorter than Finger II; disc of Finger I barely expanded; all other discs expanded, rounded to slightly truncate (Fig. 2 A); ventral pads well defined by circumferential grooves; nuptial pads absent. Tibia length 47 % of SVL; foot length 45 % of SVL; inner metatarsal tubercle present, about the same size of the outer, rounded and enlarged tubercle; subarticular tubercles round in section; plantar supernumerary tubercles indistinct; toes bearing narrow lateral fringes; webbing absent; discs of Toe I barely expanded; all other toe discs expanded, rounded to slightly truncated; toes with ventral pads well defined by circumferential grooves; relative length of toes: I Measurements of holotype (in mm). SVL 20.04; tibia length 9.4; foot length 9.0; head length 4.5; head width 7.2; eye diameter 2.3; interorbital distance 2.6; upper eyelid width 1.5; internarial distance 2.0; eye-nostril distance 1.8. Coloration of holotype in preservative. Dorsal surfaces pale brownish gray, with minute dark dots on the scapular and sacral region, distributed symmetrically on each half of the body, and with faint, dark transverse bars on the hind limbs (Fig. 1 A); upper lip with dark brown bars below eyes; dark brown supratympanic stripe present; background color of ventral surfaces khaki with a localized rusty tinge on the hidden surfaces of the hind limbs, which is also present but much fainter on the lower belly, palms and axillary region (Fig. 1 B). Coloration of holotype in life (based on observations in the field and photographs; Figs. 3 A, B). Dorsal surfaces light orangish brown, with faint, dark transverse bars on the hind limbs; flanks with same coloration as dorsum, with a dark brown postocular stripe accompanied by faint vertical supralabial stripes; background color of ventral surfaces khaki, finely mottled with dark pigment on the throat and strongly tinged with yellow on the throat and upper chest; palms, soles, ventral surfaces of hind limbs and groin intensely tinged with rusty to yellow, which becomes deep saffron on the belly as it fuses with the yellowish pigment of the throat; upper half of iris light goldenrod, lower half coppery brown. Variation. Meristic variation is presented in Tables 2 and 3. Some paratypes have two eyelid tubercles (QCAZ 46705, 46714, 46712, 46739, 46711, 26724), and others have only one (QCAZ 46725, 46704, 41444, 41606, 41616, 41620, 41672, 41694, 41738; Figs. 4, 5 B); several paratypes have no visible dorsolateral fold (QCAZ 46706, 46714, 46725, 46711, 46742, 46724) nor middorsal fold (QCAZ 46725, 46711, 46742, 46724). In ethanol, the dorsum of one paratype is densely covered throughout with symmetrically distributed dark marks (QCAZ 46742; Fig. 6 A), another one has a pale gray middorsal spot (QCAZ 46714), and yet another one (QCAZ 46710) has a light drab frontal patch. The dorsal background color of three paratypes is darker than the holotype (QCAZ 47555, 47556, 47557; Fig. 6 D) or completely dark grayish brown altogether (QCAZ 46707); three male paratypes (QCAZ 46712,47555, 47557) have no localized rusty to light reddish brown pigment on the groin, axilla and hidden surfaces of the hind limbs. Most male paratypes (QCAZ 47555, 46726, 47556, 47558, 47557, 46711, 46712, 46714 46725, 46713, 46724, 46727, 46706, 46742) have pale cream spots on the groin and posterior surface of the thighs. Throat mottling is much more extensive in two male paratypes (QCAZ 46706, 46707). In preservative, the dorsal color becomes grayer and the rusty and yellowish tinge of the ventral surfaces is extensively lost. Specimens in life have well-defined tubercles and folds on the dorsum and legs, but those skin marks are much fainter in preservative. Vocalization. The advertisement call of QCAZ 47558 (an adult male recorded on Reserva Mazar on 11 March 2010 at 22:08 h, temperature not recorded) can be characterized as a series of short, indistinctly pulsed, non-modulated notes in amplitud. The dominant frequency ranges between 2.50 and 2.56 kHz (maximum energy is 122.54 �� 1.74 dB; n = 20). The note repetition rate is 54.42 notes per minute, and the note length ranges between 0.005 and 0.006 s (Fig. 5). Internote intervals range between 0.816 and 1.782 s (mean = 1.195 �� 0.310 s; n= 20). Out of six recordings, call duration ranged between 12 and 103 s, with significant variation among note repetition rate. Vocalizing males decreased note intensity when approached by an observer. This is the first call described for a species in the P. o re s t e s group. Etymology. This specific epithet bambu refers to the habitat preference of the new species, which is found in bamboo-dominated montane forests (Chusquea sp). The epithet is a noun in apposition. Distribution and natural history. Pristimantis bambu is known only from La Libertad, Reserva Mazar, (S 2.54804, W 78.69741), at elevations between 2876 and 2989 m.a.s.l (Fig. 6). This site, a family-owed alpaca ranch, is located in the eastern mountain range of Ecuador, on the southern edge of Sangay National Park, Ca��ar province. The area is a patchwork of native evergreen montane forest (Valencia et al. 1999), shrub, secondary forest and pastures (FCT 2008). Average temperatures at La Libertad are 18 ��C and precipitation is 1200 mm per year. There are two distinct seasons: the dry season (Oct/Nov ��� Jan/Feb) and the wet season (FCT 2008). Individuals of P. bambu were active during night on leaf litter and vegetation (20���155 cm above ground) within regrowth montane forest where bamboo (Chusquea sp) is dominant. Individuals were seen active under overcast and cloudy nights, even under light drizzle, but not under heavy showers. During early morning, P. bambu was found active on the leaf litter, later during the day, individuals were found hidden in the leaf litter. Two individuals were hidden under piled up eucalyptus lumber and dead leaves far from the forest border, in pastures. Pristimantis bambu was found both near rivers and far from significant watercourses; but it was never found within primary forest. The species appears to be closely associated with secondary forests where the canopy cover is not complete, a habitat composed mainly of small trees, shrubs and bamboo. The species was also found on the forest border. Females Males (n = 5) (n = 13) SVL 24.6���26.4 (25.3 �� 0.6) 17.4���20.1 (19.2 �� 0.7) Tibia length 11.9���12.4 (12.1 �� 0.2) 8.7���9.8 (9.3 �� 0.3) Foot length 11.3���11.7 (11.5 �� 0.2) 7.5���9.1 (8.4 �� 0.4) Head length 8.3 ���9.0 (8.6 �� 0.3) 6.1���7.2 (6.7 �� 0.3) Head width 8.4���9.3 (8.7 �� 0.3) 6.2���7.1 (6.6 �� 0.3) Interorbital distance 3.0��� 3.3 (3.1 �� 0.1) 2.2���2.6 (2.4 �� 0.1) Upper eyelid width 1.6���2.1 (1.9 �� 0.2) 1.5 ���2.0 (1.7 �� 0.1) Radioulna lenght 5.4���6.3 (5.8 �� 0.3) 4.2���4.7 (4.5 �� 0.1) Eye to nostril distance 2.2���2.4 (2.3 �� 0.1) 1.5���1.9 (1.7 �� 0.1) Snout to eye distance 4.0��� 4.3 (4.1 �� 0.1) 3.0��� 3.6 (3.3 �� 0.2) Eye diameter 2.5���2.7 (2.6 �� 0.1) 2.1���2.4 (2.2 �� 0.1) Tympanum diameter 1.0��� 1.5 (1.3 �� 0.2) 0.7���1.3 (1.1 �� 0.2) Hand lenght 6.1���6.4 (6.2 �� 0.1) 4.3���5.2 (4.9 �� 0.2) Finger I lenght 3.6 ���4.0 (3.7 �� 0.2) 2.6���3.1 (2.9 �� 0.2) Females Males During March 2010, a vocalizing male was heard. When searching it, a female Pristimantis bambu was found, presumably attracted by the male's vocalization. They were both bagged together and began amplexus 72 min afterwards. During the wet season, five males were heard vocalizing simultaneously along with another species (Pristimantis pycnodermis). A gravid female released 30 unfertilized eggs inside a plastic collecting bag. Sympatric species of P. b a m b u include, P. pycnodermis, P. gagliardoi, P. riveti, and two undescribed Pristimantis species. During night, P. bambu remains still when approached by an observer; calling males may continue to vocalize even when approached, though some individuals may stop calling or foraging, even jumping away at times. At night, most individuals were found by standing 0.4���1.8 m from the vegetation and carefully scanning thin (2���10 mm in diameter) twigs, ferns, and leafs. Discussion. The most recent comprehensive phylogeny of strabomantids includes four species assigned to the Pristimantis orestes group; two of them, P. o re s t e s and P. simonbolivari, cluster together, but the other species (P. simonsii and P. melanogaster) do not (Hedges et al. 2008). Two scenarios explain the observed pattern: (i) the molecular phylogeny is accurate and the shared morphologies of the species included in the Pristimantis orestes group represent convergent evolution to cope with similar habitats (Hedges et al. 2008), or (ii) the molecular phylogeny is not congruent with the evolutionary history of the these amphibians, and the Pristimantis orestes group is monophyletic. Since only a molecular phylogeny is at hand at the moment, we favor the first scenario. Both morphology and molecules support the lineage distinctiveness of P. b a m b u. Given our taxon sampling, P. bambu is most closely related to Pristimantis sp. (Fig. 7; corrected genetic distance = 5.9���6.8 %), P. simonbolivari (corrected genetic distance = 7.7���7.8 %), and P. o re s t e s (corrected genetic distance = 8.9 %; Fig. 7)., Published as part of Arteaga-Navarro, Alejandro F. & Guayasamin, Juan M., 2011, A new frog of the genus Pristimantis (Amphibia: Strabomantidae) from the high Andes of Southeastern Ecuador, discovered using morphological and molecular data, pp. 17-29 in Zootaxa 2876 on pages 19-27, DOI: 10.5281/zenodo.205032, {"references":["Hedges, S. B., Duellman, W. E. & Heinicke, P. (2008) New world direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182.","Duellman, W. E. & Lehr, E. (2009) Terrestrial breeding frogs (Strabomantidae) in Peru. Natur und Tier - Verlag, Naturwissenschaft, Munster, 384 pp.","Lynch, J. D. (1979) Leptodactylid frogs of the genus Eleutherodactylus from the Andes of southern Ecuador. Miscellaneous Publications, Museum of Natural History, University of Kansas, 66, 1 - 62.","Valencia, R., Ceron, C., Palacios, W. & Sierra, R. (1999) Las formaciones naturales de la Sierra del Ecuador. In: Sierra, R. (Ed.), Propuesta Preliminar preliminar de un sistema de Clasificacin clasificacion de Vegetacion vegetacion para el Ecuador Continentalcontinental. Proyecto INEFAN / GEF - BIRF and Ecociencia, Quito, Ecuador, pp. 79 - 108.","FCT. (2008) Nudo del Azuay Conservation Initiative: An Introduction to the Dudas Watershed. Fundacion Cordillera Tropical, Cuenca, Ecuador."]}

Atelopus podocarpus sp. nov. Holotype. QCAZ 6801, adult female, from Lagunas del Compadre (4 �� 10 ' S, 79 ��06' W; ca. 3300 m), Parque Nacional Podocarpus, Provincia Loja, Ecuador, collected on 1 December 1994 by Jenny Rudston. Paratypes. EPN 1223 ��� 26, from Lagunas del Compadre (4 �� 10 ' 20 " S, 79 ��06' 42 " W; 3400 m), Parque Nacional Podocarpus, Provincia Loja, Ecuador, no collection date stated on field notes, but possibly collected on September 1981, by Luis H. Albuja V.; EPN 2143 ��� 45, from Cerro Toledo (4 �� 23 ' S, 79 ��06' W; 2900 m), Provincia Loja, Ecuador, collected on 11 September 1981 by Luis H. Albuja V.; EPN 3235, 3237, Mirador path nearby the Centro Administrativo Cajanuma of the Parque Nacional Podocarpus (3 �� 58 '09" S, 79 ��08'06" W; 2850���2990 m), Provincia Loja, Ecuador, obtained on 16���18 November 1986 by Ana Almendariz; KU 120389, from 15 km east of Loja (2700 m), Provincia Zamora Chinchipe, Ecuador, obtained on 10 June 1968 by John D. Lynch; KU 165004 ��� 5, from 13 km east of Loja (3 �� 58 ' S, 79 ��06' W), Abra de Zamora, Provincia Loja, Ecuador; KU 178418, from Provincia Zamora Chinchipe: 13.7 km east of Loja (3 �� 58 ' S, 79 ��06' W), Provincia Loja, Ecuador; KU 201115 ��� 7, from 3.7 km south of Saraguro (3 �� 38 ' 49 " S, 79 �� 14 ' 42 " W; 2800 m), collected on 0 8 March 1984 by J. E. Simmons; QCAZ 2838 (cleared and double stained skeleton), from Centro Administrativo Cajanuma (4 ��07' S, 79 ��09' W; 2800 m), Parque Nacional Podocarpus, obtained on 25 June 1987 by Luis A. Coloma, Mario Garc��a Saltos, and Renato Le��n. Referred specimens. KU 196633 ��� 40, from west slope Cerro Chinguela on Huancabamba���San Ignacio trail, Departamento Piura, Peru, obtained on 14���15 October 1977 by Gary R. Graves; KU 196641 ��� 42, from Cerro Chinguela, 5 Km NE Zapalache, 2900 m, Departamento Piura, Peru, obtained on 13 and 20 July 1980 respectively, by L. J. Barkley. Diagnosis. (1) A species with mean SVL in adult females 47.7 mm (39.5���52.9, SD = 3.1, n = 22) and in adult males 37.7 mm (34.4���40.9, SD = 2.6, n = 5); (2) hind limbs short, tibia length/SVL 0.322���0.424 (n = 27); (3) phalangeal formula of hand 2 - 2-3 - 3; webbing absent; (4) foot webbing formula I( 0)���(1) II( 1 1 / 2)��� (3) III( 1)���(3 ���) IV (2 2 / 3)���(1 +) V; (5) snout acuminate, protruding beyond lower jaw; (6) tympanic membrane and tympanic annulus absent; (7) dorsal surfaces of body smooth; arms and legs surfaces warty; ventral surface weakly aerolate; (8) red-orange spiculae (cream in preservative) on flanks and some extending to the dorsum; (9) vertebral neural processes inconspicuous; postorbital and cranial crests present; (10) dorsum entirely black, flanks with or without cream marks; (11) no minute gray stippling on dorsum of body; (12) venter and throat pale green and cream (cream or yellow in preservative), with or without dark marks; (13) gular region lacking warts, spiculae, or coni. Atelopus podocarpus sp. nov. is most easily confused with A. petersi from the north and east-central flanks of the Ecuadorian Cordillera Oriental of the Andes. The two species have similar color patterns and body proportions. However, A. podocarpus can be easily differentiated by having numerous minute brownred-orange spiculae (in life) on the flanks (18���71 spiculae on the flanks between the arms and legs counted in a straight line); in contrast, A. petersi has white spiculae aggregated into few small warts (1���21 spiculae per wart; 8���15 warts on flanks between arms and legs counted in a straight line) (Figs. 10, 11 A���B). Atelopus podocarpus differs from A. bomolochos (from the southern Andes of Ecuador) and A. pachydermus O. Schmidt (from south Andean Ecuador and northern Peru) in the dorsal coloration in life; A. podocarpus has a black dorsum, whereas A. pachydermus has a yellow and brown dorsum and A. bomolochos has a yellow to dark green dorsum (Coloma et al. 2007: Fig. 3 A���D). Atelopus peruensis Gray and Cannatella (from northern Peru) is slightly smaller than A. podocarpus (SVL in A. peruensis = 32.8���38.5 in adult males; 38.4���45.2 mm in adult females; Venegas 2005), has, in life, black flanks with white warts (red flanks in A. podocarpus), and a non-protruding snout (protruding in A. podocarpus). Atelopus podocarpus and A. ignescens (from the northern Andes of Ecuador) are similar in size and coloration (in preservative), but A. ignescens bears a patch of spiculae and coni on the gular and pectoral regions of females (absent in A. podocarpus). Genetically, A. podocarpus is clearly differentiated from A. ignescens, A. bomolochos, A. onorei, and A. peruensis. A molecular phylogenetic analysis shows A. halihelos Peters as the sister species of A. podocarpu s (Guayasamin et al. 2010). These two species are differentiated by size (male SVL of A. podocarpus = 34.4���40.9 vs 25.3 in holotype of A. halihelos), color pattern (in preservative, black dorsum in A. podocarpus, light brown dorsum in A. halihelos), the presence of a discontinuous dorsolateral line of warts in A. halihelos (absent in A. podocarpus), and flanks with some warts with multiple spiculae in A. halihelos (warts never have more than one spiculae in A. podocarpus). Description of holotype. (Figs. 12 A���C, virtual animation). Female. Head about as long as wide; head length and head width less than one third SVL (HLSQ /SVL = 0.276; HDWD /SVL = 0.278); snout acuminate in dorsal view, protruding beyond anterior margin of jaw; nostrils slightly protuberant; canthus rostralis distinct; loreal region concave, lips flared; interorbital region flat and smooth, eyelid without tubercles; postorbital and cranial crests present; tympanic membrane and tympanic annulus absent, tympanic area warty; postorbital and cranial crests. Forearm relatively short (RDUL /SVL = 0.291); palmar tubercle round; supernumerary palmar tubercles present; subarticular tubercles not prominent; digital tips with round pads; thumb relatively long (THBL / HAND = 0.677) having two phalanges; hand webbing absent; fingers lacking lateral fringes; length of fingers III( 0)��� (1) II 1 (1 / 2)���(3) III( 1)���(3 ���) IV( 2 2 / 3)���(1 +) V; webbing becoming fringe-like distally; length of toes IMeasurements of holotype (mm): SVL 50.8, TIBL 17.0, FOOT 19.5, HLSQ 14.0, HDWD 14.1, ITNR 4.5, EYDM 4.5, RDUL 14.8, HAND 12.4, THBL 8.4, SW 11.0 Variation. Meristic variation is given in Table 7. Low warts on the arms and legs are evident is some specimens (EPN 1225, 2143, 3235, 3237, KU 201115, 201117) and absent in others (EPN 1223, 1224, 2144, KU 201116); some individuals lack warts in the tympanic area (EPN 1223, 3237); three specimens have spiculae on the tympanic area (EPN 3235, KU 201116 ��� 17). TABLE 7. Measurements (in mm) of the type series of Atelopus podocarpus. Mean �� standard deviation (range). Abbreviations follow Gray and Cannatella (1985) and Coloma et al. (2000). They are: SVL = snout���vent length; TIBL = tibia length; FOOT = footh length; HLSQ = head length; HDWD = head width; EYDM = eye diameter; EYNO = eye���nostril distance; ITNA = internarial distance; RDUL = radio-ulna length; THBL = thumb length; SW = sacrum width. All measurements are in mm. Ventral coloration varies among individuals (Fig. 13 A���D); some have a black thick longitudinal band on the middle of the throat (EPN 1223, 1225); one individual has black throat and venter with cream marks (EPN 2143); one individual has black throat with two cream marks at the sides at the level of the articulation of the jaw (EPN 2144); generally the inner part of thighs are cream, but they may be black (EPN 2143); the inner part of shanks can be black (EPN 1223 ���25, 2143��� 44), cream (EPN 3237) or black with cream marks (EPN 3235, KU 201115 ��� 7). The venter varies from cream to yellowish (KU 201115 ��� 7). Fingers and toes are mostly black (except for the tips), but Finger II (EPN 1223 ���25, 2143���44, 3235, 3237) and Toe I (EPN 1225, 2144, 3237) and Toe II (EPN 1223 ��� 24) are entirely cream in some individuals. The inner surface of the arms is black (EPN 2143), cream (EPN 1223 ���24, 3235, 3237, KU 201115, 201117) or black with cream marks (EPN 1225, 2144, KU 201116). The dorsum and flanks may be the same color, but the flanks may have some cream marks (EPN 2144). KU 196633 ��� 42: Dorsum and flanks uniform black; throat, chest, belly, and ventral surfaces of limbs dull cream. Palmar surfaces dull cream to pale brown, but black with cream palmar tubercles in KU 196633, 19636, and 196641 ��� 42. Tips of fingers cream and inner and outer metatarsal tubercles cream in most specimens. Color in life. (Fig. 10 A���B). The following is based on color slides taken by AA (EPN 3235, dorsolateral view) and Luis H. Albuja (EPN 1223, ventral view) at Parque Nacional Podocarpus: Dorsum and dorsal surfaces of arms and legs black; flanks and spiculae in flanks orange red; face black; belly light green, throat cream with black mark in the center; palms and inner surface of arms cream; inner surface of thighs cream. KU 165004 (field color notes and slide in dorsolateral view taken by WED on 7 March 1975) from Abra de Zamora, 13 km E Loja, 2850 m: Dorsum dull brownish black with dull red tubercles laterally on body. Venter green with tan streak on throat and black stripe on throat and chest. Palmar and tarsal tubercles and tips of digits white. Iris dark brown with bronze wing around pupil. Distribution, ecology and current population status. Atelopus podocarpus is only known from paramo and subparamo habitats in the southern Cordillera Oriental of Ecuador (Provincias Loja and Zamora Chinchipe) to Departamento Piura in Cordillera de Huancabamba in northern Peru (Fig. 5). In Ecuador, it occurs in the following Natural Formations (classification proposed by Valencia et al. 1999): Herbazal Lacustre Montano, P��ramo Arbustivo, Bosque Siempre Verde Montano Alto, and Bosque de Neblina Montano. These habitats lie between 2700���3400 m (Fig. 5). Annual mean rainfall is 1000���2000 mm and annual mean temperature is 7���18 ��C (Ca��adas-Cruz, 1983). The area of its extent of occurrence is of about 2475 km 2. Habitat and other features of the Podocarpus National Park were described by Rivera-Rosi (2007). KU 120388 ��� 89 (John D. Lynch field notes, 10 June 1968) from Abra de Zamora, 15 Km E Loja, 2800 m were found under stones on dirt banks. KU 165004 (WED field notes, 7 March 1975) from Abra de Zamora, 13 km E Loja, 2850 m was found under a rock by day, in elfin, wind-swept scrub with a thick layer of moss, many lichens and liverworts, bromeliads on ground and on bushes, and bamboo. During the day of collection, the temperature was 8.5 ���18.0˚C and rainfall was 6 mm. Atelopus podocarpus is considered to be Critically Endangered (Possibly Extinct) (A 2 ace, IUCN Red List categories and criteria). The species is tagged as Possibly Extinct until further surveys confirm otherwise. The population has declined dramatically (more than 80 %) in the last two decades probably because of climate change and the impact of pathogens, which have affected many other montane species of Atelopus. The amphibian chytrid fungus has not been reported yet from Loja or Zamora Chinchipe provinces. The closest record of the chytrid is Azuay Province (Ron and Merino 2000). The risk factor of potential threat caused by the chytrid for anuran amphibian species calculated by R��dder et al. (2009: Fig. 2 C) is high at the area of its distribution. In Ecuador, the last living individual (QCAZ 6801) was seen on 1 December 1994. It was collected in an unhealthy condition at Lagunas del Compadre in Parque Nacional Podocarpus; the animal died while being transported. Collecting efforts at several localities of its distribution (Abra de Zamora in September 2001, August 2003, December 2009; nearby Saraguro in June 2008, Lagunas del Compadre in November 2009) revealed no individuals. In Peru, no individuals have been found since 20 July 1980. Etymology. The specific name is a noun in apposition and refers to the Podocarpus National Park, where this species formerly lived. Podocarpus takes its name from having the country's largest contingent of the Podocarpus or romerillo tree, the only conifer native to Ecuador. Ecuador's southernmost national park ranges from 1000 meters in the river valleys to 3600 meters in the higher reaches of the Nudo de Sabanilla mountain range, part of the larger Cordillera Oriental. This park of 360,000 hectares shelters an array of cloud forests, subparamo, paramo, a series of small lakes, and many endemic species of flora and fauna., Published as part of Coloma, Luis A., Duellman, William E., C, Ana Almend��riz, Ron, Santiago R., Ter��n-Valdez, Andrea & Guayasamin, Juan M., 2010, Five new (extinct?) species of Atelopus (Anura: Bufonidae) from Andean Colombia, Ecuador, and Peru, pp. 1-54 in Zootaxa 2574 on pages 28-34, DOI: 10.5281/zenodo.197448, {"references":["Coloma, L. A., Lotters, S., Duellman, W. E. & Miranda-Leiva, A. (2007) A taxonomic revision of Atelopus pachydermus, and description of two new (extinct?) species of Atelopus from Ecuador (Anura: Bufonidae). Zootaxa, 1557, 1 - 32.","Venegas, P. (2005) Atelopus peruensis. In: Rueda-Almonacid, J. V., Rodriguez-Mahecha, J. V., Marca, E. L., Lotters, S., Kahn, T. & Angulo, A. (Eds.) Ranas Arlequines. Conservacion Internacional. Panamericana Formas e Impresos S. A., Bogota, Colombia, 150 pp.","Guayasamin, J. M., Bonaccorso, E., Duellman, W. E. & Coloma, L. A. (2010) Genetic differentiation in the nearly extinct harlequin toads (Bufonidae: Atelopus), with emphasis on the Atelopus ignescens and A. bomolochos species complexes. Zootaxa, 2574, 55 - 68.","Gray, P. & Cannatella, D. C. (1985) A new species of Atelopus (Anura, Bufonidae) from the Andes of northern Peru. Copeia, 910 - 917.","Coloma, L. A., Lotters, S. & Salas, A. W. (2000) Taxonomy of the Atelopus ignescens complex (Anura: Bufonidae): Designation of a neotype of Atelopus ignescens and recognition of Atelopus exiguus. Herpetologica, 56, 303 - 324.","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 / GERF-BIRF y Ecociencia, Quito, 79 - 108 pp.","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.","Rivera-Rosi, J. (2007) Parque Nacional Podocarpus. In: ECOLAP & MAE (Eds.) Guia del patrimonio de areas naturales protegidas del Ecuador. ECOFUND, FAN, Darwin Net, IGM, Quito, Ecuador, 185 - 192 pp.","Ron, S. R. & Merino, A. (2000) Amphibian declines in Ecuador: overview and first report of chytridiomycosis from South America. Froglog, 42, 2 - 3."]}

Atelopus orcesi sp. nov. Holotype. MHNG 2684.75, adult gravid female, from La Alegr��a-Sibundoy (0˚ 34 ' N, 77 ˚ 31 ' W; 2400 m), Provincia de Sucumb��os, Ecuador, collected in May 1988, by Ana Mar��a Velasco. Paratype. MHNG 2559.67, adult male with same data as holotype. Diagnosis. (1) A medium-sized species with SVL in adult male 29.4 mm and in adult female 42.1 mm; (2) hind limbs short, tibia length/SVL 0.354���0.415; (3) phalangeal formula of hand 2 - 2-3 - 3 (inferred), basal webbing absent; (4) foot webbing formula I (0-0+)���(��- 1) II (0+)���(1 -) III (��- 1 -)���(��) IV (��)���(0�����) V; (5) snout acuminate, protruding beyond lower jaw; (6) tympanic membrane absent; (7) dorsal surfaces of body smooth, and warty on limbs; (8) row of yellow creamy warts on dorsolateral body of male, flanks with few, scattered, minute spiculae on female; (9) vertebral neural processes inconspicuous; (10) dorsum light to dark brown (in preservative), bearing a rudimentary X-shaped mark anteriorly and scattered small marks posteriorly; (11) minute gray stippling absent on dorsum of body, but minute white stippling present; (12) venter cream (in preservative) with a brown mark on gular-chest region; (13) gular region without warts, spiculae or coni. Atelopus orcesi sp. nov. is distinct from most species of Atelopus by the combination of two features, i.e., bearing an X-shaped mark on anterior dorsum and by male having a conspicuous dorsolateral row of warts on the body. Only A. laetissimus Ruiz-Carranza, Ardila-Robayo, and Hern��ndez-Camacho, A. mucubajiensis Rivero, A. nahumae Ruiz-Carranza, Ardila-Robayo, and Hern��ndez-Camacho, A. pinangoi Rivero, and A. sanjosei Rivero and Serna, share these two features. Atelopus cruciger Lichtenstein and Martens and A. walkeri Rivero also have these features, but females of these species have a conspicuous dorsolateral row of warts on the body (absent in female A. orcesi). Atelopus orcesi is mostly similar to two specimens (Atelopus sp. 16; Rueda-Almonacid 2005 a) from Departamentos del Choc�� and Valle from western Colombia. Although they are geographically separated and on opposite sides of the Andes, and they also differ by female of A. orcesi having a brown band on flank, the possibility that they are conspecific remains to be further investigated with a larger sample size. The morphological character that differentiates them might be polymorphic. Atelopus orcesi also is similar to A. nepiozomus Peters from Provincia Morona-Santiago in southeastern Ecuador. Both have a similar ventral color pattern. Atelopus orcesi is larger (female holotype SVL: A. orcesi = 42.1 vs A. nepiozomus = 32.4) and lacks warts on dorsum (many low, rounded warts and pustules, which may be spiculate in A. nepiozomus). Atelopus orcesi differs from Atelopus laetissimus, A. mucubajiensis, A. nahumae, and A. pinangoi by the contrasting color of the row of warts (not contrasting color in A. laetissimus, A. mucubajiensis, A. nahumae, and A. pinangoi), and by the female having a pale brown band on the flank (band on flank absent in A. laetissimus, A. mucubajiensis, and A. pinangoi). Furthermore, A. orcesi is geographically distant and geographically isolated from A. laetissimus and A. nahumae in the Sierra Nevada de Santa Marta in Colombia, and from A. mucubajiensis and A. pinangoi in the Sierra de Santo Domingo in the Cordillera de M��rida of the Venezuelan Andes. Atelopus orcesi differs from the smaller A. sanjosei (female SVL A. orcesi = 42.1 vs females SVL A. sanjosei = 31.5���32.7 mm, n = 2) (from the eastern versant of Cordillera Central in Departamento Antioquia, Colombia) by female of A. orcesi lacking the dorsolateral row of warts (present in female A. sanjosei) and by having a brown mark on gular-chest region (absent in A. sanjosei). Description of holotype. (Figs. 7 A, 8 A, 8 C, 8 D, virtual animation). Female. Head about as long as wide, HLSQ and HDWD less than one third SVL (HLSQ /SVL = 0.263, HDWD /SVL = 0.256); snout acuminate, its margin triangle-shaped with a round anterior vertex in dorsal view; profile of tip of snout in lateral view nearly straight and 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 slightly flared; interorbital and occipital regions flat, smooth; eyelid flared without tubercles; postorbital crest slightly raised, not glandular; low pretympanic and post-tympanic areas warty; tympanic membrane and tympanic annulus absent; temporal area smooth; choanae small, rounded, widely separated (29 % of HW); tongue about twice as long as wide, its posterior half not attached to floor of mouth. Forearm relatively short (RDUL /SVL = 0.264); palmar tubercle round; thenar tubercle nearly indistinct, supernumerary palmar and subarticular tubercles absent; digital tips with round pads; thumb relatively long (THBL / HAND = 0.615), apparently having two phalanges; webbing on hands absent, fingers lacking lateral fringes; relative length of fingers III( 0+)���(1 -) II( 0+)���(1 -) III( ��)���(��) IV( ��)���(��) V; relative length of toes IMeasurements of holotype (mm): SVL 42.1, TIBL 14.9, FOOT 14.4, HLSQ 11.1, HDWD 10.8, ITNR 4.3, EYDM 3.5, EYNO 2.6, RDUL 11.1, HAND 9.6, THBL 5.9, SW 12.4. Variation. (Figs. 7 B, 8 B, virtual animation). The male paratype has the following measurements: SVL 29.4, TIBL 12.2, FOOT 11.5, HLSQ 9.5, HDWD 9.1, ITNR 3.5, EYDM 3.1, EYNO 2.5, RDUL 8.6, HAND 7.1, THBL 4.0, SW 9.0. The male paratype resembles the holotype with the following exceptions. Sexual dimorphism is evident in that it is smaller than the female. It has vocal slits and keratinized nuptial pads on the dorsal and inner surfaces of the thumb. Forelimbs are relatively long and slender in female. A tubercle is absent on the forearm of the male, which lacks spiculae on the flanks. The male bears a dorsolateral row of conspicuous warts from the suprascapular to the inguinal regions on the body (see discussion), and has a more protuberant snout, bearing a keeled tip. Color variation in preservative (~ 70 % ethanol): The dorsum is paler than in the female. The X-shaped mark on the head is more continuous, and there are more scattered marks on posterior dorsum. The marks are nearly black, and there is no defined band on the flanks. Color in life. Unknown. Tadpoles. Unknown. Distribution, ecology and current population status. Atelopus orcesi is known only from the type locality on the eastern versant of Cordillera Oriental in Provincia Sucumb��os, Ecuador (Fig. 5). The type locality is at about 2400 m above sea level in Montane Cloud Forest (Valencia et al. 1999). At the type locality, annual mean temperature is 14.2 ��C and annual mean precipitation is 1199 mm (Hijmans et al. 2005). Atelopus orcesi is considered to be Critically Endangered (Possibly Extinct) (A 2 ace, IUCN Red List categories and criteria). The species is tagged as Possibly Extinct until further surveys confirm otherwise. The population probably declined dramatically (more than 80 %) in the last two decades because of climate change and the impact of pathogens, which have affected many other montane species of Atelopus. The amphibian chytrid fungus has been reported in very close proximity to the type locality near Santa B��rbara in Sucumb��os Province (= Carchi Province of Ron and Merino 2000). The risk factor of potential threat caused by the chytrid for anuran amphibian species calculated by R��dder et al. (2009: Fig. 2 C) is high at the area of its distribution. No individuals have been reported after its discovery in May 1988, despite at least one intensive search effort of about 40 person-hours at the type locality and surroundings at elevations of 2000���2800 m above sea level, in December 2009 by Elicio Tapia and Samael Padilla of QCAZ. Etymology. The specific name is a noun in the genitive case and it is a patronym for Gustavo Edmundo Orc��s Villag��mez (1903���1999), a pioneer of Ecuadorian vertebrate zoological research (Albuja V. 2004; Adler 2007). We recognize his contributions as a mentor and professor of a generation of zoologists in Ecuador, among them, two of us (LAC and AA). The specific name also is in recognition to the Fundaci��n Herpetol��gica Gustavo Orc��s that through the Vivarium, initially leaded by Jean-Marc Touzet and Ana Mar��a Velasco (the latter a collector of this species), pioneered herpetological public exhibits in Ecuador since the 1980 s., Published as part of Coloma, Luis A., Duellman, William E., C, Ana Almend��riz, Ron, Santiago R., Ter��n-Valdez, Andrea & Guayasamin, Juan M., 2010, Five new (extinct?) species of Atelopus (Anura: Bufonidae) from Andean Colombia, Ecuador, and Peru, pp. 1-54 in Zootaxa 2574 on pages 16-20, DOI: 10.5281/zenodo.197448, {"references":["Rueda-Almonacid, J. V. (2005 a) Atelopus sp. 16. 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, 152 pp.","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 / GERF-BIRF y Ecociencia, Quito, 79 - 108 pp.","Hijmans, R. J., Cameron, S. E., Parra, J. L., Jones, P. G. & Jarvis, A. (2005) Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology, 25, 1965 - 1978.","Ron, S. R. & Merino, A. (2000) Amphibian declines in Ecuador: overview and first report of chytridiomycosis from South America. Froglog, 42, 2 - 3.","Albuja, V. L. (2004) Dr. Gustavo Orces Villagomez (1903 - 1999). Politecnica, 25, 7 - 10.","Adler, K. (2007) Orces V., Gustavo (1903 - 1999). In: Adler, K. (Ed.) Contributions to the History of Herpetology. Society for the Study of Amphibians and Reptiles, Dexter, USA, 394 pp."]}

Atelopus gigas sp. nov. Holotype. KU 140320, adult female, from La Victoria, Departamento Nariño, Colombia, obtained in May 1970 by Kjell von Schneidern. According to John D. Lynch (in litt.) this locality is in Municipio de Ipiales, 38 km (by road) SE of Ipiales (0˚ 40 ' N, 77 ˚ 32 ' W; ~ 2700 m). Paratypes. KU 140315 –17, 140318 (cleared-and-stained adult female), 140319, 140321 – 29. Same data as holotype. Diagnosis. (1) A large species with SVL in adult males 43.0– 48.6 (mean = 45.6 mm, SD = 1.8082, n = 6), and in adult females 43.3–57.2 (mean = 53.8 mm, SD = 4.3232, n = 9); (2) hind limbs short, tibia length/SVL 0.346–0.413 (n = 15); (3) phalangeal formula of hand 2 - 2-3 - 3, basal webbing absent; (4) foot webbing formula I (0–½)-—(½– 1) II (½– 1 +)—(1 ½– 2 +) III (½– 1 +)—(2–3) IV (1 ½– 3)—(½– 1 ½) V; (5) snout rounded to acuminate in dorsal view, slightly protruding beyond lower jaw; (6) tympanic membrane, tympanic annulus, and stapes absent; (7) dorsal surfaces of body usually smooth, bearing few warts mostly in males; (8) warts present on forearm, flanks and dorsal surfaces of thighs and shanks; spiculae present on flanks of females; (9) vertebral neural processes inconspicuous; (10) dorsum yellowish cream to yellowish tan in preservative; (11) minute gray stippling absent on dorsum of body; (12) venter cream in preservative; (13) gular region without warts, spiculae or coni. In having a plain yellowish cream to yellowish tan dorsum in preservative, Atelopus gigas sp. nov. is most similar to A. guanujo, A. bomolochos (populations from Cutchil, Provincia Azuay, Ecuador), A. onorei Coloma, Lötters, Duellman, and Miranda-Leiva, A. ardila sp. nov. (some individuals), A. carbonerensis Rivero, A. chrysocorallus La Marca, and A. sorianoi from the Andes of Venezuela. Atelopus gigas (mean SVL of females 53.8; mean SVL of males 45.6) is significantly larger than A. guanujo (mean SVL of females 39.6; mean SVL of males 33.9). It differs from A. bomolochos and A. onorei in having coni and larger spiculae, and from A. bomolochos by lacking minute gray stippling (present in A. bomolochos). Atelopus gigas differs from A. ardila in females lacking a patch of spiculae and coni on the gular and pectoral regions (present in A. ardila), in having a nearly smooth dorsum (body smooth, bearing spiculae and coni mostly on sacral region in A. ardila), and by being significantly larger than A. ardila (mean SVL of females 46.7; mean SVL of males 38.6) (SVL of females Student’s t -test, t = 6.090, df = 45, P t -test, t = 10.932, df = 67, P Atelopus gigas differs from A. carbonerensis and A. sorianoi in having a less protuberant snout and lacking a prominent postocular crest (postocular crest present in A. carbonerensis and A. sorianoi). Furthermore, it differs from A. carbonerensis and A. chrysocorallus by having vocal slits in males (absent in A. carbonerensis and A. chrysocorallus) and lacking a row of warts on the dorsolateral surfaces of the body in males (present in A. carbonerensis and A. chrysocorallus). Description of holotype. (Figs. 6 A–C). Head slightly longer than wide, HLSQ and HDWD less than one third SVL (HLSQ /SVL = 0.277, HDWD /SVL = 0.266); snout rounded in dorsal view; in lateral view, profile of tip of snout to the anterior margin of jaw nearly straight and slightly protuberant; swollen gland on tip of snout absent; nostrils slightly protuberant, directed laterally, situated at level slightly posterior to apex of lower jaw; canthus rostralis distinct, nearly straight from eye to nostril; loreal region concave; lips not flared; interorbital region and occiput flat, smooth; eyelid flared without distinct tubercles; postorbital crest slightly raised, glandular; pretympanic and tympanic areas free of warts; tympanic membrane and tympanic annulus absent; postmandibular and temporal area with warts; choanae small, rounded, widely separated; tongue twice as long as wide, broadest at anterior half, free along its posterior half. Forearm relatively short (RDUL /SVL = 0.300); palmar tubercle round, poorly defined; supernumerary palmar tubercles distinct; thenar and subarticular tubercles low raised; digital tips with round pads; thumb relatively long (THBL / HAND = 0.779), apparently having two phalanges; webbing on hands absent, fingers having lateral keels that are more conspicuous on Finger IV; relative length of fingers III( 0+)—(½) II( ½)—(1 +) III( 1)—(2 +) IV( 2 +)—(1) V; relative length of toes I Dorsal surfaces smooth with scattered flat warts in posterolateral sacral region and on dorsal and posterior surfaces of thighs, increasing in number toward the flanks; warts on flanks and anterior and proximal upper anterior surfaces of forelimbs bear spiculae; each wart may lack or bear a single or a group (up to four) of spiculae; throat, chest, belly, undersides of limbs smooth; underside of arm rugose; cloaca opens as an inconspicuous tube at upper level of thighs, directed posteriorly; rugose margin of cloacal opening. Color in preservative (70–75 % ethanol): dorsal surfaces of head and body pale yellowish cream, becoming pale yellowish tan posteriorly on body, flanks, and dorsal surfaces of limbs; margin of cloacal opening dark brown; minute gray stippling absent on dorsum of body (viewed at 8 x magnification); warts and spiculae cream; dorsal surfaces of fingers vary from yellowish cream of inner finger and toe to yellowish tan towards outer fingers and toes; they bear a brown lines present at articular regions of phalanges, with the distal line being better defined than the rest; throat, chest, belly, and ventral surfaces of limbs uniform cream; outer metatarsal tubercle slightly lighter than adjacent areas; proximal end of tongue lacking black pigmentation, but lower lip bearing a poorly pigmented brown margin at distal end. Measurements of holotype (mm). SVL 55.9, TIBL 20.9, FOOT 21.1, HLSQ 15.5, HDWD 14.9, ITNR 5.3, EYDM 5.8, EYNO 3.9, RDUL 16.8, HAND 13.6, THBL 10.6, SW 16.5. Variation. Meristic variation is shown in Table 3. The paratypes resemble the holotype with the following noteworthy exceptions. There is variation in size, abundance and distribution of warts and spiculae. For example, a female (KU 140316) has more abundant warts and larger spiculae along dorsum and flanks than other females, whereas female KU 140317 has fewer warts and spiculae. Another female (KU 140317) has a more rugose plantar surface than other females. A juvenile female (KU 140321, SVL = 43.3) has more warts and spiculae than other specimens. A female (KU 140317) has a shorter than normal and deformed Finger IV. Six males differ among them in that KU 140319, and 140324 possess more abundant warts on dorsum of sacral region, thighs and shanks. Atelopus gigas La Victoria Sexual dimorphism is evident in that females are larger than males (Table 3). Males have vocal slits and keratinized nuptial pads on the dorsal and inner surfaces of the thumb and of Finger III, in which they are more abundant at the proximal phalanx. The forelimbs are relatively long and slender only in females, but they are short with a stout muscular area in males (Table 3); there are significant differences in RDUL between males and females (RDUL Student’s t -test, t = 4.4105, df = 13, P All paratypes lack a brown area surrounding cloacal opening. In preservative, the dorsal coloration varies from uniform yellowish cream (KU 140321) to dark brown (KU 140319). One individual (KU 140318) has two dark brown bars across mid-dorsum of the body and a few, small diffuse dark brown marks on the dorsal surfaces of the hindlimbs. In one female (KU 140317), the lower lip is more pigmented with brown. Color in life: Unknown. Presumably yellow or orange by comparison with coloration in ethanol of other preserved Atelopus. Tadpoles. Tadpoles of this species are unknown. Distribution, natural history, and conservation status. Atelopus gigas is known only from the type locality (Fig. 5). This locality currently has cultivated areas. At La Victoria, annual mean rainfall is 1190 mm and the annual mean temperature is 13.3 ºC (Hijmans et al. 2005). At time of collection, in May 1970, four out of seven adult females were gravid. One of them (KU 140318) contained 626 ovarian eggs of about 2.15 mm of diameter (n = 15, SD = 0.151). The stomach of KU 140318 contained remains of 63 arthropods [22 Coleoptera (1 Brenthidae, 7 Carabidae, 3 Curculionidae, 4 Staphylinidae, 7 undetermined), 13 Acari, 7 Hymenoptera, 4 Diptera, 1 Aracnida, 1 Blatodea, 1 Hemiptera, 1 Homoptera, 7 undetermined insects, 5 insect larvae], vegetal material, unidentified helminth endoparasites, and some undetermined material. The largest prey was an insect larva 7.1 mm long. Atelopus gigas is considered as Data Deficient under IUCN Red List categories and criteria, because no collecting efforts have been carried out at its only known locality. Nonetheless, a similar fate of either severe decline or extinction (as most highland species of Atelopus) is a reasonable presumption. The chytrid fungus has been reported in very close proximity to the type locality in an Ecuadorian locality at Carchi Province (Ron and Merino 2000).The risk factor of potential threat caused by the chytrid for anuran amphibian species calculated by Rödder et al. (2009: Fig. 2 C) is high at the area of its distribution. Etymology. Gigas is a noun in apposition from the Greek word “ gigas ” that means giant. It is used in reference to the large size of this species, which is exceeded in size only by Atelopus boulengeri.

Atelopus ardila sp. nov. Holotype. KU 200214, adult gravid female, from 7.5 km E of Pasto, on road to Laguna La Cocha (= Lago Guamuez) (ca. 1 &ring; 12 ' N, 77 &ring; 13 ' W; 2800 m), Vereda San Fernando, Departamento Nari��o, Colombia, obtained on 25 February 1984 by Patricia A. Burrowes and Benjam��n del Castillo. Paratypes. All (except ICN 3406 and specimens from Hacienda San Gerardo at Volc��n Galeras) are from the road from Pasto to Laguna de la Cocha, Vereda San Fernando, Departamento Nari��o, Colombia; KU 200207 ���12, 200215 ��� 18 (3 females, 7 males), with same data as holotype; KU 200213, male, from 7.5 km E of Pasto (2800 m) obtained on 25 February 1984 by Patricia A. Burrowes and B. del Castillo; KU 169267, female, from 8 km NE Pasto (3020 m), obtained on 24 September 1974 by William E. Duellman; KU 169268, female, from 8 km NE of Pasto (3050 m), obtained on 21 May 1975 by William E. Duellman; KU 169269 ��� 335, 169341, QCAZ 24467 ��� 71 (10 females, 61 males, 1 juvenile), from 12 km E of Pasto (3050 m), obtained on 24 September 1974 by William E. Duellman, Linda Trueb, Dana T. Duellman, and John E. Simmons, and on 2 October 1974 by William E. Duellman and John E. Simmons; KU 200239, female, from 12.8 km NE of Pasto (3020 m), obtained on 25 February 1984 by Benjam��n del Castillo; KU 154583, adult gravid female, from crest between Pasto and Laguna La Cocha, on road to Laguna La Cocha (3150 m), obtained on 2 August 1965 by R. E. Smalley; KU 200219 ��� 38 (19 females, 1 male), from El Tabano, 16 km E of Pasto (3280 m), obtained on 25 February 1984 by Patricia A. Burrowes and Benjam��n del Castillo; KU 169262 ���66, 5 females, from north shore of Laguna La Cocha, (2790 m), obtained on 2 October 1974 by John E. Simmons and Linda Trueb; ICN 3406, female, from Volc��n Galeras (Hacienda San Gerardo), obtained on August 1977 by Santiago D��az; ICN 3633, 3636���38, 4 females, from El Encano (2800 m), obtained on December 1977 by L. G��mez; ICN 450, 459, 469, 471 ��� 73, 481 (3 females, 4 males), from El Encano (2800 m), obtained on 19 January 1971 by Nancy Bastidas. Diagnosis. (1) A species with mean SVL in adult females 46.7 mm (40.3���52.2, SD = 2.852, n = 48) and in adult males 38.6 mm (34.7���41.8, SD = 1.428, n = 76); (2) hind limbs short, tibia length/SVL 0.302���0.406 (n = 121); (3) phalangeal formula of hand 2 - 2-3 - 3, webbing absent; (4) foot webbing formula I (0+- 1 -)���(��- 2 -) II (��- 1 +)���(1 - - 2 ��-) III (��- 2 -)���(1 - - 3) IV (1 - - 3)���(1 - - 2 -) V; (5) snout acuminate, barely protruding beyond lower jaw; (6) tympanic membrane and tympanic annulus absent; (7) dorsal surfaces of body smooth, bearing spiculae and coni mostly on sacral region, (8) black spiculae and coni (gray in preservative) present on flanks; (9) vertebral neural processes inconspicuous; (10) dorsum orange-red to black (pale creamy brown to black in preservative); (11) minute gray stippling present on dorsum of body; (12) venter orange to orange-red and dull yellow (cream in preservative); (13) gular region with a patch of black spiculae on gular-chest region in females (but see sexual dimorphism variation). By displaying in life either orange���red, or black or a combination of both colors, Atelopus ardila sp. nov. is most similar to A. guanujo Coloma (some individuals orange-red), A. bomolochos Peters (some individuals orange-red), A. sorianoi (orange-red), A. sp. 6 (some individuals orange, Rueda-Almonacid and Acosta 2005), A. ebenoides Rivero (black), A. nanay Coloma (black), A. marinkellei Cochran and Goin (black), A. guitarraensis Osorno-Mu��oz, Ardila-Robayo, and Ruiz-Carranza (some individuals black), A. pastuso sp. nov. (some individuals black), A. patazensis Venegas, Catenazzi, Siu-Ting, and Carrillo (orange and black), A. podocarpus sp. nov. (black), A. carrikeri Ruthven (black to orange and yellow), and A. ignescens (black). Nonetheless, by having a patch of black spiculae on females��� gular-chest region, A. ardila is distinct from all of them, except A. ignescens, A. carrikeri, and potentially A. sp. 6 (presence and characteristics of gular patch unknown). It differs from the latter by lacking a dark brown dorsal pattern. It differs from A. ignescens in SVL, foot and tibia length, and sacrum width (see below), and by nearly lacking posterolateral process on hyoid plate (well developed in A. ignescens). It differs from A. carrikeri in size and by having minute gray stippling on dorsum of body (absent in A. carrikeri); A. ardila (mean SVL of females 46.7; SD = 2.852, n = 48) is significantly smaller than A. carrikeri (mean SVL of females 57.0, SD = 4.153, n = 4; data from Coloma 1997) (SVL of females t -test, t = 6.7175, df = 50, P Atelopus ardila vs A. ignescens (see materials and methods). Three axes of the PCA accounted for 58 % of the total variation (Table 1). PC I mainly included tibia, foot, and radio ulna lengths, which are the highest loadings along this axis; in which most A. ardila individuals tended to have larger tibia, feet and forearms (Fig. 2). PC II mainly included sacrum width and internarial distance, whereas in PC III the highest loading was SVL. Although there is some overlap of the morphometric space of A. ardila vs A. ignescens (Fig. 3), significant separation occurred along two axes [overall: F(3) = 43.854, P Description of holotype. (Figs. 1 A���C, virtual animation). Head about as long as wide, HLSQ and HDWD less than one third SVL (HLSQ /SVL = 0.260, HDWD /SVL = 0.256); snout acuminate, its margin rounded in dorsal view; profile of tip of snout in lateral view curved and barely protruding 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, nearly straight from eye to nostril; loreal region concave; lips slightly flared; interorbital and occipital regions flat, smooth; eyelid flared without tubercles; postorbital crest flared, not glandular; postorbital, pretympanic and post-tympanic areas bearing coni; tympanic membrane and tympanic annulus absent; choanae small, rounded, widely separated (30.3 % of HW); tongue about twice as long as wide, its posterior half not attached to floor of mouth. Forearm relatively short (RDUL /SVL = 0.271); palmar tubercle round; thenar, supernumerary palmar, and subarticular tubercles distinct; digital tips with round pads; thumb relatively long (THBL / HAND = 0.611), apparently having two phalanges; webbing on hands absent, fingers lacking lateral fringes; relative length of fingers III( 0+)���(��) II (��)���(1 -) III (��)���(1 -) IV (0+�����)���(1) V; relative length of toes IMeasurements of holotype (mm). SVL 47.3, TIBL 15.3, FOOT 20.4, HLSQ 12.3, HDWD 12.1, ITNR 4.6, EYDM 3.6, EYNO 2.7, RDUL 12.8, HAND 12.6, THBL 7.7, SW 13.5. Variation. Meristic variation is given in Table 2. A juvenile female (KU 169341, SVL = 21.3 mm) possesses abundant spiculae and coni on the dorsum and flanks, but lacks spiculae on gular-chest region. It differs from adults by having cream flanks in contrast to gray dorsum. The paratypes resemble the holotype with the following noteworthy exceptions. Females are larger than males (Table 2). Males have vocal slits and keratinized nuptial pads variably extensive on the dorsal and inner surfaces of the thumb (= Finger II) and Finger III. The forelimbs are relatively long and slender only in females, but they are short with a stout muscular area in males (Table 2); there are significant differences in RDUL between males and females (RDUL Student���s t -test, t = 11.4579, df = 106, P Color variation in preservative (~ 70 % ethanol): Topotypic specimens and one from a nearby site (KU 169268) (4 females, 7 males) are similar to the holotype. Dorsal color generally varies from pale creamy brown (KU 200217) to dark brown (KU 200212); spiculae and coni vary from dark brown to black. A male (KU 200216) has dark brown marbling on the dorsum, and stripes at joints on the tarsi, feet and hands. The venter is uniform cream in all individuals. A thin brown line along middorsum varies from conspicuous (KU 200207) to barely visible (KU 200217). Atelopus ardila Pasto-Cocha Coloration in a series of specimens (KU 169269 ���335, 169341, 200239, QCAZ 24467 ��� 71) from 12.0��� 12.8 km east of Pasto (11 females, 61 males, 1 juvenile) varies from plain pale creamy brown (e.g., KU 169270) to plain black (e.g., KU 169322), or to a mosaic pattern of bold black marks (e.g., KU 169303). A narrow middorsal line varies from conspicuous (e.g., KU 169318) to absent (e.g., KU 169317) in males, whereas in females it is absent. The venter is uniform cream; however, in individuals that are darker dorsally the black color partially covers plantar and toe surfaces. In a series of specimens (KU 200219 ��� 38) from 16 km E of Pasto (19 females, 1 male) the dorsum varies from deep black (12 specimens, e.g., KU 200235) to dark brown (e.g., KU 200232) to a mosaic pattern of black mottling (e.g., KU 200230). In 12 specimens the flanks are paler than the dorsum. In black individuals, the black invades the pelvic patch and the palmar and plantar surfaces, nearly entirely covering the latter, except for some of the tubercles (e.g., KU 200221). In five females (KU 169262 ��� 66) from Laguna La Cocha the dorsum is mostly entirely black with diffuse dark brown mottling. Color in life. (Figs. 3, 4 A���B). The following is based on color slides taken by WED (CT 5932 ���34, 5735��� 42, 7211��� 12 of KU 169269 ���73, 169285���88, 200207 ���08, 200217), from WED field notes of 24 September and 2 October 1974, 21 May 1975, and from Patricia A. Burrowes��� field notes of 25 February 1984. The dorsal ground color of the body and limbs varies from nearly entirely yellow and red-orange to black. Single individuals or series were described as follows: KU 200207 ��� 18 (from 7.5 km E Pasto, 2800 m): red with or without black suffusion on snout and limbs; belly orange-red; flanks yellow with black lateral tubercles. KU 200239 (from 8 km NE Pasto, 3020 m): body yellow with black markings. KU 169267 (from 8 km NE Pasto, 3020 m): dorsum orange-red with black-tipped tubercles, webbing pale orange, venter deep orange, iris dull brown with minute gold flecks. KU 169268 (from 8 km NE Pasto, 3020 m): dorsum dull orange-red with black-tipped tubercles and brownish black head, venter orange. KU 169269 ��� 341 (from 12 km E Pasto, 3050 m): dorsal ground color varying from bright yellow-ochre to medium cadmium-orange and bright venetian red; dorsum plain with slight brownish suffusion and thin vertebral stripe and/or darker colored tubercles to spotted, mottled, or striped with sepia-umber (warm); venters plain with ground color; iris dull brown with minute bronze flecks. KU 200219 ��� 38 (from 16 km E Pasto, 3280 m): black dorsally; venter orange to dull yellow. KU 169262 ��� 66 (from North shore Lago de la Cocha, 2790 m): dorsum black; venter red in KU 169262. Tadpoles. G��mez Castillo (1982) described tadpoles from the R��o Pasto and tributaries (2600���3000m). Tadpoles in Gosner Stages 18���29 had total lengths of 12���21 mm. G��mez Castillo (1993) described and illustrated black tadpoles from several creeks around Lago La Cocha (= Lago Guamez). He remarked on their similarity to tadpoles of A. varius and A. ignescens described by Starret (1967) and Duellman and Lynch (1969) respectively. Distribution, ecology, natural history, and conservation status. Atelopus ardila is known only from the type locality and surrounding areas between Volc��n Galeras eastward to Laguna de la Cocha in the Pasto Massif of the Andean Cordillera in Departamento Nari��o in southern Colombia. These localities are in subparamo and paramo 2800���3280 m above sea level. The area of its extent of known occurrence is about 51.3 km 2 (Fig. 5). At the type locality, annual mean rainfall is 1388 mm and the annual mean temperature is 12 ��C (Hijmans et al. 2005). At the time of collection, individuals (KU 200207 ��� 18) were crawling along a grassy irrigation ditch; others (KU 200219 ��� 38) were crawling on black rocks in a subparamo area with a steep incline, orange soils, black loose rocks, bunch grasses, Baccharis, and lots of mosses; an amplectant pair (yellow male, red female) also was found (P.A. Burrowes field notes, 25 February 1984). KU 169268 was under a yucca stump by day in subparamo with dense woody bushes and bamboo (WED field notes, 21 May 1975). KU 169285 ��� 341 were crawling on the ground (open, mossy, grassy) after a rain; some were under rocks by day. Others (KU 169262 ��� 66) were crawling across a road by day (WED field notes, 2 October 1974). A female (KU 169267) was under a rock by day (temperature: 7.5 ���13.0&ring;C; no rainfall) in a subparamo area with dense woody bushes and bamboo. Other individuals (KU 169269 ��� 84) were in a rocky stream, where most were under rocks at the edge of the streambed; a few individuals were crawling amidst rocks on the ground by day (WED field notes, 24 September 1974). Del Castillo Ch. (1982) and G��mez Castillo (1982; 1993) provided data on the reproductive biology of individuals from several localities and at ex situ conditions. Del Castillo Ch. (1982) induced oviposition using hormones, described behavior, physiology and early development of individuals and eggs from El Encano (locality stated in G��mez Castillo 1993: 169). G��mez Castillo (1982) also described courtship, mating and reproduction of individuals from El Encano, San Fernando, and 11 km N of Pasto. G��mez Castillo (1993) provided data on the reproductive biology of individuals from Laguna La Cocha and vicinity and reported additional descriptions of induced oviposition. Three types of calls of individuals from several populations from Departamento Nari��o (presumably a mixture of A. ardila and A. pastuso) were briefly described by G��mez Castillo (1982: 61), two of them as peeps and another as a trill. Before axillary amplexus, it was observed that the male walks or jumps over the female from behind or from the side. Two males embracing a female were observed after exposing several males to a female. Amplectant pairs were found throughout the year at El Encano, San Fernando, Veredas Motil��n, and Santa Rosa (G��mez Castillo 1982; 1993). At El Encano a pair remained in amplexus for at least 22 days. Amplexus among pairs maintained in captivity lasted more than 2��� 3 months; in one of these cases a pair was in amplexus for more than 74 days, and the male remained in the amplectant position for two days after the female died. Amplectant pairs were observed on two occasions under water (about 25 cm below the surface)���one in October 1978 at El Encano, and another in May 1979 at San Fernando. Another amplectant pair was observed under water at Quebrada Funduyacu (G��mez Castillo 1993). Tadpoles have been found throughout the year, but they are less abundant in the summer from June to August. Tadpoles were found in clear, non-contaminated waters in mountain creeks, Laguna La Cocha, and the R��o Pasto and tributaries, but they were not observed near human habitations in urban areas (G��mez Castillo 1982; 1993). G��mez C. and Ramos O. (1982) and G��mez and Ramos O. (1982) analyzed the stomach content of 30 individuals of five populations (San Fernando, 2600 m; El Encano, 2850 m; Daza, 2800 m, Volc��n Galeras, 3150 m; Col��n-Mocoa, 2200 m) from Municipio de Pasto and Intendencia de Putumayo, presumably belonging to Atelopus ardila (under the name A. ignescens). No voucher specimens were indicated, but we assume that the specimens were A. ardila based on the localities, measurements and color description of the animals provided. They report the following diet: Diptera 38 %, Coleoptera 33.2 %, Hymenoptera 25.4 %, Lepidoptera and Homoptera in smaller proportions. Chrysomelidae (Coleoptera) were the most abundant prey (26.5 %), followed by Drosophilidae (Diptera), Pteromalidae (Hymenoptera), Braconidae (Hymenoptera), Anthomyiidae (Diptera), Muscidae (Diptera), and Tephritidae (Diptera). Atelopus ardila is considered to be Critically Endangered (Possibly Extinct) (A 2 ace, IUCN Red List categories and criteria). The species is tagged as Possibly Extinct until further surveys confirm otherwise. The population has declined dramatically (more than 80 %) in the last two decades probably because of climate change and the impact of pathogens, which have affected many other montane species of Atelopus. No chytrid fungus reports or climate data for its area of distribution are available. The closest report of occurrence of the chytrid fungus is from Provincia Carchi near the Ecuador-Colombia border (Ron and Merino 2000). The risk factor of potential threat caused by the chytrid for anuran amphibian species calculated by R��dder et al. (2009: Fig. 2 C) is high at the area of its distribution. Although this species previously was abun, Published as part of Coloma, Luis A., Duellman, William E., C, Ana Almend��riz, Ron, Santiago R., Ter��n-Valdez, Andrea & Guayasamin, Juan M., 2010, Five new (extinct?) species of Atelopus (Anura: Bufonidae) from Andean Colombia, Ecuador, and Peru, pp. 1-54 in Zootaxa 2574 on pages 3-12, DOI: 10.5281/zenodo.197448, {"references":["Rueda-Almonacid, J. V. & Acosta, A. R. (2005) Atelopus sp. 6. In: Rueda-Almonacid, J. V., Rodriguez-Mahecha, J. V., Lotters, J. V., La Marca, E., Kahn, T. & Angulo, A. (Eds.) Ranas arlequines. Conservacion Internacional. Panamericana Formas e Impresos S. A., Bogota, Colombia, 142 pp.","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.","Gomez Castillo, L. (1982) Aspectos ecologicos, morfologicos y de conducta de las ranas del genero Atelopus del Departamento de Narino, Colombia. Revista Humboldt, 1, 53 - 64.","Gomez Castillo, L. (1993) Aspectos de la ecologia y ciclo vital de la rana Atelopus ignescens (Cornalia) de los alrededores del Lago Guamuez, Narino Colombia. Revista de Investigaciones Universidad de Narino, Pasto, 7 (VI), 168 - 177.","Starrett, P. H. (1967) Observations on the life history of frogs of the family Atelopodidae. Herpetologica, 3, 195 - 204.","Duellman, W. E. & Lynch, J. D. (1969) Descriptions of Atelopus tadpoles and their relevance to atelopodid classification. Herpetologica, 25, 231 - 240.","Hijmans, R. J., Cameron, S. E., Parra, J. L., Jones, P. G. & Jarvis, A. (2005) Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology, 25, 1965 - 1978.","Del Castillo Ch., B. (1982) Comportamiento reproductor y primeros estadios embrionarios de Atelopus ignescens (Cornalia). In: II Congreso Grancolombiano y VI Jornadas Nacionales de Ciencias Biologicas. Vol. I. Sociedad Ecuatoriana de Biologia (ed.). Editorial Escuela Superior Politecnica del Litoral, Guayaquil, Ecuador. 127 - 132 pp.","Gomez, C. L. & Ramos, O. A. (1982) Alimento de las ranas Atelopus ignescens (Cornalia) de Narino y Putumayo (Colombia). In: II Congreso Grancolombiano y VI Jornadas Nacionales de Ciencias Biologicas. Vol. I. Sociedad Ecuatoriana de Biologia (ed.). Editorial Escuela Superior Politecnica del Litoral, Guayaquil, Ecuador. 167 - 173 pp.","Ron, S. R. & Merino, A. (2000) Amphibian declines in Ecuador: overview and first report of chytridiomycosis from South America. Froglog, 42, 2 - 3.","Ardila-Robayo, M. C. & Maldonado-Silva, R. A. (2004) Sapito arlequin de Narino << Atelopus ignescens >>. In: Rueda- Almonacid, J. V., Lynch, J. D., & Amezquita, A. (Eds.) Libro rojo de los anfibios de Colombia. Serie Libros Rojos de Especies Amenazadas de Colombia. Conservacion Internacional Colombia, Instituto de Ciencias Naturales-Universidad Nacional de Colombia, Ministerio de Medio Ambiente, Panamericana Formas e Impresos S. A., Bogota, Colombia, 384 pp.","Cepeda-Quilindo, B. & Rueda-Almonacid, J. V. (2005) Atelopus complejo ignescens. 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, 135 pp.","Mueses-Cisneros, J. J. (2005) Fauna anfibia del valle de Sibundoy, Putumayo-Colombia. Caldasia, 27, 229 - 242.","Gray, P. (1983) Morphometrics of the Atelopus ignescens Complex (Anura: Bufonidae). M. Sc. dissertation, Lawrence, University of Kansas. 225 pp.","Coloma, L. A., Lotters, S. & Salas, A. W. (2000) Taxonomy of the Atelopus ignescens complex (Anura: Bufonidae): Designation of a neotype of Atelopus ignescens and recognition of Atelopus exiguus. Herpetologica, 56, 303 - 324.","Ron, S. R., Duellman, W. E., Coloma, L. A. & Bustamante, M. R. (2003) Population decline of the Jambato toad Atelopus ignescens (Anura: Bufonidae) in the Andes of Ecuador. Journal of Herpetology, 37, 116 - 126.","Lotters, S. (1996) The Neotropical Toad Genus Atelopus. Checklist - Biology - Distribution. M. Vences & F. Glaw, Koln, Germany, 143 pp."]}

Atelopus angelito: New country record for Ecuador Atelopus angelito Ardila-Robayo and Ruiz-Carranza was named and described from the subparamo of the P��ramo de las Papas at Cordillera Central of the Andes in Departamento del Cauca, Colombia, at Municipio de San Sebasti��n, near Valencia, ca. 1 �� 51 ' N, 76 �� 47 ' W, 2900���3000 m. Their description was based on five adult males, one female and three juveniles. Additionally, Ardila-Robayo (2005) provided data on conservation. Two other females were recorded (under the name Atelopus sp. 14) from R��o La Plata, Comunidad de Mor��n, Reserva Ecol��gica El ��ngel, Provincia Carchi, Ecuador, by Y��nez-Mu��oz and Altamirano B. (2005). Herein, these specimens are also referred to Atelopus angelito. They provided SVL���s for two females, a description of color in life, two photographs (top one, KU 178417), and conservation data. Herein we report additional data of the latter individual KU 178417 (Figs. 4 H, 14), an adult gravid female that we consider to be conspecific with A. angelito. It was found under a stone 14 km (airline) SE Maldonado (ca. 0�� 49 ' N, 78 �� 3 ' W, 2500 m), Provincia Carchi, Ecuador, by John D. Lynch, on 31 May 1977. The Ecuadorian localities extend the geographic range of A. angelito 183 km (airline) southwest of its type locality. The color in life of KU 178417 was described by Lynch (field notes, 31 May 1977) as follows: pea green above with black markings; flanks yellow with black, venter orange with black; iris dark brown with yellow edging on pupil. Its measurements and ratios (in mm) are: SVL 49.2, TIBL 17.1, FOOT 19.4, HLSQ 14.1, HDWD 14.3, ITNR 4.6, EYDM 4.5, EYNO 3.5, RDUL 14.0, HAND 11.2, THBL 8.4, SW 16.5, HDWD /SVL 29.1 %, HLSQ /SVL 28.7 %, HLSQ / HDWD 98.6 %, EYNO / EYDM 77.7 %, TIBL /SVL 34.8 %, FOOT /SVL 39.4 %. A comparison of the female (KU 178417) with specimens of the original description of Atelopus angelito does not reveal any major differences, but there are subtle differences in size, amount of foot webbing, and coloration. The only adult female available from the type locality is smaller (SVL = 41.0 mm) than the Ecuadorian female (SVL = 49.2 mm). Nonetheless, the two additional adult females (deposited at Museo Ecuatoriano de Ciencias Naturales, MECN) recorded from Ecuador by Yanez-Mu��oz and Altamirano B. (2005) have SVLs of 41.3 and 43.8 mm. Thus, adult female size of the topotypic female is very close to the range of variation of Ecuadorian females. The extent of webbing described in the diagnosis and depicted (of a male) in Fig. 3 by Ardila-Robayo and Ruiz-Carranza (1998) is greater than that in the Ecuadorian female. However, the amount of webbing in several specimens (including a female) from the type locality depicted in Fig. 4 by Ardila-Robayo and Ruiz-Carranza (1998) is similar to that in the Ecuadorian female. In the types the flanks are black and creamy white, whereas the flanks in Ecuadorian females are black and yellow. These color differences can be attributed to either intraspecific or geographic inter-population variation; however, sample sizes are too small to know with certainty. Distribution, ecology, and conservation status. Atelopus angelito is known only from paramo and subparamo habitats at elevations of 2500���3000 m on the eastern slopes of Cordillera Central in Departamento del Cauca in southern Colombia to the western slopes of Cordillera Occidental in Provincia Carchi in northern Ecuador. The area of its extent of occurrence is of about 533 km 2. In Ecuador, it occurs in the Bosque de Neblina Montano Natural Formation (according to the classification proposed by Valencia et al. 1999). These localities lie in the high massif of the Nudo de Pasto, as defined by Duellman (1979). At the Ecuadorian locality (14 km SE Maldonado) annual mean rainfall is 1051 mm and the annual mean temperature is 14.2 ��C (Hijmans et al. 2005). The Ecuadorian localities of Atelopus angelito are in close proximity to one locality of A. pastuso at the western versant of Cordillera Occidental (Fig. 5); thus, they might have occurred in parapatry or sympatry at these localities. Atelopus angelito is considered to be Critically Endangered (Possibly Extinct) (A 2 ace, IUCN Red List categories and criteria). The species is tagged as Possibly Extinct until further surveys confirm otherwise. The population has declined dramatically (more than 80 %; see below) in the last two decades probably because of climate change and the impact of pathogens, that have affected many other montane species of Atelopus. The last individual of this species was collected 12 April 1995 in Colombia (Ardila-Robayo 2005) and there have been no additional efforts to find it. In Ecuador, the last two living individuals were collected on 22 July 1988, and no additional records exist since that date, in spite of seven occasional searching efforts by QCAZ parties (10 June 1989, 2��� 3 March 1993, 10 May 1996, 23 October 1999, 17 September 2008, 24 September 2008, 28 September 2009) and intensive searching efforts in 2003 and 2004 by MECN parties at seven nearby sites in Provincia Carchi (Y��nez-Mu��oz and Altamirano 2005). Presence of the amphibian chytrid fungus was reported in an Ecuadorian locality near the localities for Atelopus angelito and for A. pastuso from 42 km W of Tulc��n, Provincia Carchi, in 1993 (Merino-Viteri 2001)., Published as part of Coloma, Luis A., Duellman, William E., C, Ana Almend��riz, Ron, Santiago R., Ter��n-Valdez, Andrea & Guayasamin, Juan M., 2010, Five new (extinct?) species of Atelopus (Anura: Bufonidae) from Andean Colombia, Ecuador, and Peru, pp. 1-54 in Zootaxa 2574 on pages 34-35, DOI: 10.5281/zenodo.197448, {"references":["Ardila-Robayo, M. C. (2005) Atelopus angelito. 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, 56 pp.","Yanez-Munoz, M. & Altamirano B., M. (2005) Atelopus sp. 14. 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, 150 pp.","Ardila-Robayo, M. C. & Ruiz-Carranza, P. M. (1998) Una nueva especie de Atelopus A. M. C. Dumeril & Bribon 1841 (Amphibia: Bufonidae) de la Cordillera Central Colombiana. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 83, 281 - 285.","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 / GERF-BIRF y Ecociencia, Quito, 79 - 108 pp.","Duellman, W. E. (1979) The herpetofauna of the Andes: Patterns of distribution, origins, differentiation, and present communities In: Duellman, W. E. (Ed.) The South American herpetofauna: Its origin, evolution, and dispersal. Monographs of the Museum of Natural History, The University of Kansas, Lawrence, 371 - 459 pp.","Hijmans, R. J., Cameron, S. E., Parra, J. L., Jones, P. G. & Jarvis, A. (2005) Very high resolution interpolated climate surfaces for global land areas. International Journal of Climatology, 25, 1965 - 1978.","Merino-Viteri, A. (2001) Analisis de posibles causas de las disminuciones de poblaciones de anfibios en los Andes del Ecuador. Tesis de Licenciatura, Quito, Ecuador, Pontificia Universidad Catolica del Ecuador. 66 pp."]}

Pristimantis minimus Ter��n-Valdez & Guayasamin, new species Holotype. QCAZ 41612, an adult male (Fig. 1, 2 B) from the Cordillera del C��ndor, Miazi Alto (04.25656 S, 78.622456 W; 1300���1315 m.a.s.l.), Cant��n El Pangui, Provincia Zamora Chinchipe, Ecuador (Fig. 3), collected by Juan M. Guayasamin, Elicio E. Tapia, and Holger Braun on 10 April 2009. Paratypes. All paratypes (45 males, 25 females) were collected in two areas of the Cordillera del C��ndor, east and west of R��o Nangaritza, within the Cant��n Nangaritza, Provincia Zamora Chinchipe, Ecuador (Fig. 3); airline distance between the two localities is 6 km. Specimens collected east of R��o Nangaritza are: 48 adult males (QCAZ 40744 ��� 45, 41033, 41380, 41422, 41429, 41444, 41496, 41498, 41500, 41511, 41537, 41549, 41551, 41553, 41606, 41614, 41616 ��� 17, 41619���20, 41623, 41630 ��� 31, 41633, 41635, 41643, 41650, 41660 ��� 61, 41664���66, 41668, 41672, 41690 ��� 91, 41693���95, 41701, 41711, 41713, 41716, 41721 ��� 22, 41738), and 25 adult females (QCAZ 41378, 41426, 41430 ��� 33, 41443, 41497, 41499, 41548, 41600, 41602, 41607, 41613, 41621, 41634, 41651, 41667, 41670 ��� 71, 41673, 41689, 41714, 41718, 41723) from the type locality, Miazi Alto (04.25656 S, 78.622456 W; 1300���1315 m.a.s.l.), collected by Juan M. Guayasamin, Holger Braun, Elicio E. Tapia on 06��� 12 April 2009; QCAZ 41495, cleared-and-stained (04.24584�� S, 78.61497�� W; 1415 m.a.s.l.); QCAZ 41537, 41548 ��� 49, 41551, 41553 (04.24663�� S, 78.62119�� W; 1432 m.a.s.l.); QCAZ 41643, 41650 ��� 51 (04.25026�� S, 78.61746�� W; 1256 m.a.s.l.); QCAZ 41511, 41600, 41602, 41606 ���07, 41660 ��� 61, 41664���68, 41670 ��� 73 (04.25628�� S, 78.62225�� W; 1260���1300 m.a.s.l.); QCAZ 41613, 41614, 41616 ��� 21, 41623, 41630 ��� 31, 41633���35 (04.25656�� S, 78.62246�� W; 1314 m.a.s.l.). The following specimens were collected on mountains west of R��o Nangaritza by Elicio E. Tapia, Jessica L. Deichmann, Sylvia Ald��s on 17��� 18 April 2009: QCAZ 41378, 41380, 41426, 41430 ��� 31, 41443 (04.25523�� S, 78.67314�� W; 1300 m.a.s.l.); QCAZ 40744 ��� 45, 41433, 41689 ��� 91, 41693���95, 41721 ��� 23, 41738 (04.25789�� S, 78.68166�� W; 1554 m.a.s.l.); QCAZ 41701 (04.25669�� S, 78.67798�� W; 1418 m.a.s.l.); QCAZ 41711, 41713 ��� 14, 41716, 41718 (04.25971�� S, 78.68630�� W; 1685 m.a.s.l.). Generic placement. Some of the main traits shared by all Pristimantis include: (1) head as wide as body; (2) cranial crests absent; (3) dentigerous process of vomers present; (4) terminal discs on digits expanded, with circumferential grooves and terminal phalanges T-shaped; (5) Toe V larger than Toe III (Hedges et al. 2008). All traits observed in the new species agree with the diagnosis of Pristimantis. However, no synapomorphy has been yet identified for the genus Pristimantis and, indeed, three other genera (Eleutherodactylus, Ischnocnema, Craugastor) share the same diagnostic characters. In the genus Craugastor (family Craugastoridae), species have a sexually dimorphic tympanum (larger in males), although sexual dimorphism for the tympanum has not been determined in the subgenus Hylactophryne (Hedges et al. 2008); most species in Pristimantis do not present a sexually dimorphic tympanum (Hedges et al. 2008). Given that many clades have restricted geographic ranges, distribution provides an indirect clue to determine the generic placement of newly discovered taxa (see Guayasamin et al. 2009). For example, Craugastor is distributed mostly in Central America, marginally reaching northwestern South America (northeastern Honduras southeastward through Nicaragua, Costa Rica, and Panama to northwestern Ecuador; modified from Hedges et al. 2008); Eleutherodactylus (family Eleutherodactylidae) is distributed throughout the West Indies, peninsular Florida, southern Texas, northern Central America (Frost 2009); Ischnocnema (family Brachycephalidae) is restricted to the Atlantic Coastal Forest in eastern Brazil and in extreme southeastern Brazil and northern Argentina (Hedges et al. 2008); whereas, Pristimantis is found in southern Central America, most of South America, and on islands in the Lesser Antilles closest to the mainland of South America (Trinidad and Tobago, Grenada, St. Vincent; Hedges et al. 2008). Therefore, the newly discovered Pristimantis, based on geography, is likely to be part of Pristimantis, an hypothesis that should be tested with molecular data. Diagnosis. A species of Pristimantis having the following characters: (1) skin on dorsum smooth to slightly shagreen, lacking tubercles or folds; venter shagreen, lacking discoidal or thoracic folds (Figs. 1, 2); (2) tympanic membrane absent, tympanic annulus present but not visible externally; (3) snout short, rounded in dorsal and lateral views; (4) upper eyelid lacking enlarged tubercles, when present tubercles are low and almost undistinguishable from surrounding skin texture; upper eyelid slightly smaller or as wide as interorbital distance; cranial crests absent; (5) dentigerous process of vomers well developed, the number of teeth in each process varies from 1 to 4; (6) males with a small subgular vocal sac, vocal slits present, nuptial pads absent; (7) Finger I shorter than Finger II; discs on fingers expanded and rounded, except for Finger I that is only slightly expanded, disc on Finger III acute (Fig. 4 A, C, Fig 5 A, B); (8) fingers lacking lateral fringes; outer palmar tubercle bifid; distinct supernumerary tubercles (Fig. 4 C), (9) ulnar and tarsal tubercles absent; (10) heel lacking tubercles; (11) toes lacking lateral fringes; webbing absent; Toe V slightly longer than Toe III; toe discs expanded, rounded (Figs. 4 B, D); (12) inner metatarsal tubercle oval, about twice the size of outer metatarsal tubercle; supernumerary plantar tubercles present (Fig. 4 D); (13) in ethanol, dorsal coloration variable (Figs. 1, 6): brown, pale brown-cream, brown-reddish, brown-grayish or gray, with dark brown scapular and sacral marks; white mid-stripe can be present or absent; brilliant silver and/or bronze chromatophores distributed on dorsum; venter cream to brown; throat pale brown to brown; minute white and brown dots on venter and throat seen under magnification. In life (Fig. 2), dorsum brown-orange to yellowish brown with darker marks, flanks as dorsum but with small white dots; venter grayish brown with minute white dots; belly partially translucent; red iris with thin black reticulation; (14) SVL in females 15.3���18.9 mm (mean = 16.9 �� 1.0, n = 25), in males 9.5���13.7 mm (mean = 12.2 �� 0.8, n = 46). Similar species. Pristimantis minimus is easily distinguished from most species in Pristimantis by its minute body size. Other small South American Pristimantis, although larger than P. minimus, include P. coronatus (SVL = 15.3 mm in the only known specimen, a female; Lehr & Duellman 2007), P. andinognomus (SVL = 10.0���14.5 mm in adult males; 12.6���17.9 mm in adult females; Lehr & Coloma 2008), and P. trachyblepharis (SVL = 12.1���15.8 mm in adult males; 15.8���19.2 mm in adult females; Lynch & Duellman 1980). Pristimantis coronatus has an occipital fold, ulnar tubercles, conical tubercles on the upper eyelid, and a red-orange coloration on flanks and groin (absent in P. minimus). Pristimantis andinognomus presents dorsolateral and postocular folds, conical tubercles on the upper eyelid and heel, tarsal tubercles, and a distinctive tympanic membrane (absent in P. minimus). Finally, P. trachyblepharis is distinguished by lacking vocal slits (present in P. m i n i m u s) and having an externally evident tympanum (concealed in P. m i n i m u s). Other small species of Pristimantis from Ecuador are notably different from P. m i n i m u s. Males of P. caeruleonotus are slightly larger (11.6���14.6 mm in males; 20.5���22.6 mm in females; Lehr et al. 2007) than P. minimus. Additionally, P. caeruleonotus presents a visible tympanum and lateral fringes in fingers, characteristics that are absent in P. minimus. From other relatively small species, P. minimus is differentiated mainly by the smaller size, but also the absence of conical upper eyelid tubercles, which are present in P. colodactylus (Lynch, 1979), P. hectus (Lynch & Burrowes, 1990), P. leoni (Lynch, 1976) and P. pyrrhomerus (Lynch, 1976). Additionally, P. l e o n i and P. pyrrhomerus bear ridges on the dorsum and inner and outer tarsal tubercles (absent in P. minimus). Pristimantis species that were found in sympatry with P. minimus include P. diadematus, P. trachyblepharis, and P. katoptroides; at Cordillera del C��ndor, these three species have a larger body size that does not overlap with P. minimus [in P. trachyblepharis, SVL (males) = 13.7���16.2 mm, n = 15, SVL (females) = 18.9���22.2 mm, n = 6; in P. diadematus, SVL (males) = 17.9���26.9 mm, n = 8, SVL (females) = 26.7���32.2 mm, n = 6; P. katoptroides, SVL in one adult female = 32.3 mm]. Moreover, P. katoptroides is easily distinguished from all other Pristimantis by having a green venter with white reticulation and a blue groin. Description of the holotype. Adult male (QCAZ 41612; Figs. 1, 2 B). Head as wide as body, slightly longer than wide, lacking tubercles; head width 35 % of SVL; head length 38 % of SVL; snout short, rounded in dorsal and lateral views (Fig. 2); tongue oval, with a small notch on its posterior border; eye diameter slightly larger than eye-nostril distance; nostrils slightly protuberant, directed anterolaterally; canthus rostralis straight in dorsal view and in profile; loreal region weakly concave; upper eyelid width 74 % of interorbital distance, lacking tubercles; tympanic membrane absent; tympanic annulus present, but not visible externally. Vocal slits straight, located at the level of the junction of the upper and lower lips, between tongue and margin of jaw; vocal sac small, restricted to throat. Skin on dorsum shagreen without tubercles or folds (Fig. 1); skin on venter shagreen, discoidal or thoracic folds absent (Fig. 2 B); cloacal sheath absent; cloacal region without tubercles. Ulnar tubercles absent; outer palmar tubercle bifid, about the same length as oval inner palmar tubercle; subarticular tubercles rounded; distinct, non-conical supernumerary palmar tubercles, lower than subarticular tubercles; fingers lacking lateral fringes; Finger I shorter than Finger II; disc of Finger I slightly expanded, all other discs expanded (Fig. 4 A); ventral pads well defined by circumferential grooves; nuptial pads absent. Tibia length 54 % of SVL; foot length 46 % of SVL; surfaces of legs smooth, lacking tubercles; inner metatarsal tubercle ovoid, about twice the size of outer metatarsal tubercle; subarticular tubercles rounded, well defined; plantar supernumerary tubercles present, smaller than subarticular tubercles; toes lacking lateral fringes; webbing absent; discs expanded, round to acute, similar to those on fingers; toes with ventral pads well defined by circumferencial grooves; relative length of toes: I Measurements of holotype (in mm). SVL 11.9; tibia length 5.6; foot length 5.5; head length 4.5; head width 4.2; eye diameter 1.8; interorbital distance 1.4; upper eyelid width 1.0; internarial distance 1.5; eyenostril distance 1.2. Coloration of holotype in preservative. Background color of dorsum light brown-cream with a continuous large dark brown mark from snout to sacral region, only divided by a white, thin middorsal stripe and a white interorbital band (Fig. 1); upper lip with dark brown bars below eyes; dark brown supratympanic fold; silver and bronze brilliant sparkling distributed on dorsum; dark brown mark surrounding the cloaca; minute white dots on the flanks; dorsally, arms and legs with pink hue, barred thighs, forearm with one brown stripe; ventral coloration (including arms and legs) cream with numerous minute brown dots and scarce white dots seen under magnification (Fig. 1). Coloration of holotype in life (based on observations in the field and on photographs; Fig. 2 B). Brownorange dorsum, with a thin middorsal line and a large brown mark; flanks with same coloration as dorsum, but with small white dots; venter grayish brown with minute white dots; belly translucent, therefore the heart, liver, and viscera are partially visible through the skin; red iris with thin black reticulation. Variation. Meristic variation is presented in Table 1. Some paratypes have the discs of the fingers more expanded than the holotype (QCAZ 41422, 41432, 41511, 41691, 41711, 41716; Fig. 5 A), and others have the tip of the discs slightly acuminate (QCAZ 40744, 40745, 41444, 41606, 41616, 41620, 41672, 41694, 41738; Fig. 5 B). In ethanol, two paratypes have two white dorsolateral stripes starting at the nostrils and ending just anterior to the cloaca (QCAZ 41430, 41537; Fig. 6 A). Most paratypes do not have the white, thin middorsal stripe (QCAZ 40744, 40745, 41378, 41380, 41422, 41426, 41429, 41431, 41432, 41433, 41443, 41444, 41497, 41511, 41548, 41551, 41553, 41602, 41620, 41631, 41635, 41650, 41660, 41661, 41666, 41689, 41690, 41691, 41693, 41694, 41695, 41701, 41711, 41713, 41714, 41716, 41718, 41721, 41722, 41723, 41738; Fig. 6 D). The background coloration of the dorsum varies from gray (QCAZ 41422, 41429, 41433, 41444, 41498, 41499, 41500, 41548, 41613, 41650, 41661, 41664, 41667, 41671, 41672, 41673, 41691; Fig. 6 B) to brown-grayish (QCAZ 40744, 41380, 41443, 41549, 41600, 41602, 41607, 41631, 41643, 41665, 41666, 41668, 41670, 41690, 41695, 41713, 41714; Fig. 6 C), brown (QCAZ 40745, 41378, 41431, 41432, 41511, 41614, 41616, 41617, 41620, 41621, 41623, 41426, 41630, 41633, 41634, 41651, 41660, 41689, 41693, 41694, 41701, 41711, 41716, 41718, 41721, 41722, 41723; Fig. 6 D) and brown-reddish (QCAZ 41619, 41496, Fig. 6 E). Some paratypes have a scapular mark W-shaped and a sacral V-shaped mark (QCAZ 41378, 41422, 41429, 41433, 41444, 41497, 41548, 41551, 41553, 41602, 41660, 41661, 41667, 41671, 41672, 41691, 41738; Fig. 6 B). The venter of most paratypes (specially those with brown, brown-grayish or brown-reddish dorsum) is cream brown, being the throat darker than the belly (Figs. 6 F, 6 G); in preservative, the belly is not as translucent as in life. Specimens in life show low and scarce tubercles on the upper eyelid, dorsum and legs, but those tubercles are not visible in preservative., Published as part of Ter��n-Valdez, Andrea & Guayasamin, Juan M., 2010, The smallest terrestrial vertebrate of Ecuador: A new frog of the genus Pristimantis (Amphibia: Strabomantidae) from the Cordillera del C��ndor, pp. 53-68 in Zootaxa 2447 on pages 54-57, DOI: 10.5281/zenodo.195104, {"references":["Hedges, S. B., Duellman, W. E. & Heinicke, P. (2008) New world direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182.","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. 2009. Phylogenetic systematics of glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Frost, D. R. (2009) Amphibian Species of the World: an Online Reference. Version 5.3 Available from: http: // research. amnh. org / herpetology / amphibia / (22 March, 2010).","Lehr, E. & Duellman, W. E. (2007) A diminutive new species of Pristimantis (Amphibia: Anura: Leptodactylidae) from northern Peru. Salamandra, 43, 165 - 171.","Lehr, E. & Coloma, L. A. (2008) A minute new Ecuadorian Andean frog (Anura: Strabomantidae, Pristimantis). Herpetologica, 64, 354 - 367.","Lynch, J. D., & Duellman, W. E. (1980) The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Miscellaneous Publications. Museum of Natural History, University of Kansas, 69, 1 - 86.","Lynch, J. D. (1979) Leptodactylid frogs of the genus Eleutherodactylus from the Andes of southern Ecuador. Miscellaneous Publications. Museum of Natural History, University of Kansas, 66, 1 - 62.","Lynch, J. D. & Burrowes, P. A. (1990) The frogs of the genus Eleutherodactylus (Family Leptodactylidae) at the La Planada Reserve in southwestern Colombia with descriptions of eight new species. Occasional Papers of the Museum of Natural History, University of Kansas, 136, 1 - 31.","Lynch, J. D. (1976) Two new species of frogs of the genus Euparkerella (Amphibia: Leptodactylidae) from Ecuador and Peru. Herpetologica, 32, 48 - 53."]}

Family: Centrolenidae Glassfrogs have a Neotropical distribution and are particularly diverse in the northern Andes. All centrolenids have a partial or complete ventral transparency, a dilated medial process on Metacarpal III, and deposit their eggs out of water (usually on the upper or under sides of leaves; Hayes & Starrett 1980; Ruiz-Carranza & Lynch 1991; Guayasamin et al. 2009). They are usually found at night on vegetation along streams when reproducing. Herein, we follow the taxonomy proposed by Guayasamin et al. (2009) that recognizes 12 genera and 147 species within Centrolenidae. In YBS, there are five identified (Centrolene bacatum, C. buckleyi, Nymphargus posadae, N. siren, N. wileyi) and two unidentified (Centrolene sp, Cochranella sp) species of glassfrogs. guacamayo Rhinella margaritifera 2000���2200 Open pasture, Small ponds, ditches, 3 / 4 / 2 /0/ 1 Nocturnal disturbed habitat and pools in pasture Centrolenidae Centrolene bacatum 2000���2200 Primary forest and Leaves 1 / 9 /0/0/0 Nocturnal bamboo along streams Centrolene buckleyi 2000���2200 Primary forest along Leaves 0/ 2 /0/0/ 1 Nocturnal streams Centrolene sp 1 2100 Primary forest along Leaves 0/ 3 /0/0/0 Nocturnal streams Cochranella sp 1 2100 Primary forest along Leaves 1 /0/0/0/0 Nocturnal streams Nymphargus posadae 2000���2200 Primary forest along Ferns, leaves 1 / 3 /0/0/0 Nocturnal streams Nymphargus siren 2100 Primary forest along Leaves 0/ 2 /0/0/0 Nocturnal streams Pristimantis eriphus 2000���2300 Primary and secondary Leaves, ferns, 10 / 3 / 9 / 14 / Nocturnal forest bromeliads 27 Pristimantis cf. eriphus 2000���2200 Primary and secondary Leaves 0/ 1 / 1 /0/ 1 Nocturnal forest Pristimantis inusitatus 2000���2200 Primary and secondary Leaves, ferns 10 / 7 / 2 / 2 / 14 Nocturnal forest Pristimantis gladiator 2270���2500 Primary forest Low vegetation, 0/0/0/0/ 2 Nocturnal forest floor Pristimantis leucopus 2000���2200 Primary forest High leaves above 1 / 2 /0/0/0 Nocturnal streams Pristimantis cf. petersi 2000���2200 Primary forest Leaves and low 7 / 6 / 9 / 49 / 27 Nocturnal vegetation Pristimantis w-nigrum 2000���2200 Pasture, forest edge, Hidden in grass in 0/ 7 / 3 / 7 / 8 Nocturnal secondary and primary pastures, forest floor, forest low vegetation Pristimantis sp 1 2000���2200 Primary forest Low vegetation 0/0/0/ 36 /0 Nocturnal Caeciliidae Caecilia orientalis 2100 Pasture and secondary Fossorial, under 1 / 3 /0/ 1 / 2 Unknown forest rotten logs, comes to surface during hard rains Centrolene bacatum (Plate 1). Described by Wild (1994). This species is easily distinguished from other amphibians in YBS by having a green dorsal coloration with few white tubercles, some of which form a line that extends from below the eye to the insertion of the arm (Wild 1994; Guayasamin et al. 2006 a). As all species included in the genus Centrolene, C. bacatum has a venter that is posteriorly translucent. Also, adult males of this species have a humeral spine (absent in all Nymphargus species, except N. grandisonae and some individual in N. griffithsi, both species found on the western slope of the Andes). Centrolene bacatum is smaller than C. buckleyi [25.3���30.5 mm (n = 23) in adult males of C. buckleyi]. In YBS, SVL in C. bacatum males, 19.3���21.8 mm (n = 11); SVL = 20.4 mm in one adult female. Active, calling males of Centrolene bacatum are frequently heard in YBS during the night; however, they usually call from areas close to the canopy; females deposit the egg masses on the upper and undersides of leaves, in vegetation along streams. Centrolene buckleyi (Plate 1). ��� Described by Boulenger (1882). Among the glassfrogs found at YBS, Centrolene buckleyi is easily distinguished by having green dorsal surfaces sharply demarcated laterally from white lower flanks, a humeral spine (in males), white upper lip, a moderate adult size (SVL in males 25.3��� 30.5 mm; in females 29.8���34.4 mm) and a sloping snout in lateral profile (Guayasamin et al. 2006 a). Remarks: Genetic analyzes suggests that Centrolene buckleyi represents a species complex (Guayasamin et al. 2008), a hypothesis that needs further examination with a dense sampling in Ecuador and Colombia. Nymphargus posadae (Plate 2). ��� Described by Ruiz-Carranza and Lynch (1995). This species can be differentiated from other glassfrogs by lacking webbing between fingers, lacking humeral spines, having a green dorsum with small greenish-white warts, and by having a white upper lip, and a white line on the ventrolateral border of arm, Finger IV, tarsus and Toe V (Guayasamin et al. 2006 a). Nymphargus posadae is differentiated from Centrolene buckleyi and C. bacatum mainly by lacking humeral spines, which are present in the adult males of the two Centrolene species. Also, N. posadae lacks webbing between fingers, whereas there is some webbing between the two externalmost fingers of C. bacatum and C. buckleyi. At YBS, there are two additional glassfrogs that belong to the genus Nymphargus (i.e., N. siren and N. wileyi). These three species share the absence of webbing between fingers. Nymphargus siren differs from the other two species (and any other amphibian in YBS) by presenting small yellow spots on the dorsum, whereas N. wileyi has a uniform green dorsum (in N. posadae, green dorsum with small greenish-white warts). SVL in males is 30.7��� 34.1 mm (x = 32.3, n = 6); in females 30.2���33.3 mm (x = 31.4, n = 4). Remarks: Males of Nymphargus posadae collected in YBS are smaller (SVL = 30.7���31.9 mm, n = 3) than Colombian specimens (SVL = 32.7���34.1 mm, n = 3; Ruiz-Carranza & Lynch 1995; Guayasamin et al. 2006 a). Nymphargus siren (Plate 2). ��� Described by Lynch and Duellman (1973). In YBS, this glassfrog is the only amphibian that has a green dorsum with small yellow spots. Additionally, N. siren lacks webbing between fingers, lacks humeral spines, and is relatively small (SVL Nymphargus siren seems to be a rare species in YBS; only two males have been found during the two years of surveys. Nymphargus wileyi (Plate 2). ��� Described by Guayasamin et al. (2006 a). Nymphargus wileyi differs from other Glassfrogs by having a uniform green dorsum, white renal peritoneum, and by lacking membranes between Fingers III and IV. In adult males, SVL 23.4���26.5 mm (x = 24.6; n = 11); 25.1���28.1 mm in adult females (x = 26.3; n = 4)., Published as part of Guayasamin, Juan M. & Funk, Chris, 2009, The amphibian community at Yanayacu Biological Station, Ecuador, with a comparison of vertical microhabitat use among Pristimantis species and the description of a new species of the Pristimantis myersi group, pp. 41-66 in Zootaxa 2220 on pages 43-46, DOI: 10.5281/zenodo.190060, {"references":["Hayes, M. P. & Starrett, P. H. (1980) Notes on a collection of Centrolenid Frogs from the Colombian Choco. Bulletin of the Southern California Academy of Science, 79, 89 - 96.","Ruiz-Carranza, P. M. & Lynch, J. D. (1991) Ranas Centrolenidae de Colombia I. Propuesta de una nueva clasificacion generica. Lozania, 57, 1 - 30.","Guayasamin, J. M., Castroviejo-Fisher, S., Trueb, L., Ayarzaguena, J., Rada, M. & Vila, C. (2009) Phylogenetic systematics of glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1 - 97.","Wild, E. R. (1994) Two new species of centrolenid frogs from the Amazonian slope of the Cordillera-Oriental, Ecuador. Journal of Herpetology, 28, 299 - 310.","Guayasamin, J. M., Bustamante, M. R., Almeida-Reinoso, D. & Funk, W. C. (2006 a) Glass frogs (Centrolenidae) of Yanayacu Biological Station, Ecuador, with the description of a new species and comments on centrolenid systematics. Zoological Journal of the Linnean Society, 147, 489 - 513.","Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum. Trustees, British Museum (Natural History), London, 495 pp.","Guayasamin, J. M., Castroviejo-Fisher, S., Ayarzaguena, J., Trueb, L. & Vila, C. (2008) Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 48, 574 - 595.","Ruiz-Carranza, P. M., & Lynch, J. D. (1995) Ranas Centrolenidae de Colombia V: cuatro nuevas especies de Cochranella de la Cordillera Central. Lozania, 62, 1 - 23.","Lynch, J. D., & Duellman, W. E. (1973) A review of the centrolenid frogs of Ecuador, with descriptions of new species. The University of Kansas Museum of Natural History Occasional Papers, 16, 1 - 66."]}

Noblella coloma Guayasamin & Ter��n, new species Holotype. QCAZ 26307, adult male (Figs. 1, 2), collected from Reserva Flor��stica R��o Guajalito (0�� 14 ' S, 78 �� 49 ' W; 1800���2000 m), 3 km NW from Km 59 on the old road from Quito to Santo Domingo, collected by Rub��n D. Jarr��n on 18 October 2003. Paratopotypes. QCAZ 7277, adult female, collected by Leonora Orejuela on 22 Abril 1995; QCAZ 8701, juvenile female, collected by Santiago Burneo on 15 October 1995; QCAZ 11614, juvenile, collected by ��talo G. Tapia, date unknown; QCAZ 32702, juvenile, collected by Cristina E. Toapanta on 16 December 2006; QCAZ 6466, sex unknown, collected by George Fletcher on 23 October 1994, cleared-and-stained; QCAZ 7412, sex unknown, collected by Juan M. Guayasamin on 18 June 1995, cleared-and-stained; QCAZ 40579, juvenile female, collected by Diana Pazmi��o on 15 March 2009. Generic placement. Duellman (1991), De la Riva et al. (2008), and Hedges et al. (2008) list the following traits as characteristic of Noblella: (1) head narrower than body; (2) tympanic membrane differentiated (except in N. duellmani); (3) cranial crests absent; (4) dentigerous processes of vomers absent; (5) ���S��� condition of adductor muscle; (6) terminal discs on fingers and toes not expanded or slightly expanded; discs and circumferential grooves present distally (except in N. duellmani); terminal phalanges narrowly T- shaped; (7) Finger I shorter than, or equal in length to, Finger II; (8) Toe III shorter than Toe V; tips of at least Toes III���IV pointed; (9) subarticular tubercles not protruding; (10) conspicuous tarsal tubercle; (11) dark inguinal spots present (except in N. duellmani); (12) small size (SVL Noblella; therefore, the placement of Noblella coloma in the genus Noblella is justified. The only other possibility for the generic placement of the new species is Barycholos, a genus endemic to South America that has a close evolutionary and morphological affinity with Noblella (Heinicke et al. 2007; Hedges et al. 2008). However, species in Barycholos have vomers with small dentigerous processes (absent in Noblella), Finger I longer than II (Finger I shorter than, or equal in length to, Finger II in Noblella), and projecting subarticular tubercles (not projecting in Noblella). Diagnosis. Noblella coloma (Figs. 1, 2) presents the following characteristics: (1) skin of the dorsum finely shagreen; (2) tympanic annulus and membrane clearly visible and differentiated, supratympanic fold covering upper border of tympanum (Fig. 2 C); (3) snout rounded in dorsal and lateral view (eye to nostril distance 48.8���58.9% of eye diameter, Fig. 2 C); (4) dentigerous processes of vomers absent; (5) fingers not expanded distally; finger tips acuminate; Finger I slightly shorter than Finger II (Fig. 3); nuptial pad not visible; fingers lacking circumferential grooves; (6) distal phalanges T-shaped; phalangeal formula of hands: 2, 2, 3, 3 (Fig. 5); (7) hands with one weakly defined supernumerary tubercle; subarticular tubercles rounded, not prominent; ulnar tubercles absent; (8) one elongated tarsal tubercle (Fig. 3); two prominent metatarsal tubercles; supernumerary plantar tubercles absent; toes slightly expanded and acuminate distally; (9) Toe V shorter than Toe III; distal portions of circumferential grooves present on Toes III���V; (10) phalangeal formula of feet: 2, 2, 3, 4, 3 (Fig. 5); (11) in life, dorsum uniform brown, except for distinctive suprainguinal marks; flanks with dark brown band that narrows as it approaches groin; venter orange with minute brown and white spots (Fig. 1); (12) in one adult male, SVL 14.6 mm; in one adult female, 16.0 mm. Noblella coloma differs from all the other species in Noblella by lacking most of the dorsal color marks (i.e., interobital bar, scapular and sacral chevrons; Fig. 2), and by having a bright orange venter (Fig. 1). Additionally, the dorsum of N. coloma is shagreen, whereas the dorsum of N. duellmani and N. lynchi is pustular, and N. pygmaea has a tubercular dorsal skin. Noblella coloma is similar to N. duellmani, N. heyeri, N. lynchi and N. pygmaea in having three phalanges in Finger IV, in contrast with N. carrascoicola, N. lochites, N. myrmecoides, and N. ritarasquinae that have two phalanges in Finger IV. A tympanum is evident in N. coloma, N. heyeri, N. lynchi, N. myrmecoides, N. peruviana, and N. pygmaea, and is absent in N. duellmani, N. carrascoicola, and N. ritarasquinae. In preservative, N. coloma has a dark throat and a uniform cream venter, whereas the venter is brown with cream mottling in N. duellmani, N lynchi, N. heyeri, and N. carrascoicola (Fig. 6). Comparison of the new species with Noblella peruviana are difficult because of the paucity of information regarding the morphology of the later. All what we know about N. peruviana comes from the brief description provided by Noble (1921) and De la Riva et al. (2008), both based on five specimens collected more than a century ago. Characters mentioned in Noble���s original description include: tympanum present, digits pointed with no terminal disks, well-developed and elongated tarsal tubercle, skin feebly granular above and smooth below, dorsal coloration uniform gray brown or with dark brown blotches, and broad band of dark brown extending on each side from the nostril to the lumbar region. Additionally, De la Riva et al. (2008) mentioned that the types of N. peruviana have the tips on Toes II, III and IV distinctively pointed, and that, at least in some paratypes, a black inguinal spot is present. The traits listed above for N. peruviana are present in several other species of the genus; therefore, comparisons are limited. However, Noble (1921) mentioned that, in preservative, the ventral coloration in N. peruviana was reddish brown; in contrast, the ventral coloration of N. coloma is cream. In addition, N. coloma and N. heyeri are the only species in the genus found on the western slopes of the Andes (Table 1); this being a biogeographical criterion that could be useful to distinguish among species. Description of holotype. Adult male (QCAZ 26307); head as long as wide; snout round in dorsal and lateral views (Fig. 2 C); canthus rostralis straight; loreal region concave; upper eyelid about 50 % of interorbital distance; eye-nostril distance 59 % of eye diameter; tympanum visible almost completely, tympanic membrane well differentiated from surrounding skin; supratympanic fold covering upper border of tympanum. Dentigerous processes of vomers absent. Skin of dorsum finely shagreen; venter smooth; ulnar tubercles absent; palmar tubercle oval, about 1.5x the size of thenar tubercle; one supernumerary tubercle weakly marked; proximal subarticular tubercles large, rounded; fingers not expanded distally, finger tips slightly acuminate (Fig. 3), circumferential grooves absent; relative lengths of fingers I heyeri Ecuador, Peru: Andes of northern Peru (33 km SW of 2000���3100 m Lynch (1986), this work Huancabamba) and southern Ecuador (Provincia de Loja). Color of holotype in alcohol. Dorsum grayish brown to brown with dark brown inguinal spots; scapular and sacral chevrons absent. Sides of head uniform black continuous posteriorly onto flanks; labial bars absent; each flank with a wide longitudinal dark brown stripe that narrows posteriorly. Forearms and hind limbs with well-defined dark brown marks (Fig. 2). Throat dark brown; venter cream with minute dark spots (visible under magnification). Coloration in life. Dorsum orange-brown, with a thin dark brown middorsal stripe and dark brown inguinal spots; interobital bar and scapular and sacral marks absent. Venter orange with minute white and brown spots (Fig. 1). Sides of head dark brown; flanks with dark brown stripe that narrows posteriorly. Iris dark brown with orange ring around pupil (based on photographs of QCAZ 32702, holotype, by Luis A. Coloma). Variation. Throat cream (QCAZ 32702) or cream with minute brown spots visible under magnification (QCAZ 11614). There is no evident sexual dimorphism. For measurements see Table 2. 7277 26307 8701 40579 11614 32702 Sex Adult female Adult Male Juv. Female Juv. female Juvenile Juvenile Osteology. The following osteological description of Noblella coloma is based on a cleared-and-double stained individual (QCAZ 6466, SVL = 14.1 approximately; Figs. 4, 5). The specimen lacks information regarding its sex, but because of its SVL we assume that it could be an adult or subadult male or juvenile female. Drawings of hand and foot are based on QCAZ 7412. Cranium. Shape and proportions. The skull is widest posterior to the orbit at the level of the articulation of the maxilla with the quadratojugal (Fig. 4); at this level, the skull is as wide as long. The rostrum is short, accounting for less than a quarter (17 %) of the medial length of the skull. At the level of midorbit, the width of the braincase is about 35 % of the greatest width of the skull and 38 % of the medial skull length. Neurocranium. The anterior neurocranium comprises large olfactory capsules, which are cartilaginous, and the anterior part of the braincase. The medial walls of the nasal capsules are narrowly separated, with the internasal septum apparently being composed of a thin plate of cartilage between the capsule walls; a minute, ossified septomaxilla is present, but obscured by the staining of the anterior nasal capsule cartilages (Fig. 4). The cartilaginous planum antorbitale has a slight anterolateral orientation in dorsal and ventral aspects. Ventrally, the planum is invested by a simple, thin and relatively long neopalatine (Fig. 4). The distal end of the neopalatine is clearly separated from the maxilla; the medial end articulates with the sphenethmoid (Fig. 4). The braincase and otic capsules are moderately ossified. Ventrally, the bony sphenethmoid is not fused medially (Fig. 4). There is a moderately broad separation between the sphenethmoid and prootics, within which the optic fenestra lies (Fig. 4). The optic fenestra lies in cartilage, except for its posterior margin that is formed by the prootic bone. Dorsally and ventrally, the exoccipitals are separated from each other and from the prootics (Fig. 4). The contralateral prootics are completely separated from each other; each one bears a relatively long prootic foramen, and two small foramina, the oculomotor and trochlear. The frontoparietal fontanelle is partially exposed medially between the frontoparietal (Fig. 4). Dorsally, the otic capsule is well ossified, and bears a moderately broad, cartilaginous crista parotica. The epiotic eminences are prominent; the anterior eminence is slightly longer than the posterior and the angle between the two arms is approximately 90 &ring;. Ventrally, the lateral wall of the otic capsule is cartilaginous. The cartilaginous operculum lies posteriorly adjacent to the pars interna plectri of the stapes in the fenestra ovalis, which is formed entirely in cartilage. The pars media plectri is a long, slender, ossified element that extends anterolaterally beneath the crista parotica and behind the palatoquadrate and ventral ramus of the squamosal to terminate in the club-shaped cartilaginous pars externa plectri located in the middle of the incomplete tympanic annulus (Fig. 4). Dorsal investing bones. Dorsal investing bones are poorly developed. The nasals are thin and widely separated from one another; they cover a large area of the nasal capsules dorsally. The frontoparietals are well-developed bones, separated medially, and separated from the prootics. The frontoparietal bears a lamina perpendicularis, the vertical component forming the dorsolateral edge of the braincase (Fig. 4). Ventral investing and palatal bones. Owing to the thinness of the bone, the posterior margin of the parasphenoid is difficult to determine accurately. The cultriform process (ca. 39 % the width of the braincase) extends anteriorly reaching about the midlength of the sphenethmoid (Fig. 4). The anterior end of the process is tapered; the parasphenoid is separated from the neopalatines. The lateral margins of the process are approximately parallel; anteriorly, the margins converge gradually towards one another and make contact with the medial margin of the sphenoid. The parasphenoid alae are moderately long (about equal to the width of the cultriform process), slightly posterolaterally oriented, and distally truncate beneath the midwidth of the otic capsules. A posteromedial process is present, but distinctly separated from the margin of the foramen magnum (Fig. 4). The small vomers are broadly separated from one another medially. Each is composed of an arcuate bone bordering the medial and small portions of the anterior and posterior margins of the choana. The prechoanal and postchoanal ramus are slightly expanded anteriorly; the dentigerous process is absent (Fig. 4). The neopalatines are thin and underlie the plana antorbitalae, covering the posterior surfaces and dorsomedial surfaces of these structures (Fig. 4). The bones are unornamented, straight, and articulate with the sphenethmoid; each neopalatine is separated from the maxilla. Maxillary arcade. The maxillary arcade bears teeth on the premaxillae and maxillae. The maxillae are in close proximity to the quadratojugals, but have no contact (Fig. 4). The partes palatinae of the maxillae are narrow and barely evident. The premaxilla bears prominent partes palatinae; the posterolateral palatinae process is more robust than the posteromedial process (Fig. 4). There is a simple, juxtaposed articulation between the anterior end of the maxilla and the premaxilla. The pars facialis of the maxilla is well developed anteriorly, having a preorbital process that covers the planum antorbitale along the lateral aspect of the olfactory capsule (Fig. 4). Suspensory apparatus. The triradiate pterygoid bears a curved anterior ramus that is oriented anterolaterally toward the maxilla, with which it has an area of close proximity. The pterygoid is separated from the maxilla by the pterygoid cartilage, which lies along the medial margin of the maxilla in the orbital region. The medial and posterior rami of the pterygoid are about equal in length; however, the medial ramus is more robust than the posterior. The edge of the medial ramus overlaps the lateral edge of the ossified margin of the prootic. The squamosal is T-shaped; the otic ramus is much more longer than the zygomatic ramus (otic ramus 3 x the length of the zigomatic ramus). The otic ramus overlaps a large area of the crista parotica. The ventral ramus invests the lateral surface of the palatoquadrate, and articulates with the quadratojugal (Fig. 4). Postcranium Vertebral column. There are eight presacral vertebrae (Fig. 5). Presacral I is notably shorter than the posterior presacrals, and Presacral III is relatively longer than the other presacral vertebrae. All of the presacrals are non-imbricate. The vertebral profile in increasing order of overall width is: I Pectoral girdle. The prezonal portion is represented by a short cartilaginous omosternum. The clavicles are oriented anteromedially, with the medial tips distinctly separated from one another and located at about the same level of the anterolateral end of the clavicle that articulates with the scapula. The coracoid is stout, with the glenoidal and sternal ends about equally expanded. The midshaft width is about 13 % the length of the coracoid, and about 41 % the width of the expansion of the sternal end of the bone. The pectoral fenestra is oval and about two times as wide as it is deep. The scapula is long with a prominent pars acromialis that is not separated from the pars glenoidalis. The clavicle is 75 % the length of the scapula. The suprascapula is mineralized, with the ossified cleithrum apparent as a slender bone along the leading edge of the suprascapular blade and with an expansion of its proximal end. The sternum is mineralized and narrowly separated from the epicoracoid (Fig. 5). Pelvic girdle. The long, slender ilial shafts bear conspicuous dorsolateral crests. The ilial prominence is broad and low; the pubes is cartilaginous. The round acetabulum is ossified (Fig. 5). Manus and pes. The phalangeal formulae for the hand and foot are standard: 2 - 2-3 - 3 and 2 - 2-3 - 4 - 3, respectively. In increasing order of length, the order of the digits on the hand is: I Etymology. The specific name coloma is a patronym for Luis Aurelio Coloma in recognition of his continual efforts in studying and protecting amphibians, and mentoring students (us included). The specific epithet is used as a substantive in apposition. Distribution and conservation. Noblella coloma is known only from the type locality, Reserva Flor��stica R��o Guajalito (0�� 14 ' S, 78 �� 49 ' W; 1800���2000 m), 3 km NW, by path, from Km 59 on the Quito-Chiriboga- Santo Domingo road (Fig. 7). All individuals of N. coloma have be, Published as part of Guayasamin, Juan M. & Ter��n-Valdez, Andrea, 2009, A new species of Noblella (Amphibia: Strabomantidae) from the western slopes of the Andes of Ecuador, pp. 47-59 in Zootaxa 2161 on pages 48-57, DOI: 10.5281/zenodo.188980, {"references":["Duellman, W. E. (1991) A new species of leptodactylid frog, genus Phyllonastes, from Peru. Herpetologica, 47, 9 - 13.","Hedges, S. B., Duellman, W. E. & Heinicke, P. (2008) New world direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182.","Noble, G. K. (1921) Five new species of Salientia from South America. American Museum Novitates, 29, 1 - 7.","Lynch, J. D. (1986) New species of minute Leptodactylid frogs from the Andes of Ecuador and Peru. Journal of Herpetology, 20, 423 - 431."]}

Pristimantis bicantus, new species Holotype. Adult female QCAZ 19024, collected at Yanayacu Biological Station, trail to San Isidro Lakes (0�� 36 ' S, 77 �� 53 'W; 2150 m), Provincia Napo, Ecuador, on 10 January 2001 by Fernando Nogales and Diego Almeida-Reinoso. Paratopotypes. All specimens were collected along different trails within Yanayacu Biological Station, Provincia Napo, Ecuador, or nearby the scientific station, at elevations between 2100 and 2300 m: High trail, QCAZ 19003 ���08, collected by Mart��n R. Bustamante on 2 January 2001; QCAZ 19021 ��� 23, collected by Claudia Arcanjo on 2 January 2001; QCAZ 19025 ��� 28, collected by WCF on 2 January 2001; QCAZ 39813, 39818��� 19, collected by Mart��n R. Bustamante on 3 January 2005. Macuca loma, QCAZ 18996, 18999, collected by Mart��n R. Bustamante and WCF on 29 December 2000; QCAZ 19044 ���45, 19056��� 57, collected by Mart��n R. Bustamante on 14 January 2002; QCAZ 18979, 18985���89, 18997, collected by WCF on 11 January 2001; QCAZ 18990 ���91, 18998, collected by WCF on 0 1 January 2001; QCAZ 18983 ��� 84, collected by WCF on 11 January 2001; QCAZ 18997 ��� 98, 19000 ���002, collected by Mart��n R. Bustamante and WCF on 0 1 January 2001. Macuca loma creek number 4, QCAZ 22379, collected by Mart��n R. Bustamante and Galo Diaz on 25 October 2002. San Isidro lakes, QCAZ 19017 ��� 18, collected by Fernando Nogales and Diego Almeida-Reinoso on 10 January 2001; QCAZ 39820, collected by Fernando Nogales and Diego Almeida- Reinoso on 11 January 2005. Small San Isidro lake, QCAZ 18980, 18981, collected by WCF on 10 January 2001. Trail to San Isidro lakes, QCAZ 18982, 18993, 19019 ��� 20, collected by WCF on 10 January 2001. Upper old station creek, QCAZ 19039 ���43, 19046��� 55, collected by Mart��n R. Bustamante on 17���18 January 2002. Upper trail, QCAZ 19030 ��� 38, collected by Mart��n R. Bustamante on 15 January 2002; QCAZ 22378, 22380��� 81, collected by Mart��n R. Bustamante and Galo Diaz on 15 August 2002. Waterfall trail, QCAZ 19029, collected by WCF on 5 January 2002; QCAZ 39814 ��� 17, collected by Mart��n R. Bustamante, Diego Almeida-Reinoso, and Fernando Nogales on 0 6 January 2005. Nearby scientific station, QCAZ 18978, collected by Diego Almeida-Reinoso and Fernando Nogales on 16 January 2001. Generic placement. As defined by Lynch and Duellman (1997) and Hedges et al. (2008), the Pristimantis (Pristimantis) myersi group contains small frogs (adult females P. leoni and P. ocreatus); cranial crests are absent; vocal slits are present (except in P. floridus); and vomerine teeth are present. The morphology of Pristimantis bicantus agrees with all the diagnostic traits of the P. myersi species group (Lynch & Duellman 1997; Hedges et al. 2008), except that the tympanic annulus is relatively larger in males than in females, and tarsal tubercles are absent. Diagnosis. (1) Skin on dorsum shagreen with small scattered, rounded warts; upper flanks with numerous low warts; occipital and dorsolateral folds usually absent (but see variation; Figs. 1, 2); (2) tympanic membrane and tympanic annulus well differentiated (Fig. 1); tympanic annulus sexually dimorphic; tympanum in males 6.87���10.22 % (x = 7.99, n = 32) SVL, tympanum in females 4.28���6.15 % (x = 5.23, n = 33) SVL; (3) snout rounded in dorsal and lateral views (Figs. 1, 2); (4) upper eyelid bearing many low tubercles; upper eyelid about 43.5���85.7 % (x = 70.2) IOD; cranial crests absent; (5) dentigerous processes of the vomer evident, each processes bearing 2���8 teeth; (6) males with vocal slits, nuptial pads absent; (7) Finger I shorter than Finger II; discs on Finger I and II not expanded, discs on Finger III and IV slightly expanded (Fig. 1); (8) fingers lack lateral fringes; (9) ulnar tubercles absent; (10) heel and inner and outer edges of tarsus lacking tubercles; (11) inner metatarsal tubercle oval, about 1.5���2.5 �� the length of round outer metatarsal tubercle; (12) toes lack lateral fringes; webbing absent; Toe V slightly longer than Toe III; discs slightly expanded, disc on Toe IV slightly larger than those on outer fingers (Fig. 1); (13) in preservative, dorsum brown with darker markings, throat whitish cream to pale brown with two dark brown marks on center, venter whitish cream to pale brown, groin cream, posterior surfaces of thighs cream to pale brown (Fig. 2); (14) SVL in males 12.0��� 15.8 mm (x = 14.2 �� 1.013, n = 32), in females 17.0��� 21.7 mm (x = 19.0 �� 1.117, n = 33). Pristimantis bicantus is distinguished from most species in the genus Pristimantis by having a Toe V slightly longer than Toe III, a Toe V that reaches about to the level of the penultimate subarticular tubercle on Toe IV, and discs on fingers and toes not expanded or slightly expanded. Pristimantis bicantus is most likely to be confused with species of the myersi group (Lynch & Duellman 1997; Hedges et al. 2008), which contains the following species: Pristimantis festae (Peracca, 1904), P. floridus (Lynch & Duellman, 1997), P. gladiator (Lynch, 1976 b), P. hectus (Lynch & Burrowes, 1990), P. leoni (Lynch, 1976 b), P. myersi (Goin & Cochran, 1963), P. ocreatus (Lynch, 1981), P. pyrrhomerus (Lynch, 1976 b), P. repens (Lynch, 1984), P. scopaeus (Lynch, Ruiz-Carranza & Ardila-Robayo, 1996), and P. xeniolum (Lynch, 2001). The main characters distinguishing Pristimantis bicantus from P. f l o r i d u s, P. m y e r s i, and P. pyrrhomerus are the absence of tarsal tubercles (present in P. floridus, P. myersi, and P. pyrrhomerus); these species also occur in different biogreographic areas (Table 2). Additionally, Pristimantis bicantus differs from P. hectus and P. m y e r s i by lacking ulnar tubercles (present in P. hectus and P. myersi). Pristimantis bicantus is distinguished from P. leoni by lacking ulnar and tarsal tubercles (present in P. leoni), and bearing low tubercles on upper eyelid (subconical tubercles in P. leoni). Pristimantis bicantus differs from P. f e s t a e by having slightly expanded discs on outer fingers (fingers not expanded in P. f e s t a e), lacking lateral fringes on toes (present in P. f e s t a e), and having a pale cream venter in preservative (black venter usually with cream spots in P. f e s t a e). Pristimantis bicantus can be distinguished from P. ocreatus by possessing pads on fingers (pads absent on inner fingers in P. ocreatus), and having slightly expanded discs on toes and outer fingers (fingers and toes not expanded in P. ocreatus). Pristimantis bicantus differs from P. repens by lacking a single subconical tubercle on the upper eyelid (present in P. re p e n s; Lynch 1984), having, in life, a translucent venter with a gray-orange hue (olive-brown venter in P. repens; Lynch 1984); additionally, P. re p e n s is found in p��ramo and subp��ramo habitats (Lynch 1984), whereas P. bicantus inhabits cloud forests. Pristimantis bicantus is distinguished from P. scopaeus by having an evident tympanic membrane and tympanic annulus (not visible in P. scopaeus; Lynch et al. 1996). Pristimantis bicantus is distinguished from P. xeniolum by having dentigerous process of the vomer with teeth (teeth absent in P. xeniolum; Lynch 2001), lacking ulnar and tarsal tubercles (present in P. xeniolum; Lynch 2001), and, in males, having vocal slits (absent in P. xeniolum; Lynch 2001). Finally, P. bicantus is most likely to be confused with P. g l a d i a t o r. Differences between these two species are subtle, but not overlapping. Pristimantis bicantus has teeth on the vomers (teeth absent in P. gladiator), lacks ulnar and tarsal tubercles (low tubercles present in P. g l a d i a t o r), and, in life has a gray groin with or without reddish or salmon hue (groin black with red-orange spots in P. gladiator). Additionally, P. gladiator is found at elevations between 2270 and 2910 m (Lynch 1976 b; personal observation), whereas P. bicantus seems to be restricted to elevations between 2100���2300 m. Given the importance of the Andean mountains in promoting allopatric speciation (Lynch & Duellman 1997; Guayasamin et al. 2008; Bonaccorso 2009), it is relevant to note that only three species of the Pristimantis myersi group (i.e., P. bicantus, P. festae, P. gladiator) inhabit the same biogeographical region (Amazonian slope of the Andes; Table 2); therefore, comparisons between the new species and P. f e s t a e and P. g l a d i a t o r are, likely, more important when testing the validity of the new species. Description of holotype. Adult female (QCAZ 19024, SC 408) with head narrower than widest part of body; head longer than wide (head width 91.1 % head length); snout rounded in dorsal and lateral views, relatively short (snout-to-eye distance 18.6 % SVL), without papilla at tip (Fig. 1); in lateral view, canthus rostralis distinct; loreal region slightly concave; nostrils slightly protuberant, directed laterally; interorbital area flat, broader than upper eyelid (upper eyelid width 69.6 % interorbital distance); cranial crests absent; upper eyelid bearing several nonconical tubercles; tympanic membrane well-defined, with dark brown pigmentation on upper third; tympanic annulus distinct, round; supratympanic fold obscuring anterodorsal and posterodorsal edges of annulus (Fig. 1); tympanum diameter 47.6 % eye length; two nonconical postrictal tubercles. Choanae small, nearly elliptical, not concealed by palatal shelf of maxillary; dentigerous process of the vomer oblique, widely separated, posteromedial to choanae, each bearing 3���5 small teeth; tongue slightly longer than wide, granular, without notch in posterior border. Skin of head shagreen; dorsum shagreen, with scattered small tubercles, some of which are aligned over a W-shaped occipital mark, but not forming folds; upper flanks with numerous low warts; venter slightly areolate; discoidal fold absent; anal sheath absent. Forearm slender; radio-ulna length 24.5 % SVL; ulnar tubercles absent; ulnar fold absent; hand length longer than radio-ulna length (hand length 26.1 % SVL); fingers without lateral fringes; finger lengths I Measurements of holotype (mm).��� Adult female (QCAZ 19024); SVL = 18.8; tibia length = 10.2; foot length = 9.0; head length = 7.9; head width = 7.2; interorbital distance = 2.3; upper eyelid width = 1.6; internarial distance = 2.4; eye-to-nostril distance = 1.8; snout-to-eye distance = 3.5; eye diameter = 2.1; tympanum diameter = 1.0; eye-to-tympanum distance = 0.9; radio-ulna length = 4.6; hand length = 4.9; and Finger-I length = 2.9. Variation in external morphology. Meristic variation of the type series is presented in Table 4. There is conspicuous variation in the number of teeth on the dentigerous process of the vomer (2���8 teeth), usually larger individuals have more teeth. Tympanum in males larger than in females (tympanum in males 6.9��� 10.2 % SVL, tympanum in females 4.3���6.2 % SVL). Snout with small papilla at tip (females, QCAZ 10103, 16209, 19029). Some individuals (QCAZ 16203, 16209, 19031, 19040���2, 19045, 19048, 19051, 19053, 19057) present a series of low tubercles that form low occipital and dorsolateral folds (Fig. 2). Color in life. The following description includes the color variation found in Pristimantis bicantus. Dorsum light to dark brown usually with an indistinct dark interorbital bar and with or without dark chevron markings; some individuals with light dorsolateral line and some with pale snout; dorsal surfaces of legs with contrasting dark and light bars; distinct dark line extending from posterior edge of eye, through top half of tympanum, to above insertion of arm; flanks light brown or gray with or without diagonal dark bars extending towards groin; uniform dark brown patch with lighter colored border on posterior surface of thighs below cloacal opening; groin and hidden surfaces of thighs gray with or without a reddish or salmon hue; venter translucent with a light gray-orange hue and small white and black speckles, especially laterally and anteriorly; iris pale yellow with dark red medial horizontal bar and orange ring around pupil (Plate 3). Color variation (in ethanol).��� Dorsally, anterior half of head cream (females, QCAZ 19042, 19051; males, QCAZ 19026). Dorsum grayish cream (females, QCAZ 16201, 19039; males, QCAZ 18981, 18983��� 84, 19019, 19027, 19035, 19038, 19044), pale brown (female QCAZ 19005), or uniform pale brown (female, QCAZ 19056; males, QCAZ 18994, 18997). Interorbital bar, W-shaped mark, and chevron absent (females, QCAZ 16186, 19056; males, QCAZ 18994, 18997). Dorsolateral stripes, external border of upper eyelid, and canthal stripe cream (female, QCAZ 10102, 19029, 19049, 19051; males, QCAZ 18980, 18982, 18996, 19004, 19031, 19036, 19052). Dorsolateral stripes dark gray (males, QCAZ 18994, 18997). Flanks whitish cream (females, QCAZ 16209, 19049; males, QCAZ 18981, 19030, 19032, 19035, 19038, 19044), grayish cream (female, QCAZ 19039; males, QCAZ 18980, 18983, 19052, 18984), or brown (female, QCAZ 19001). Groin and flanks brown with cream flecks (female, QCAZ 19001). Posterior surfaces of thighs creamy gray (female, QCAZ 16201; males, QCAZ 19019, 19030���02, 19035, PUCE 330, 334), or brown with cream flecks (females, PUCE 335, QCAZ 19001). Throat, venter, and groin whitish cream (females, PUCE 329, 344, QCAZ 16186, 16209, 19003, 19005���06, 19029, 19033, 19046, 19048��� 49; males, QCAZ 18981, 18984, 18986, 19004, 19019, 19026, 19030 ���02, 19038, 19044), creamy gray (females, QCAZ 16201, 19039, 19042; males, QCAZ 18980, 19052), whitish gray (females, QCAZ 19050 ��� 51), pale brown (female, QCAZ 18978), or brown (female, QCAZ 10102). Throat with no dark brown marks (females, QCAZ 10102, 19003, 19048, 19029, 19050; males, QCAZ 18992 ���94, 19004, 19019, 19025���26, 19030, 19036, 19038), brown with darker marks and cream flecks (males, QCAZ 18982, 18996���97, 19034), or whitish cream (male, QCAZ 18983). Venter gray (male, QCAZ 18983). Palms and soles gray (females, QCAZ 19001, 19029, 19051; male, QCAZ 19001). Etymology. The specific name bicantus is derived from the Latin bi-, meaning ���two��� and cantus, meaning ���song��� in reference to the species��� call, which has two distinct call types. Distribution. Pristimantis bicantus is known only from the cloud forests surrounding Yanayacu Biological Station (0�� 36 ��S, 77 �� 53 ��W) at elevations between 2100 and 2300 m (Fig. 3). Ecology. Pristimantis bicantus is one of the two most abundant frog species at Yanayacu (Pristimantis cf. petersi is the other most abundant frog; Table 1). Ninety-four individuals were found at night during approximately two years of inventory work (2000���2002), of which 65 were adults, 23 were juveniles, one was a subadult, and stage was not determined for the remaining five. Most were found in primary forest, although 31 % were found in secondary forest, suggesting this species is more tolerant of disturbance than most other Pristimantis at Yanayacu. Pristimantis bicantus used low leaves 0���80 cm (x = 32 cm) above the ground as substrate at night (Fig. 4). Of the eight P. bicantus females dissected to count and measure ovarian ova, the number of large ova was 13���20 (x = 16.3), each with a diameter of 1.1���2.6 mm (x = 2.1 mm). Some females also had a few to several very small ova. Museum number mean, standard deviation, and sampling size (n). Females Males SVL 17.0��� 21.7 11.8���15.8 18.9 �� 1.1 14.2 �� 1.0 n = 36 n = 32 Foot 8.0��� 9.5 5.5���6.8 8.7 �� 0.3 6.4 �� 0.4 n = 26 n = 12 Head length 7.2���8.6 5.1���6.5 8 �� 0.3 6.0 �� 0.4 n = 26 n = 12 Head width 6.2���7.8 5.0��� 6.2 7.2 �� 0.3 5.3 �� 0.3 n = 26 n = 12 Interorbital distance 2.0��� 2.6 1.7���2.2 2.3 �� 0.1 1.9 �� 0.1 n = 36 n = 32 Upper eyelid width 1.0��� 1.9 1.0��� 1.7 1.6 �� 0.2 1.3 �� 0.2 n = 36 n = 32 Eye-to-nostril distance 1.5���2.2 1.3���1.7 1.9 �� 0.1 1.4 �� 0.1 n = 26 n = 12 Snout-to-eye distance 3.0��� 3.9 2.2���2.9 3.3 �� 0.2 2.5 �� 0.2 n = 26 n = 12 Eye diameter 2.1���2.8 1.7���2.5 2.4 �� 0.2 2.0 �� 0.3 n = 26 n = 12 Tympanum 0.8���1.4 0.9���1.5 1 �� 0.2 1.2 �� 0.2 n = 36 n = 32 Radioulna length 4.4���5.3 2.9���, Published as part of Guayasamin, Juan M. & Funk, Chris, 2009, The amphibian community at Yanayacu Biological Station, Ecuador, with a comparison of vertical microhabitat use among Pristimantis species and the description of a new species of the Pristimantis myersi group, pp. 41-66 in Zootaxa 2220 on pages 49-56, DOI: 10.5281/zenodo.190060, {"references":["Lynch, J. D. & Duellman, W. E. (1997) Frogs of the genus Eleutherodactylus in western Ecuador. Systematics, Ecology, and biogeography. Special Publications of the Natural History Museum, University of Kansas, Lawrence, 23, 1 - 236.","Hedges, S. B., Duellman, W. E. & Heinicke, P. (2008) New world direct-developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182.","Peracca, M. G. (1904) Rettili ed Amfibii in viaggio del Dr. Enrico Festa nell'Ecuador e regioni vicine. Bolletino dei Musei di Zoologiaed Anatomia Comparata della Universita di Torino, xix, 1 - 41.","Lynch, J. D. (1976 b) Three new leptodactylid frogs (genus Eleutherodactylus) from the Andean slopes of Colombia and Ecuador. Herpetologica, 32, 310 - 317.","Lynch, J. D. & Burrowes, P. A. (1990) The frogs of the genus Eleutherodactylus (Family Leptodactylidae) at the La Planada Reserve in southwestern Colombia with descriptions of eight new species. Occasional Papers of the Museum of Natural History, University of Kansas, Lawrence, 136, 1 - 31.","Goin, C. J. & Cochran, D. M. (1963) Two new genera of leptodactylid frogs from Colombia. Proceedings of the California Academy of Sciences, 4, 31, 499 - 505.","Lynch, J. D. (1981) Leptodactylid frogs of the genus Eleutherodactylus in the Andes of northern Ecuador and adjacent Colombia. Miscellaneous Publications, Museum of Natural History, University of Kansas, Lawrence, 72, 1 - 46.","Lynch, J. D. (1984) A new species of Eleutherodactylus (Amphibia: Anura: Leptodactylidae) from southern andean Colombia. Herpetologica, 40, 234 - 237.","Lynch, J. D., Ruiz-Carranza, P. & Ardila-Robayo, M. (1996) Three new species of the Eleutherodactylus (Amphibia: Leptodactylidae) from high elevations of the Cordillera Central of Colombia Caldasia, 18, 329 - 342.","Lynch, J. D. (2001) A small amphibian fauna from a previously unexplored paramo of the Cordillera Occidental in western Colombia. Journal of Herpetology, 35, 226 - 231.","Guayasamin, J. M., Castroviejo-Fisher, S., Ayarzaguena, J., Trueb, L. & Vila, C. (2008) Phylogenetic relationships of glassfrogs (Centrolenidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution, 48, 574 - 595.","Bonaccorso, E. (2009) Historical biogeography and speciation in the Neotropical highlands: molecular phylogenetics of the jay genus Cyanolyca. Molecular Phylogenetics and Evolution, 50, 618 - 632.","Funk, W. C., Almeida-Reinoso, D., Nogales-Sornosa, F. & Bustamante, M. R. (2003) Monitoring population trends of Eleutherodactylus frogs. Journal of Herpetology, 37, 245 - 256."]}

Family: Dendrobatidae The family Dendrobatidae is distributed in the Neotropics and contains 267 species (AmphibiaWeb 2009; including species of Aromobatidae and Dendrobatidae sensu Grant et al. 2006). Unlike most anurans, poison dart frogs are diurnal and often times show colorful, aposematic coloration. Also, dendrobatids present elaborated reproductive modes, with different types of parental care (Zimmerman, 1990). In YBS, only one species of dendrobatid has been found, Hyloxalus pulchellus. We follow the taxonomy proposed by Grant et al. (2006), as modified by Santos et al. (2009). Hyloxalus pulchellus (Plate 2). Described by Jiménez de la Espada (1871). This species is characterized by having a dark brown dorsum, thin cream lateral stripe, dark brown to black flanks, and a bluish-white venter with grayish-black reticulation. In adult males, SVL 17.3–21.5 mm (n = 44); 19.6–24.4 mm (n = 16) in adult females (Coloma 1995). Remarks. Hyloxalus pulchellus exhibits morphological and color variation along its distribution —inter- Andean páramos of southern Colombia and northern Ecuador, and on the eastern slopes of the Andes from southern Colombia to the headwaters of the Río Pastaza in Ecuador, at elevation between 1590 and 2970 m) — and it has been suggested that it might represent a complex of species (Coloma 1995).

Family: Strabomantidae This family is restricted to tropical and subtropical South America (Hedges et al. 2008) and has a recognized diversity of 556 species (AmphibiaWeb 2009). All strabomantids seem to have direct development, a trait that might have facilitated their diversification in moist areas where standing water is rare (e.g., cloud forests). Most species in this family lack webbing between fingers and toes. Strabomantids represent the most diverse clade in YBS, with 6 described and 3 undescribed species. We follow the taxonomy proposed by Heinicke et al. (2007) and Hedges et al. (2008). Pristimantis eriphus (Plate 3). Described by Lynch and Duellman (1980). This species is distinguished from other anurans by having many small conical tubercles on dorsal skin, one prominent tubercle on the upper eyelid, conical ulnar and tarsal tubercles, and a conical tubercle on heel. In life, Pristimantis eriphus has a pale green dorsum with dark olive to green-brown markings; its flanks and hidden surfaces of limbs have a black and white pattern; the venter is white to greenish-white with dark brown to black reticulations; the iris is orange-red to reddish copper. At YBS, SVL in adult males is 16.3–21.4 mm (n = 16); in adult females, SVL 18.6–33.2 mm (n = 16). Pristimantis gladiator (Plate 4). Described by Lynch (1976 b). In life, Pristimantis gladiator has a brown to orange-brown dorsum with dark brown markings, and a black groin with orange to red spots. Other diagnostic traits of P. gladiator include the presence of low ulnar and tarsal tubercles and low, non-pungent tubercles on the upper eyelid. In adult males, SVL 14.9–16.9 mm (n = 5); adult females unknown (updated from Lynch 1976 b). None of the other frogs at YBS have a black groin with orange-red spots. This small species has been found by day under rocks and logs (Lynch 1976 b). Pristimantis inusitatus (Plate 4). Described by Lynch and Duellman (1980). This species is distinguished from other frogs mainly by its dorsal color pattern (see below) and by having conical ulnar and tarsal tubercles, a conical tubercle on heel, and a conical tubercle on the upper eyelid. In life, Pristimantis inusitatus has a sexually dimorphic coloration. Females have a green dorsum with yellow to yellow-green tubercles, a pale lemon yellow to yellow groin, white throat with green markings, white venter, and yellow-cream to yellow iris with black reticulations. Males differ by having a brown-green to pink-brown dorsum with dark brown marks, clearly banded limbs, a pale yellow throat with small white spots, and a white venter (translucent on its posterior portion). At YBS, SVL in adult males is 13.1–17.5 mm (n = 15); in adult females, SVL 22.2 –26.0 mm (n = 10). Pristimantis leucopus (Plate 5). Described by Lynch (1976 c). A species diagnosed by having an upper eyelid with few small flat tubercles or lacking tubercles, low and inconspicuous ulnar and tarsal tubercles, and by lacking tubercles on the dorsum. In life, this species has a green to brown dorsum with dark brown marks, flanks mostly dark gray with greenish-brown mottling, and throat, chest, and belly reddish-orange with brown mottling that is most conspicuous on the throat. The groin and ventral surfaces of legs are dark gray to black. Pristimantis leucopus is one the largest species found at YBS; SVL in adult males is 29.7–37.8 mm; in adult females, SVL 39.2 –44.0 mm (Lynch & Duellman 1980; this work). Pristimantis cf. petersi (Plate 5). Described by Lynch and Duellman (1980). This species is characterized by having smooth dorsal skin, a papilla at the tip of the snout, and one conical tubercle on the upper eyelid. In life, Pristimantis petersi usually has a pale green to dull green dorsum with no distinct markings (Lynch & Duellman 1980), but individuals found at YBS have a brown to orange-brown dorsum with dark brown markings. At YBS, SVL in adult males is 15.7–21.2 mm (n = 10); in adult females, SVL 24.0– 27.5 mm (n = 5). Remarks. Because of the differences in color pattern between the usual Pristimantis petersi and the population found at YBS, we consider the identification as tentative. Additionally, individuals (especially females) found at YBS are larger than those reported in the literature (in 35 males, SVL = 14.5–19.9 mm; in 8 females, SVL = 20.3–23.1 mm; Lynch & Duellman 1980). In Funk et al. (2003), Pristimantis cf. petersi was referred to as Eleutherodactylus sp. 2. Pristimantis w-nigrum (Plate 5). Described by Boettger (1892). This frog is easily recognized by having, in life, a color pattern of yellow with black markings on the groin, anterior and posterior surfaces of thighs, and concealed shank. Other distinctive traits include having a light brown dorsum with brown markings, a prominent tympanum, and a first finger that is longer than the second. At YBS, SVL in adult males is 31.9– 40.2 mm (n = 5); no adult females have been found. Lynch and Duellman (1980) report the following body size for the species: in adult males, SVL = 29.3–46.8 mm (n = 32); in adult females, SVL = 44.4–56.6 mm (n = 15). Remarks. Pristimantis w-nigrum is unusual among Andean amphibians by having a broad distribution that includes the Pacific and Amazonian slopes of the Andes, as well as the high Andes of Colombia and Ecuador (800–3200 m; Lynch & Duellman 1997). Given the importance of the Andes in the speciation of amphibians (Lynch & Duellman 1997; Guayasamin et al. 2008), it is likely that P. w-nigrum actually represents a species complex. Caecilia orientalis (Plate 5). — Described by Taylor (1968). Family Caeciliidae. Caecilians are subterranean or aquatic amphibians found throughout much of the tropics (Duellman & Trueb 1994). At YBS, only one species of caecilian has been reported, Caecilia orientalis (Funk et al. 2004), which is easily identified by having an elongated and limbless body with a blue-gray to lavender dorsal coloration. Funk et al. (2004) described the egg clutch and nest site of C. orientalis, confirming that this caecilian is oviparous. Caecilia orientalis is the main prey item of the Barred Hawk, Leucopternis princeps (Greeney et al. 2008). At YBS, SVL in adult males is 320–356 mm (n = 2); in females, SVL = 348–458 mm (n = 2). FIGURE 4. Microhabitat use of sympatric species of Pristimantis at Yanayacu Biological Station. Box plots of the vertical distance above the ground where frogs were encountered at night. Separate box plots are shown for adults and juveniles for species in which at least 10 individuals of each stage were found. Grey = all individuals; black = adults; and white = juveniles. The boundaries of boxes indicate the 25 th and 75 th percentiles; the line within the box is the median; error bars are the 10 th and 90 th percentiles; and points below and above error bars are outliers. Sample sizes (numbers of frogs) are shown above plots in parentheses. Species grouped with a horizontal dashed line and letter did not have significantly different height means (for all individuals; Tukey-Kramer method, P > 0.05).

Family: Bufonidae This group of toads contains some of the most familiar anurans, including the cane toad (Rhinella marina; previously Bufo marinus) and the jambato (Atelopus ignescens). Most of them have a warty appearance and are diurnal, terrestrial, and have aquatic larvae (e.g., Rhinella, Atelopus), but others are nocturnal, arboreal, and undergo direct development (e.g., Osornophryne). Bufonids are found natively in all continents, except Australia (where it has been introduced) and Antarctica. The family Bufonidae contains 538 species (AmphibiaWeb 2009). The taxonomy of amphibians has undergone several recent changes and there is debate on which names should be used. For bufonids, we follow the general proposal by Frost et al. (2006), as modified by Chaparro et al. (2007). In YBS, there are two species of bufonids, Osornophryne guacamayo and Rhinella margaritifera (previously Bufo margaritifer). Osornophryne guacamayo (Plate 1). Described by Hoogmoed (1987). This species can be easily recognized by having a blackish dorsal coloration with, usually, two yellow-gray dorsolateral lines, which are not clearly differentiated from the rest of the dorsum; yellow venter with dark brown to black spots; dorsal skin of body and limbs with numerous tubercles of different sizes; limbs long and slender; fingers connected by a thick web; feet heavily webbed with Toe IV and V much longer than Toes I, II, and III; and by lacking parotoid glands and tympanum (Hoogmoed 1987). At YBS, SVL in adult females is 36.1���39.3 mm (n = 3) and 22.7 mm in one adult male. Osornophryne guacamayo has been found on leaves during the night (Table 1). Although most bufonids reproduce in water, O. guacamayo places its eggs on the ground, where they undergo direct development into froglets. At YBS, the only species that is likely to be confused with O. guacamayo is Rhinella margaritifera, which has a visible tympanum, parotoid glands, dorsolateral row of red tubercles (traits absent in O. guacamayo), and lacks conspicuous webbing among fingers (present in O. guacamayo). Additionally, in R. margaritifera, only Toe IV is elongated, whereas in O. guacamayo Toes IV and V are conspicuously longer that the rest of the toes. Rhinella margaritifera (Plate 1). Described by Laurenti (1768). A moderate-sized toad distinguished by having dorsal skin scattered with tubercles, and a row of red conical tubercles that begins on the posterior edge of the parotoid gland and ends on the groin. The fingers lack webbing, whereas the toes are about onehalf webbed. This is a terrestrial and diurnal species that has a dorsal coloration that mimics leaf litter. At YBS, SVL in adult females is 57.4���68.7 mm (n = 4) and 40.9���46.1 mm (n = 5) in adult males. Within YBS, Rhinella margaritifera can only be confused with Osornophryne guacamayo (see previous species account). Remarks: One of the oldest problem in the taxonomy of South American amphibians is the Rhinella margaritifera species complex. Hoogmoed (1986, 1989) recognized several species within this complex (previously referred to as the Bufo typhonius complex), but he did not formally recognize most of them. More recently, Fouquet et al. (2007) suggested that R. margaritifera might be composed of as many as 11 species. Herein, we use the name R. margaritifera realizing that it represents a species complex until the systematics of this group of toads is resolved., Published as part of Guayasamin, Juan M. & Funk, Chris, 2009, The amphibian community at Yanayacu Biological Station, Ecuador, with a comparison of vertical microhabitat use among Pristimantis species and the description of a new species of the Pristimantis myersi group, pp. 41-66 in Zootaxa 2220 on page 43, DOI: 10.5281/zenodo.190060, {"references":["Chaparro, J. C., Pramuk, J. & Gluesenkamp, A. (2007) A new species of arboreal Rhinella (Anura: Bufonidae) from cloud forest of southeastern Peru. Herpetologica, 63, 203 - 212.","Hoogmoed, M. S. (1987) New Osornophryne (Amphibia: Anura Bufonidae) from the Atlantic versant of the Andes in Ecuador. Zoologische Medelingen, 61, 209 - 242.","Laurenti, J. N. (1768) Specimen medicum, exhibens synopsin Reptilium emendatum cum experimentis circa venena et antidota Reptilium Austriacorum. Wien, Trattnern, 214 pp.","Hoogmoed, M. S. (1986) Biosystematic studies of the Bufo \" typhonius \" group: a preliminary report. In Z. Rocek (Ed), Studies in Herpetology. Charles University, Prague, pp 147 - 150.","Hoogmoed, M. S. (1989) On the identity of some toads of the genus Bufo from Ecuador, with additional remarks of Andinophryne colomai Hoogmoed, 1985 (Amphibia: Anura: Bufonidae). Zoologische Verhandelingen, Rijksmuseum van Natuurlijke te Leiden, 250, 1 - 32.","Fouquet, A., Vences, M., Salducci, M., Meyer, A., Marty, C., Blanc, M. & Gilles, A. (2007) Revealing cryptic diversity using molecular phylogenetics and phylogeography in frogs of the Scinax ruber and Rhinella margaritifera species groups. Molecular phylogenetics and Evolution, 43, 567 - 582."]}

Centrolene gorzulae (Ayarzag��ena, 1992) Centrolenella gorzulae Ayarzag��ena 1992: 19. Centrolene gorzulai Duellman 1993: 35. Centrolenella auyantepuiana Se��aris and Ayarzag��ena 1995 ��� 1993 ���: 122. Centrolene gorzulae Ruiz-Carranza and Lynch 1995: 2. Hyalinobatrachium auyantepuiana Ayarzag��ena and Se��aris ��� 1996 ��� 1997: 13. Cochranella auyantepuiana Myers and Donnelly 1997: 16. Cochranella auyantepuiana Duellman 1999: 300. Centrolene papillahallicum Noonan and Harvey 2000: 295. Centrolene lema Duellman and Se��aris 2003: 247, new synonym. Centrolene gorzulai Myers & Donnelly 2008: 29. Holotype. MHNLS 11221 (adult male), Venezuela, Estado Bol��var: Central sector of Auyan-tepui (05�� 56 ��� N, 62 �� 34 ��� W; 1850 m). Diagnosis. This species is placed in the genus Centrolene because adult males have humeral spines (Ruiz- Carranza & Lynch 1991; Duellman & Se��aris 2003). It differs from all other species of Centrolene by having a trilobate liver covered by a white peritoneum and a mostly transparent parietal peritoneum, with a small white anterior portion (the ���bib-like white patch��� of Duellman & Se��aris 2003), which is only detectable in dissected specimens. Characterization. (1) dentigerous process on vomer and vomerine teeth absent; (2) snout subtruncate to truncate in dorsal view, truncate to slightly sloping in lateral profile; (3) tympanum distinct, small, oriented dorsolaterally with a slight posterior inclination; (4) dorsal surfaces shagreened; males with small spicules visible under magnification; (5) ventral skin strongly granular, cloacal ornamentation consisting of small enameled tubercles lateral to vent and enlarged paired round tubercles below vent; (6) parietal peritoneum mostly transparent, a small anterior portion white (the ���bib-like white patch��� of Duellman & Se��aris 2003), only detectable in dissected specimens; pericardial peritoneum white; hepatic and visceral peritonea white, urinary bladder transparent; (7) liver trilobate; (8) humeral spine in adult males present; (9) finger webbing II 2 -��� 3 III 2 +���(2 -��� 2) IV; (10) toe webbing I (1 ��� 1 ��)���(2 ��� 2 +) II 1���2 + III 1 ���(2 ��� 2 +) IV (2 ��� 2 +)��� 1 V; (11) fringe on postaxial edge of Finger IV present, enameled, metacarpal fold present, weakly enameled, ulnar fold absent; fringe on postaxial edge of Toe V present, enameled, metatarsal fold present, weakly enameled, tarsal fold usually absent or very weak; (12) nuptial excrescences finely granular, white, consisting of a dense group of glands and situated in the medial, dorsolateral internal side of Finger I (Type-V); prepollex slightly enlarged; prepollical spine projecting (spine not exposed); (13) when appressed, Finger II = Finger I; (14) diameter of eye about 2 X width of disc on Finger III; (15) in life, dorsum dark green with scattered paler flecks; (16) in preservative, dorsum lavender entirely covered with minute melanophores; (17) in life, iris metallic copper with dark brown reticulations; an incomplete pale yellow ring around pupil; (18) distribution of melanophores on digits variable, but always more numerous on Fingers III���IV than I and II; Toes IV���V with many melanophores, rarely present on Toes I���III, if present, few and restricted to the two distal phalanges; in life, hands and feet bluish green to green, tips of fingers and toes yellowish green; (19) males call from the upper side of leaves; note single and pulsed (3���7 pulses per call), advertisement call of 0.02��� 0.05 s duration, dominant frequency of 4416.97���5157.48 Hz; (20) combat behavior unknown; (21) clutches deposited on vegetation overhanging streams, in moss or between two leaves, clutch size 15��� 22 eggs (n = 4); none of the clutches observed were guarded; (22) tadpole unknown; (23) SVL in males 19.2���22.5 mm (n = 11); in females 20.9 ���22.0 mm (n = 4). Natural history and distribution. The ecology of Centrolene gorzulae was recently discussed by Kok and Castroviejo-Fisher (2008), on the basis of specimens from Kaieteur National Park, Guyana. The species was recently discovered at a new locality, the southeast slope of Mount Maringma on the border of Guyana and Brazil (ca. 1400 m elevation), and additional observations were made on clutch deposition. Four clutches were collected between 23���24 November 2007; three of them were found among moss on overhanging branches 1.5 ���2.0 m above water (Fig. 5), the fourth clutch was found deposited between two leaves (attached to both), about 1.5 m above water. Many males were calling at night in the vicinity of the clutch deposition sites, but no male-male interaction was observed. Clutches contained 15��� 22 eggs at various stages of embryogenesis, and no guarding males were detected. One female (IRSNB 14327, 20.9 mm SVL), collected on 22 November 2007, contained 13 enlarged, partly pigmented, ovarian eggs, the largest with a diameter of 2.0 mm. Centrolene gorzulae was thought to be endemic to Auyan-tepui (Ayarzag��ena 1992; Se��aris & Ayarzag��ena 2005; Myers & Donnelly 2008). Kok and Castroviejo-Fisher (2008) extended its distribution to Guyana. Here we report C. gorzulae from three localities in Venezuela (Auyan-tepui, Atapare, and Sierra de Lema) and Guyana (Peters Mountain, Kaieteur National Park, and Mount Maringma) (Fig. 6). This species probably occurs in adjacent areas of Brazil. Centrolene gorzulae has a broad altitudinal distribution (~ 450���1850 m) and has been found only on vegetation overhanging small streams., Published as part of Castroviejo-Fisher, Santiago, Guayasamin, Juan M. & Kok, Philippe J. R., 2009, Species status of Centrolene lema Duellman and Se��aris, 2003 (Amphibia: Centrolenidae) revealed by Integrative Taxonomy, pp. 16-28 in Zootaxa 1980 on pages 19-24, DOI: 10.5281/zenodo.185278, {"references":["Ayarzaguena, J. (1992) Los Centrolenidos de la Guayana Venezolana. Publicaciones de la Asociacion de Amigos de Donana, 1, 1 - 48.","Duellman, W. E. (1993) Amphibian species of the world: additions and corrections. University of Kansas Special Publications, Museum of Natural History, 21, 1 - 372.","Ruiz-Carranza, P. M. & Lynch, J. D. (1995) Ranas Centrolenidae de Colombia VIII. Cuatro nuevas especies de Centrolene de la Cordillera Central. Lozania, 65, 1 - 16.","Ayarzaguena, J. & Senaris, J. C. (\" 1996 \" [1997]) Dos nuevas especies de Cochranella (Anura; Centrolenidae) para Venezuela. Publicaciones de la Asociacion de los Amigos de Donana, 8, 1 - 16.","Myers, C. W. & Donnelly, M. A. (1997) A Tepui Herpetofauna on a Granitic Mountain (Tamacuari) in the Borderland Between Venezuela and Brazil: Report from the Phipps Tapirapeco Expedition. American Museum Novitates, 3213, 1 - 71.","Duellman, W. E. (1999) Distribution patterns of amphibians in South America. In: Duellman, W. E. (Ed.), Patterns of Distribution of Amphibians. A Global Perspective. The Johns Hopkins University Press, Baltimore and London, pp. 255 - 328.","Noonan, B. P. & Harvey, M. B. (2000) A new species of glass frog (Anura: Centrolenidae) from the Highlands of Guyana. Herpetologica, 56, 294 - 302.","Duellman, W. E. & Senaris, J. C. (2003) A new species of glass frog (Anura: Centrolenidae) from the Venezuelan Guayana. Herpetologica, 59, 247 - 252.","Myers, C. W. & Donnelly, M. A. (2008) The summit herpetofauna of Auyantepui, Venezuela: Report from the Robert G. Goelet American Museum-Terramar Expedition. Bulletin of the American Museum of Natural History, 308, 1 - 147.","Ruiz-Carranza, P. M. & Lynch, J. D. (1991) Ranas Centrolenidae de Colombia I. Propuesta de una nueva clasificacion generica. Lozania, 57, 1 - 30.","Kok, P. J. R. & Castroviejo-Fisher, S. (2008) Glassfrogs (Anura: Centrolenidae) of Kaieteur National Park, Guyana, with notes on the distribution and taxonomy of some species of the family in the Guiana Shield. Zootaxa, 1680, 25 - 53.","Senaris, J. C. & Ayarzaguena, J. (2005) Revision taxonomica de la Familia Centrolenidae (Amphibia; Anura) de Venezuela. Publicaciones del Comite Espanol del Programa Hombre y Biosfera - Red IberoMaB de la UNESCO, Sevilla, 337 pp."]}

Guayasamin, Juan M., Funk, Chris (2009): The amphibian community at Yanayacu Biological Station, Ecuador, with a comparison of vertical microhabitat use among Pristimantis species and the description of a new species of the Pristimantis myersi group. Zootaxa 2220: 41-66, DOI: 10.5281/zenodo.190060

Osornophryne puruanta new species Holotype.��� QCAZ 11471 (Fig. 1), an adult female, obtained near Laguna de Puruanta (00�� 13 ' N, 77 �� 57 ' W, 3000 m.a.s.l.), Cordillera de Pimampiro, Provincia de Imbabura, Ecuador, by A. G. Gluesenkamp and D. A. Gluesenkamp on 17 December 1997. Paratypes.��� QCAZ 7684 ��� 85, adult females, obtained near Laguna de Puruanta (00�� 13 ' N, 77 �� 57 ' W, 3500 m.a.s.l.), Cordillera de Pimampiro, Provincia de Imbabura, Ecuador, by A. Vallejo, J. W. Izquierdo, and D. Almeida on 16 November 1996; QCAZ 13271, adult female, obtained near Laguna de San Marcos (00��07' 35 " N, 77 �� 55 ' 50 " W, 3400 m.a.s.l.), on the border between Provincia de Sucumb��os and Provincia de Pichincha, Ecuador, by S. R. Ron and L. A. Coloma on 29 June 1999; QCAZ 13320, adult female, obtained near Laguna de San Marcos, by J. M. Guayasamin on 21 July 1999; EPN 7081 ��� 83, adult females, obtained near Laguna de San Marcos, by A. Almend��riz and V. Corte on 20 September 1999. Diagnosis.���The following traits characterize Osornophryne puruanta: (1) females with large body size (SVL 40.5���47.1, x = 43.4 �� 2.3, n = 8), males unknown; (2) head rounded in dorsal view, rounded to slightlypointed in lateral view (Fig. 2); (3) tip of snout usually bears papilla; (4) skin on dorsum and flanks relatively smooth, interspersed with numerous glandular pustules some forming ridges, and lacking conical tubercules (Fig. 2); (5) dorsum with discontinuous dorsolateral, parasagittal, paravertebral, and sacral ridges; (6) dorsum pale brown to reddish-brown in life (pale brown in preservative); (7) venter brown with yellowish-brown spots in life (dark gray with pale yellow pustules or pale yellow with grayish-brown reticulations in preservative); and (8) six presacral vertebrae (atlas not fused to Presacral Vertebra II). Comparison with congeneric species.��� Osornophryne puruanta can be distinguished from other members of the genus by its large body size (adult female SVL> 40 mm; O. puruanta differs from O. antisana in having six discrete presacral vertebrae (five in O. antisana), wrinkled skin in the occipital area (smooth and flat in O. antisana), and a relatively more rounded snout (pointed in O. antisana). Osornophryne puruanta can be distinguished from O. bufoniformis by having a snout that is relatively more rounded in dorsal view (with a sharp angle at level of nostril in O. bufoniformis, Fig. 3), fleshy and wrinkled palmar and plantar surfaces (tuberculate in O. bufoniformis, Fig. 3), and a relatively narrower urostyle (Fig. 4; also, see Ruiz-Carranza & Hern��ndez-Camacho 1976:Fig. 8). Osornophryne puruanta can be distinguished from O. guacamayo by the presence of six discrete presacral vertebrae (five in O. guacamayo), a pustular upper eyelid (strongly tuberculate in O. guacamayo), elongate Toe IV (Toes IV and V elongate in O. guacamayo), and a mostly brown dorsum lacking dorsolateral lines (blackish brown, typically with two oblique yellow dorsolateral lines in adult female O. guacamayo). Osornophryne puruanta differs from O. percrassa by a more rounded snout in lateral view (truncate in O. percrassa), dorsum smooth with interspersed pustules (covered with small and large pustules in O. percrassa), a ventral color pattern composed of small, yellowish cream pustules (composed of large, irregular yellowish gray blotches in O. percrassa), and a pale brown dorsum in preservative (black in O. percrassa). Osornophryne puruanta differs from O. sumacoensis in having an atlas that is discrete and not fused to Presacral II (fused in O. sumacoensis), yellowish-brown or reddish-brown dorsum in life (dark brown or black in O. sumacoensis) and a venter that is brown with yellowish-cream pustules in life (black or blue with black spots in O. sumacoensis). Osornophryne puruanta differs from O. talipes by having a snout that is relatively more rounded in dorsal and lateral views (highly acuminate in O. talipes), and a head that is wider than long (about as wide as long in O. talipes). Description of holotype.��� Head wider than long (HL 91.85 % HW). Snout with papilla at tip, rounded in dorsal and lateral views. Tip of snout extending slightly beyond anterior margin of upper jaw. Canthus rostralis prominent, forming a crest angled medially from anterior margin of orbit to narial region in dorsal view, convex in lateral view, widened around naris. Nostrils directed laterally. Interorbital region smooth, depressed. Upper eyelid weakly rugose with small, flat pustules and pustular margin forming a glandular ridge continuous with canthus rostralis. Infraorbital region smooth with few scattered, inconspicuous glandular pustules. Tympanum absent. Occipital region bearing large, glandular warts. Loreal region smooth, slightly concave. Lips smooth, prominent. Tongue oval, pigmented at base. Choanae small, round, vomerine teeth absent. Skin on dorsum lacking conical tubercules, but interspersed with glandular pustules of different sizes. Some pustules forming conspicuous non-continuous ridges. Postorbital ridge oblique, extending from posterior margin of eyelid to paravertebral ridge. Glandular ridge along iliac crest, continuous with paravertebral ridge. Dorsolateral ridge extending from posterolateral margin of eyelid to paravertebral ridge at superior margin of cloaca. Flank of abdomen rugose, bearing numerous small warts interspersed with few large, pustular warts. Venter slightly rugose with numerous clusters of small, round pustules. Gular fold pronounced. Cloaca directed posteroventrally, positioned on inconspicuous cloacal tube. Supracloacal wart azygous. Limbs slender. Skin on dorsal surfaces of limbs warty. Dorsal skin of forelimb with many small, tuberculate warts. Dorsal surface of hind limb with larger, pustular warts. Ventral surfaces of thighs covered with tuberculate warts; shanks with low, rounded, pustular warts. Palmar and plantar surfaces bearing numerous tubercles. Tips of fingers and toes free of webbing, fleshy pad on ventral surface of each tip. Length of fingers: III> IV> II> I; length of toes: IV> V> III> II> I. Inguinal fat bodies absent. Color of the holotype in preservative.��� Dorsum yellowish pale brown. Venter dark brown with yellowish clusters of pustules. Cloacal area black. Palmar and plantar surfaces reddish-brown. Color of the holotype in life.��� Dorsum yellowish-brown. Iris dark brown with few golden flecks. Venter reddish-brown with clusters of cream pustules. Flanks with clusters of yellow pustules on. Palmar and plantar surfaces pale pink. Measurements of the holotype (in mm).��� SVL = 41.3; TIB = 13.5; FL = 16.2; HW = 13.5; HL = 12.4; IOD = 5.0; IND = 3.5; EN = 2.7; ED = 2.9; NR = 2.4; EW = 3.0. Morphometric ratios for holotype are followed by ranges of seven paratypes in parentheses: TIB/SVL = 0.33 (0.29���0.34); HW/HL = 1.09 (1.09���1.25); HL/SVL = 0.3 (0.25���0.29); HW/SVL = 0.33 (0.30���0.34). Osteological characteristics.���Six discrete presacral vertebrae present, atlas and axis free, width of transverse processes and sacral diapophyses: Sacrum = IV> III>V ��� VI> II. Transverse processes nearly perpendicular to notochordal axis in Presacrals V and VI, directed anteriorly in Presacral II, and posteriorly in Presacrals III, IV. The bony sacral diapophyses are broadly expanded and fused to the urostyle, which is expanded laterally and bears a low dorsal crest throughout a third of its length (Fig. 4 A). The carpus is composed of a radiale, ulnare, Element Y, Carpal 1, and a large postaxial element assumed to represent a fusion of Carpals 2���4. The prepollex is composed of one small bone that articulates with the proximal end of Metacarpal I. All carpal elements are tightly articulated. Phalangeal formula of hand: 2 - 2-3 - 3 (Fig. 4 B); phalangeal formula of foot variable: 1-2 - 2-4 - 2 (QCAZ 7684) or 1-2 - 2-4 - 1 (QCAZ 13320). Posterior sternal elements covered by m. vagina recti (see da Silva and Mendelson 1999). Stapes absent. Va r ia ti o n. ��� Snout without papilla at tip, rounded in dorsal and lateral views (QCAZ 7685). Snout slightly protruding in lateral view (EPN 7083). Atlas and presacral Vertebra II partially fused (QCAZ 7684). Variation in measurements and proportions is presented in Table 1. In preservative, tongue slightly pigmented at base (QCAZ 13320) or unpigmented (QCAZ 13271, EPN 7082 ��� 83); venter mostly dark gray with pale yellow pustules (QCAZ 7685, EPN 7083) or mostly pale yellow with grayish-brown reticulations (QCAZ 13271, QCAZ 13320, EPN 7081 ��� 82). In life, dorsum pale reddish brown (QCAZ 13271, QCAZ 13320); iris dark gray with white flecks (QCAZ 13271, QCAZ 13320, EPN 7082 ��� 83); palmar and plantar surfaces pink (QCAZ 13271, QCAZ 13320) or red (EPN 7082 ��� 83). Etymology.��� The specific name is an indeclinable noun and refers to the Laguna de Puruanta (or Puruhanta) in the vicinity of the type locality. Ecology.��� Osornophryne puruanta is known from eight specimens, all adult females. Two specimens were collected active at night, one on an epiphytic bromeliad (QCAZ 7685) one on the ground (QCAZ 7684) and the remaining six specimens were found under logs (QCAZ 11471, QCAZ 13271, and QCAZ 13320) or in the bases of plants of the family Poaceae (EPN 7081 ��� 83) during the day. This species is probably nocturnal since no individuals were observed active during the day despite extensive search efforts. It is likely that Osornophryne species are semi-fossorial, at least during the day. Clutch size in Osornophryne puruanta is comparable to that reported for O. guacamayo (35��� 50 eggs; Gluesenkamp and Acosta, 2001). One specimen of O. puruanta (QCAZ 7684) had 30 mature ovarian eggs with a maximum diameter of 3.15 mm (x = 2.73 �� 0.14), and another (QCAZ 13320) had 31 mature ovarian eggs with a maximum diameter of 3.61 mm (x = 3.05 �� 0.26). Osornophryne puruanta, as other member of the genus, is presumed to have terrestrial eggs that undergo direct development (Ruiz-Carranza & Hern��ndez- Camacho 1976). Stomach contents of QCAZ 7684 consisted of a beetle (Chrysomelidae), hymenopteran wings, a coleopteran larvae, and incidental plant material. Anurans present at the type locality include Pristimantis buckleyi, P. trepidotus, Phrynopus brunneus, and three unidentified species of Pristimantis. Vegetation at Laguna de Puruanta consists of dense forest patches dominated by Miconia trees, which are covered with a variety of epiphytic plants including arboreal bromeliads of the genus Tillandsia, ferns of several genera (Polipodium, Jamesonia, and Elaphoglossum; Polypodiaceae), orchids (Epidendrum and Pleurothallis), and mosses. Patches of forest are surrounded by members of the Ericaceae (Pernettya, Cavendishia, Psammisia, and Va c c i n u m), Asteraceae (Baccharis and Gynoxis), and Melastomataceae (Brachyotum) families. Flat, wet areas are dominated by grasses (Cortaderia and Festuca), Hypericaceae (Hypericum laricifolium) and Asteraceae (Loricaria, Hypochaeris, and Werneria) (Museo Ecuatoriano de Ciencias Naturales 1987). Arborescent ferns (Blechnum) and terrestrial bromeliads (Puya) are present in the area. The anuran community at Laguna de San Marcos includes Pristimantis curtipes, P. devillei, P. trepidotus, Gastrotheca sp., and Atelopus ignescens (not observed since 1985). The forest canopy reaches 12 m and is dominated by Gaiadendorn punctatum (Loranthaceae), Weinmmania fagaroides (Cunnoniaceae), Oreopanax sp. (Araliaceae), and Miconia sp. (Melastomataceae). Numerous epiphytes (primarily mosses, orchids, lichens, and bromeliads) cover the trees. The middle stratum of the forest is composed of the genera Miconia (Melastomataceae), Berberis (Berberidaceae), and Gynoxys (Asteraceae). The lower stratum includes terrestrial ferns and species of the genera Piper (Piperaceae), Loricaria (Asteraceae), Pernnetya (Ericaceae), Ribes (Grossulariaceae), and Brachyotum (Melastomataceae) (S. R. Espinosa and R. Mont��far, pers. comm.). Both localities are in areas classified as Evergreen High Montane Forest (Valencia et al. 1999) or Very Wet Montane Forest life zone (Holdridge 1967), and have annual temperatures between 7 and 12 ��C and an annual rainfall of 1000���2000 mm (Ca��adas-Cruz 1983). Distribution.��� This species is known only from Laguna de Puruanta (00�� 12 ' N, 77 �� 57 ' W, 3000���3500 m) on the western slopes of the Cordillera de Pimampiro, Provincia Imbabura, and the vicinity of Laguna de San Marcos (00��07' 35 " N, 77 �� 55 ' 50 " W, 3400 m). Laguna de San Marcos is located on the eastern slopes of the Filo de Talcas, approximately 6 km S Laguna de Puruanta, on the border between Provincia Pichincha and Provincia Sucumb��os (Fig. 5)., Published as part of Gluesenkamp, Andrew G. & Guayasamin, Juan M., 2008, A new species of Osornophryne (Anura: Bufonidae) from the Andean highlands of northern Ecuador, pp. 18-28 in Zootaxa 1828 on pages 19-25, DOI: 10.5281/zenodo.183157, {"references":["Ruiz-Carranza, P. M., & Hernandez-Camacho, J. I. (1976) Osornophryne, genero nuevo de anfibios bufonidos de Colombia y Ecuador. Caldasia, 11, 93 - 148.","da Silva, H. R., & Mendelson III, J. R. (1999) A new organ and sternal morphology in toads (Anura: Bufonidae): descriptions, taxonomic distribution, and evolution. Herpetologica, 55, 114 - 126.","Gluesenkamp, A. G., & Acosta, N. (2001) Sexual dimorphism in Osornophryne guacamayo with notes on natural history and reproduction in the species. Journal of Herpetology, 35, 148 - 151.","Museo Ecuatoriano de Ciencias Naturales (1987) Expedicion cientifica de reconocimiento ecologico de la Laguna de Puruhanta. Revista Geografica, 25, 7 - 28.","Valencia, R., Ceron, C., Palacios, W., & Sierra, R. (1999) Las formaciones naturales de la sierra del Ecuador. Pp. 79 - 108. In: R. Sierra (Ed.), Propuesta Preliminar de un Sistema de Clasificacion de Vegetacion para el Ecuador Continental. Proyecto INEFAN / GEF-BIRF y EcoCiencia. Quito, Ecuador.","Holdridge, L. R. (1967) Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica.","Canadas-Cruz, L. (1983) El mapa bioclimatico y ecologico del Ecuador. Banco Central del Ecuador, Quito, Ecuador."]}

Cochranella pulverata (Peters, 1873) Hyla pulverata Peters, 1873: 614. Holotype: ZMB 7842, according to Duellman, 1977: 194. Type locality: ���Chiriqui���, Panama; at the time of the description "Chiriqui" included both Atlantic and Pacific versants of extreme western Panama according to Myers, 1982: 5. Centrolene pulveratum��� Dunn, 1931: 393. Centrolenella pulveratum��� Taylor, 1949: 258. Cochranella pulverata��� Taylor, 1951: 35. Cisneros-Heredia and McDiarmid, 2006, Zootaxa, 1244: 1. Centrolenella pulverata��� Savage 1967: 328. Hyalinobatrachium pulveratum��� Ruiz-Carranza and Lynch, 1991: 24. Cochranella petersi��� Goin, 1961: 96. Holotype: BM 1902.5. 27.24. Type locality: "Rio Durango, [Province of Esmeraldas] N. W. Ecuador ". New synonymy. Centrolenella petersi��� Goin, 1964: 1. Hyalinobatrachium petersi��� Ruiz-Carranza and Lynch, 1998: 573. Diagnosis. Cochranella pulverata differs from all other Glassfrogs by having a dorsum green with small white spots, completely transparent ventral parietal peritoneum, white hepatic and gastrointestinal peritonea, a sloping snout in lateral profile, and by lacking humeral spines (Fig. 2). Characterization. (1) dentigerous process of the vomer with 2���4 teeth; (2) snout rounded in dorsal aspect, sloping in lateral profile; (3) tympanum visible, relatively small, its diameter 20.2���23.3 % of eye diameter; tympanic annulus visible except for dorsal border covered by supratympanic fold; tympanic membrane differentiated and translucent, pigmented as surrounding skin; (4) dorsal surfaces shagreen; males with small spicules visible under magnification; (5) ventral surfaces granular, thighs below vent lacking pair of enlarged tubercles; (6) ventral parietal peritoneum completely transparent; pericardium and gastrointestinal peritoneum white; (7) bulbous liver covered by white peritoneum; (8) humeral spines absent; (9) webbing between Fingers I and II absent or basal; webbing formula for outer fingers: II (1 +��� 1 1 / 3) ��� (2 4 / 5 ��� 3 ���) III (1 1 / 3 ��� 1 2 / 3) ��� (1 +��� 2 ���) IV; (10) feet about two-thirds webbed; webbing formula: I (1 ��� ��� 1) ��� (1 2 / 3 ��� 2 ���) II (1 ��� ��� 1) ��� (1 3 / 4 ��� 2 ���) III (1 ��� 1 +) ��� (1 2 / 3 ��� 2 +) IV (2 ��� ��� 2 +) ��� (1 ��� ��� 1 +) V; (11) metacarpal, ulnar, metatarsal, and tarsal enameled folds present, having low tubercles that give them an undulated shape; (12) nuptial pad Type-I in adult males; concealed prepollex; (13) Fingers I and II about equal in length (FII/FI = 0.980���1.025); (14) disc of Finger III small, its width 20.1��� 23.5 % of eye diameter; (15) in life, dorsum green with small white flecks and dots; bones green; (16) in preservative, dorsum cream to light lavender with small un-pigmented or white flecks and dots; (17) iris graywhite with thin dark gray reticulations and minute yellow flecks; a thin yellow to cream circumpupillary ring borders the pupil; (18) melanophores partially covering dorsal surface of Finger IV, absent from Fingers I���III; (19) males call from the upper side of leaves; call usually emitted as a series of three notes (each note = 0.05 s), with a short pause between them (internode pause = 0.5��� 0.8 s), dominant frequency of 5600���6200 Hz (Savage & Starrett 1967; Iba��ez et al. 1999; Kubicki 2007); (20) fighting behavior unknown; (21) egg masses deposited on the upper side of leaves; no parental care provided (Kubicki 2007); (22) in tadpoles, tooth row formula 2 / 3; A 2 tooth row is broadly separated in the center (Hoffmann 2004); (23) in adult males, SVL 22.0��� 24.5 mm (n = 13); in adult females, SVL 25.3���28.3 mm (n = 5; McCranie & Wilson, 2002; this work)., Published as part of Guayasamin, Juan M., Cisneros-Heredia, Diego F. & Castroviejo-Fisher, Santiago, 2008, Taxonomic identity of Cochranella petersi Goin, 1961 and Centrolenella ametarsia Flores, 1987, pp. 25-34 in Zootaxa 1815 on page 27, DOI: 10.5281/zenodo.182892, {"references":["Peters, W. C. H. (1873) Uber eine neue Schildkrotenart, Cinosternon Effeldtii und einige andere neue oder weniger bekannte Amphibien. Monatsberichte der Koniglich preussischen Akademic der Wissenschaften zu Berlin, 1873, 603 - 618.","Taylor, E. H. (1951) Two new genera and a new family of tropical American frogs. Proceedings of the Biological Society of Washington, 64, 33 - 40.","Cisneros-Heredia, D. F. & McDiarmid, R. W. (2006) A new species of the genus Centrolene (Amphibia: Anura: Centrolenidae) from Ecuador with comments on the taxonomy and biogeography of Glassfrogs. Zootaxa, 1244, 1 - 32.","Ruiz-Carranza, P. M. & Lynch, J. D. (1991) Ranas Centrolenidae de Colombia I: propuesta de una nueva clasificacion generica. Lozania, 57, 1 - 30.","Goin, C. J. (1961) Three new centrolenid frogs from Ecuador. Zoologischer Anzeiger, 166, 95 - 104.","Ruiz-Carranza, P. M. & Lynch, J. D. (1998) Ranas Centrolenidae de Colombia XI. Nuevas especies de ranas de cristal del genero Hyalinobatrachium. Revista de la Academia Colombiana de Ciencias Exactas, Fisicas y Naturales, 22, 571 - 586.","Ibanez, R., Rand, A. S. & Jaramillo, C. A. (1999) Los anfibios del Monumento Natural Barro Colorado, Parque Nacional Soberania y areas adyacentes. Mizrachi, E. and Pujol, S. A. Santa Fe de Bogota. 187 pp.","Kubicki, B. (2007) Ranas de vidrio de Costa Rica / Glass frogs of Costa Rica. Editorial INBio. Santo Domingo de Heredia. C. R., 304 pp.","Hoffmann, H. (2004) Description of the previously unknown tadpole of Hyalinobatrachium pulveratum (Anura: Centrolenidae). Revista de Biologia Tropical, 52, 219 - 228.","McCranie, J. R. & Wilson, L. D. (2002) The Amphibians of Honduras. Society for the Study of Amphibians and Reptiles. Ithaca, New York, USA."]}

Cochranella oyampiensis (Lescure, 1975) Centrolenella oyampiensis Lescure, 1975: 100. Holotype: MNHNP 1973.1673. Type locality: "village Zidok (Haut- Oyapock), Guyane Fran��aise". Centrolenella ametarsia��� Flores, 1987: 185. Holotype: MCZ A 96522. Type locality: ���the headwaters of R��o Caiwima, a tributary of the R��o Amaca-Yacu, ca. 70 km NNE Puerto Nari��o, Amazonas, Colombia (approximately 3 �� 20 ' S, 70 �� 20 ' W)���. New synonymy. Cochranella ametarsia��� Ruiz-Carranza and Lynch 1991: 21. Diagnosis. Among Glassfrogs, Cochranella oyampiensis is unique by having a green dorsum with small black spots, a small size (adults = 20 mm), transparent hepatic peritoneum, white gastrointestinal peritoneum, anterior third of ventral parietal peritoneum white, and a distinct prepollex. The only Glassfrog that could be confused with C. oyampiensis is C. helenae, which differs by having a yellow iris (grayish white with a fine dark reticulation in C. oyampiensis), dorsum light greenish yellow with dark punctuations (green with dark with dark punctuations in C. oyampiensis; Fig. 2), and a mostly white hepatic peritoneum (hepatic peritoneum mostly transparent, showing the brown liver, except for some iridophores on the upper border in C. oyampiensis; Fig. 3). Characterization. (1) dentigerous process of the vomer with one tooth or lacking teeth; (2) snout rounded in dorsal and lateral views; (3) tympanum visible, moderate in size, its diameter 25.8���35.4 % of eye diameter; tympanic annulus visible except for posterodorsal border covered by supratympanic fold; tympanic membrane differentiated and translucent, pigmented as surrounding skin; (4) dorsal surfaces shagreen; males and females lack spinules; (5) ventral surfaces granular, a pair of enlarged tubercles below the vent; (6) anterior 25���40 % of ventral parietal peritoneum white, posterior portion transparent; pericardium and gastrointestinal peritoneum white; (7) lobed liver covered by an almost completely transparent peritoneum except for its anterior part that may be covered by a thin layer of iridophores; (8) humeral spines absent; (9) webbing between Fingers I���III absent, moderate between outer fingers; webbing formula: III (2 ��� ��� 2 1 / 3) ��� (1 +��� 2 ���) IV; (10) webbing between toes moderate; webbing formula: I 1 ��� (2 ��� ��� 2) II (1 ��� 1 +) ��� (2 ��� 2 1 / 4) III (1 +��� 1 1 / 2) ��� 2 + IV (2 ��� 2 1 / 3) ��� 1 V; (11) low ulnar fold, lacking iridophores; low inner tarsal fold present, lacking iridophores; outer tarsal fold absent; (12) nuptial pad Type I in males; distinct prepollex (distal portion separated from Finger I); (13) Fingers I slightly longer than Finger II (FII/FI = 0.840���0.921); (14) disc of Finger III moderate, its width 31.0��� 42.3 % of eye diameter; (15) in life, dorsum green with small dark flecks; bones green; (16) in preservative, dorsum lavender with dark flecks; (17) iris grayish white with a fine dark reticulation; (18) melanophores covering dorsal surface of Fingers III and IV, absent from Fingers I and II; (19) males call from the upper side of leaves; single and double note advertisement call of 0.10��� 0.15 s duration, emphasized frequency of 4640��� 5160 Hz (Zimmerman & Bogart 1984); (20) fighting behavior unknown; (21) eggs deposited on the upper- or underside of leaves (Lima et al., 2005); parental care unknown; (22) tadpoles unknown; (23) in adult males, SVL 17.1���20.1 mm (= 18.8 �� 1.250, n = 6); in two adult females, SVL 19.8���19.9 mm; Lima et al. (2005) provide the following data for the species in central Amazonia: SVL in males 17���21 mm, in females 21���24 mm. Remarks. Our conclusions are the result of the analysis of morphological traits; however, we cannot rule out the existence of morphologically cryptic species, a possibility that has to be addressed with acoustic and/ or molecular data, unavailable now. The wide distribution of the C. oyampiensi s (from the Guianas across western Amazonia; Lescure 1975; Flores 1987; Lescure & Marty 2001; Lima et al. 2005; Guayasamin et al. 2006; Cisneros-Heredia & McDiarmid 2007 b; Kok & Castroviejo 2008) opens the possibility of testing hypotheses of diversification through phylogeographic studies., Published as part of Guayasamin, Juan M., Cisneros-Heredia, Diego F. & Castroviejo-Fisher, Santiago, 2008, Taxonomic identity of Cochranella petersi Goin, 1961 and Centrolenella ametarsia Flores, 1987, pp. 25-34 in Zootaxa 1815 on pages 29-32, DOI: 10.5281/zenodo.182892, {"references":["Lescure, J. (1975) Contribution a l'etude des amphibiens de Guyane Francaise. V. Les centrolenidae. Bulletin de la Societe Zoologique de France, 100, 385 - 394. Lescure, J. and Marty, C. 2000. Atlas des Amphibiens de Guyane. Patrimoines Naturels, Paris, France.","Flores, G. (1987) A new Centrolenella from the Amazonian lowlands of Colombia. Journal of Herpetology, 21, 185 - 190.","Ruiz-Carranza, P. M. & Lynch, J. D. (1991) Ranas Centrolenidae de Colombia I: propuesta de una nueva clasificacion generica. Lozania, 57, 1 - 30.","Zimmerman, B. & J. P. Bogart. (1984) Vocalizations of primary forests frog species in central Amazon. Acta Amazonica 14 (3 / 4): 473 - 519.","Lima, A., Magnusson, W. E., Menin, M., Erdtmann, L. K., Rodrigues, D. J., Keller, C., & Hodl, W. (2005) Guide to the Frogs of the Reserva Adolpho Ducke: Central Amazonia. Atemma Design Editorial, Manaus, Brasil.","Cisneros-Heredia, D. F. & McDiarmid, R. W. (2007 b) Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. Zootaxa, 1572, 1 - 82. Fabrezi, M. & Alberch, P. (1996) The carpal elements of anurans. Herpetologica, 52, 188 - 204.","Kok, P. J. R. & Castroviejo-Fisher, S. (2008) Glassfrogs (Anura: Centrolenidae) of Kaiteur National Park, Guyana, with notes on the distribution and taxonomy of some species of the family in the Guiana Shield. Zootaxa, 1680, 25 - 53."]}

Status of Centrolenella ametarsia. In the description of Centrolenella ametarsia (now in the genus Cochranella), Flores (1987) noted its close morphological similarity to Cochranella oyampiensis. To differentiate the two species, he listed the following traits: fully webbed outer fingers in C. oyampiensis (III 2 1 / 3 ��� 2 ��� IV in C. ametarsia), white ventral parietal peritoneum (transparent in C. ametarsia), transparent visceral peritoneum (white in C. ametarsia), concealed tympanum (lower two-thirds exposed in C. ametarsia), and concealed prepollex (distinct in C. ametarsia). Cisneros-Heredia and McDiarmid (2007 b) and Kok and Castroviejo-Fisher (2008) suggested that C. ametarsia could be a junior synonym of C. oyampiensis. After examination of the holotypes of C. oyampiensis and C. ametarsia, as well as recently collected specimens from the surroundings of their type localities and additional areas (Appendix 1), we found that the differences listed for C. ametarsia and C. oyampiensis by Flores (1987) are the product of a limited sample size and preservation artifacts. Individuals found in Leticia, near the type locality of Cochranella ametarsia, and in Amazonian Ecuador have a webbing formula of the hand that contains the variation observed in both holotypes [i.e., ametarsia and oyampiensis; III (2 ��� ��� 2 1 / 3) ��� (1 +��� 2 ���) IV]. Flores (1987) mentioned that C. ametarsia has a completely transparent ventral parietal peritoneum. This interpretation is erroneous given that in the types of C. ametarsia and C. oyampiensis all iridophores responsible for its white coloration are dissolved in the preservation liquid. In the newly collected material (except ICN 50847 where iridophores are also lost), the anterior 1 / 4 ��� 1 / 3 of the ventral parietal peritoneum is white, being the posterior portion transparent. In all the examined specimens of C. ametarsia and C. oyampiensis, most of the tympanum (lower three-fourths) is exposed, and a distinct prepollex is present. Additionally, Cochranella oyampiensis does not have a white hepatic peritoneum as reported by Se��aris and Ayarzag��ena (2005); the specimens examined by these authors are in fact C. helenae (Ayarzag��ena, 1992; see Kok & Castroviejo-Fisher 2008). We conclude that the two species are morphologically indistinguishable and consider Centrolenella ametarsia Flores, 1987, a junior synonym of Centrolenella oyampiensis Lescure, 1975. Below, we provide a synonymy, diagnosis, and characterization of Cochranella oyampiensis., Published as part of Guayasamin, Juan M., Cisneros-Heredia, Diego F. & Castroviejo-Fisher, Santiago, 2008, Taxonomic identity of Cochranella petersi Goin, 1961 and Centrolenella ametarsia Flores, 1987, pp. 25-34 in Zootaxa 1815 on page 28, DOI: 10.5281/zenodo.182892, {"references":["Flores, G. (1987) A new Centrolenella from the Amazonian lowlands of Colombia. Journal of Herpetology, 21, 185 - 190.","Cisneros-Heredia, D. F. & McDiarmid, R. W. (2007 b) Revision of the characters of Centrolenidae (Amphibia: Anura: Athesphatanura), with comments on its taxonomy and the description of new taxa of glassfrogs. Zootaxa, 1572, 1 - 82. Fabrezi, M. & Alberch, P. (1996) The carpal elements of anurans. Herpetologica, 52, 188 - 204.","Kok, P. J. R. & Castroviejo-Fisher, S. (2008) Glassfrogs (Anura: Centrolenidae) of Kaiteur National Park, Guyana, with notes on the distribution and taxonomy of some species of the family in the Guiana Shield. Zootaxa, 1680, 25 - 53.","Senaris, J. C. & Ayarzaguena, J. (2005) Revision taxonomica de la Familia Centrolenidae (Amphibia; Anura) de Venezuela. Publicaciones del Comite Espanol del Programa Hombre y Biosfera - Red IberoMaB de la UNESCO. Sevilla, Espana, 337 pp.","Ayarzaguena, J. (1992) Los centrolenidos de la Guayana venezolana. Publicacion de la Asociacion de Amigos de Donana, 1, 1 - 48.","Lescure, J. (1975) Contribution a l'etude des amphibiens de Guyane Francaise. V. Les centrolenidae. Bulletin de la Societe Zoologique de France, 100, 385 - 394. Lescure, J. and Marty, C. 2000. Atlas des Amphibiens de Guyane. Patrimoines Naturels, Paris, France."]}