**Tadpole:** Not described.

**Distribution** (Figure 46): *Centrolene lynchi* is known from the Pacific slope of the Cordillera Occidental of the Andes in Ecuador and southern Colombia. In Colombia, the species has been reported in only one locality (Reserva La Planada, 7 km route of Chucunés, 1780 m; as *C*. *scirtetes* [86], but see Taxonomic Remarks); specimens cited as *C*. *lynchi* by Coloma et al. [126] from Risaralda Department, Colombia, actually corresponds to *C*. *quindianum* (Marco Rada, pers. obs.). In Ecuador, *Centrolene lynchi* is known from seven localities from the Pacific slope of the Cordillera Occidental of the Andes at elevations of 1140–1852 m (see Specimens Examined), with a potential distribution of 1442 km2. The habitat of the species in Ecuador is within the Western Foothill Forest and the Western Montane Forest regions.

**Conservation status:** Globally, *Centrolene lynchi* is currently listed as *Endangered* by the IUCN [126]. Arteaga et al. [87] also suggested the *Endangered* category for Ecuadorian populations. Although the distribution of the species is larger than previously thought (see Distribution and Remarks), the species has not been found in some of its historic localities in the last 20 years, suggesting that the species has suffered population declines (i.e., San Francisco de Las Pampas). In recent years, reproductive populations of *C*. *lynchi* have been found at Reserva Las Gralarias (2009–2019) and at Bosque Protector Río Guajalito (2006) ([87], JMG pers. obs., DFCH pers. obs., Mario Yánez-Muñoz pers. comm.). The amphibian chytrid fungus *Batrachochytrium dendrobatidis* has been found infecting *C*. *lynchi* at Reserva Las Gralarias, but no recent declines have been observed [92].

**Figure 46.** Distribution of *Centrolene lynchi* in Ecuador (yellow dots).

**Evolutionary relationships** (Figure 24): *Centrolene lynchi* is sister to a clade formed by *C*. *sabini* and an unidentified species of *Centrolene*.

**Taxonomic remarks:** *Centrolene lynchi* was described by Duellman [127] from a locality 4 km NE (by road) of Dos Ríos, Provincia de Pichincha. After a few years, Flores [46] described *Centrolene gemmata* from San Francisco de Las Pampas, Provincia de Cotopaxi. Flores [46] mentioned that *Centrolene lynchi* differs from *Centrolene gemmata* mainly by: (i) Having a snout truncated in dorsal view and round in profile (snout round in dorsal view and slightly anteroventrally sloping in *C*. *gemmata*); (ii) having slightly more webbing on the hands (webbing formula of IV 2—2 V in *Centrolene lynchi;* webbing formula of IV (2–2<sup>1</sup>/3)—(2−–2<sup>+</sup>) V in *C*. *gemmata*); and (iii) in overall head shape, with a squatter, more semicircular head shape when viewed from above, with little or no post-cephalic constriction, and nostrils only very slightly protuberant, in contrast to the more elongated, circular head shape of *gemmata* when viewed from above, with a marked post-cephalic constriction and very protuberant nostrils. We have examined the complete type series of *Centrolene lynchi* and three paratypes of *C*. *gemmata* (MCZ A-104397, A-104074, A-104077) and, as noted by Cisneros-Heredia and McDiarmid [17], find that the differences mentioned above are not consistent. Several of the characteristics present in *C*. *gemmata* fall into the variation observed in *Centrolene lynchi*, and others are the product of preservation artifacts. It is

clear that the type material of *C*. *gemmata* was poorly preserved, causing dehydration of the specimens, which usually produces changes in head shape (i.e., post-cephalic constriction and protuberant nostrils present in the type series of *C*. *gemmata*). The examined type series of *C*. *gemmata* has a snout that is truncated to slightly sloping in profile; the exact same variation is present in the type material of *Centrolene lynchi*. Also, the hand webbing formula in the two species are nearly identical [III (2–2<sup>1</sup>/4)—(2−–2<sup>+</sup>) IV in *Centrolene lynchi*, and III (2–2<sup>1</sup>/3)—(2−–2<sup>+</sup>) IV in *C*. *gemmata*]. Therefore, none of the characters listed by Flores (1985) to differentiate *C*. *gemmata* from *Centrolene lynchi* are valid. Based on the evidence mentioned above, we consider these to represent the same species and place *Centrolene gemmata* in the synonymy of *Centrolene lynchi*.

Duellman and Burrowes [86] described *Centrolenella scirtetes* from a locality 1.4 km SW Tandayapa (Ecuador, male holotype KU 202720) and from Reserva La Planada (Colombia, two female paratypes IND-AN 1405 and 1533). For unknown reasons, *C*. *scirtetes* was never compared with specimens of *Centrolene lynchi*. As pointed out by Cisneros-Heredia and McDiarmid [17], the holotype of *C*. *scirtetes* is undistinguishable from the holotype of *C*. *lynchi* (KU 164691). Consequently, herein we formally place *C*. *scirtetes*, as defined by its holotype, in the synonymy of *Centrolene lynchi*. We have not examined the paratypes of *C*. *scirtetes*, but material from Colombia identified as *C*. *scirtetes* (ICN 12172–74) and resembling the description and photograph of the paratypes provided by Duellman and Burrowes (1989) are indistinguishable from individuals of *Nymphargus gri*ffi*thsi*, in which the dorsum has black flecks and the humeral crista ventralis presents a distal prolongation. Accordingly, we conclude that the Colombian material assigned to *C*. *scirtetes* is in fact representative of *N*. *gri*ffi*thsi*.

**Specimens examined:** *Centrolene lynchi:* Ecuador: *Provincia de Cotopaxi:* San Francisco de Las Pampas, just NW of junction of río Las Juritas and río Toachi (0.433 S, 78.9667 W; ca. 1500 m), MCZ A-104397, A-104074, A-104077; *Provincia de Pichincha:* 1.4 km SW of Tandayapa (0.033 S, 78.7667 W; 1820 m), KU 202720; Tandapi (0.4164 S, 78.7989 W; 1460 m), KU 118036, 118047–50, 178095–104; 2.1 km E Tandapi (0.4258 S, 78.7853 W; 1500 m), MCZ A-93313–14, 95742; Reserva Las Gralarias (0.01675 S, 78.73165; 1852 m), QCAZ 40191–2, 40194. *Provincia de Santo Domingo de los Tsáchilas:* stream 4 km northeast (by road) of Dos Ríos (0.3028 S, 78.8678 W; 1140 m), KU 164691 (holotype), KU 164692–99 (paratypes); 14.4 km ENE La Palma on the road La Palma-Chiriboga (0.25 S, 78.846 W; 1380 m), MCZ A-91455.

*Centrolenella medemi* Cochran and Goin, 1970 [96]. Holotype: USNM 152277.

Type locality: "Puerto Asís, upper Río Putumayo, [Comisaría] Putumayo, Colombia" (apparently in error; see Distribution).

*Centrolene medemi—*Ruiz-Carranza and Lynch, 1991 [6].

*"Centrolene" medemi*—Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, and Vilà, 2009 [1].

**Common names:** English: Medem's Glassfrog. Spanish: Rana de Cristal de Medem. **Etymology:**Theepithet*medemi*honorsDr.FredMedem,whodiscoveredthespecies[96].

 **Identification:** *"Centrolene" medemi* is unique among Ecuadorian glassfrogs by having a dark olive-green dorsum with large greenish–cream spots (Figure 47), wide disc on Finger III (>80% of eye diameter), small tympanum (<20% of eye diameter), and fully webbed foot. Adults are robust and relatively large (SVL 25.5–44.3 mm).

**Figure 47.** *"Centrolene" medemi* in life. Adult female (KU 164493) from 2 km SSW of junction between Río Reventador and Baeza-Lumbaqui road, Ecuador. Photo by William E. Duellman.

**Diagnosis:** (1) Teeth on dentigerous process of the vomer present or absent, each process bearing zero to four teeth; (2) snout round in dorsal profile, truncated to slightly protruding in lateral profile (Figure 48); (3) tympanum small, tympanum diameter 17.1%–18.5% of eye diameter, dorsal third of tympanum covered by supratympanic fold, tympanic membrane pigmented and not clearly differentiated from surrounding skin; (4) dorsal surfaces of males and females smooth to shagreen; males with small spicules on dorsum and flanks; (5) pair of slightly enlarged subcloacal warts; (6) anterior half of the ventral parietal peritoneum white (condition P2); silvery white pericardium; no iridophores in peritonea covering the intestines, stomach, testes, kidneys, gall bladder, and urinary bladder (condition V1); (7) liver tetralobed, two large ventral lobes covering two smaller lobes; hepatic peritoneum lacking iridophores (condition H0); (8) in males, small humeral spines present; (9) hand

webbing: Webbing between Fingers I, II, and III absent or basal, extensive webbing between Fingers III and IV; webbing formula as follows: II (1+–2)—3+ III (1<sup>1</sup>/4–2)—(0–11/4) IV; (10) fully webbed foot: I (0–0+)—(0+–1) II (0–0+)—(0–1) III (0–0+)—(0+–1) IV (1−–1)—0<sup>+</sup> V; (11) ulnar and tarsal folds low; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger I slightly shorter than Finger II (Finger I about 91%–98% length of Finger II); (14) disc of Finger III large, 80%–91% of eye diameter; (15) in life, dorsal surfaces of head, body, and limbs olive green to greyish brown with large (up to 2.8 mm) cream spots (Figure 47); bones bluish green or green; (16) in ethanol, dorsal surfaces of head, body, and limbs pale brown with large cream spots; (17) iris greenish brown with black reticulation; (18) melanophores covering dorsal surfaces of fingers and toes; (19) calling behavior unknown; (20) fighting behavior unknown; (21) eggs deposited on rocks along streams; parental care unknown; (22) tadpoles unknown; (23) medium body size; in adult males, SVL 25.5–30.8 mm; in adult females, SVL 34.7–44.3 mm.

**Figure 48.** *"Centrolene" medemi*. (**A**) Head in lateral view, KU 164493. (**B**) Hand in ventral view, KU 164494. Illustrations by Juan M. Guayasamin.

**Variation:** The size of the nuptial pad varies in males and can be restricted to the dorsolateral area of the base of Finger I or extend until reaching the typical Type I morphology.

**Color in life** (Figure 47): At night, dorsum black with bluish green spots; venter dull blue. By day, dorsum dark olive-brown with pale green spots; flanks cream. Lining of mouth and tongue pale blue. Bones green. Iris dull greyish–bronze with minute black flecks. Webbing pale yellowish tan (W. E. Duellman field notes, 19 March 1975).

**Color in ethanol:** Dorsal surfaces of head, body, and limbs pale brown with large cream spots. White lining on the anterior half of the ventral parietal peritoneum; silvery white pericardium; no iridophores in peritonea covering the liver, intestines, stomach, testes, kidneys, gall bladder, and urinary bladder.

**Biology and ecology:** During the night, adults have been found on rocks, in rock crevices, and on rock cli ffs along streams; juveniles were found on palm leaves or in rock crevices [128]. The oviductal eggs of one female (KU 164493) are dark brown. It is likely that females of *"Centrolene" medemi* deposit their eggs on rocks, as reported in species with similar microhabitats (e.g., *Centrolene geckoidea*, *"Centrolene" petrophilum* [113,129]). Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 49): *"Centrolene" medemi* is known from localities on the eastern and western slopes of the Cordillera Oriental of the Colombian Andes and the Amazonian slopes of the Cordillera Oriental of Ecuador at elevation between 790 and 1800 m [101,128,130]. In Ecuador, *"Centrolene" medemi* has been registered from a single stream nearby Volcán Reventador at 1490 m (Specimens Examined), within the Eastern Montane Forest ecoregion. Although this species was described from a locality in lowlands of Amazonian Colombia (Puerto Asís, upper Río Putumayo at about 280 m [96]), all subsequent records are from Andean localities. Based on this evidence, Ruiz-Carranza et al. [101] rejected the record from Amazonian Colombia.

**Figure 49.** Distribution of *"Centrolene" medemi* in Ecuador (yellow dot).

**Conservation status:** Globally, *"Centrolene" medemi* is currently listed as *Endangered* by the IUCN [131]. In Ecuador, the only record is from a stream nearby Volcán Reventador on 19 March 1975 [130], and area that has been visited several times during the last 10 years. We sugges<sup>t</sup> that the species should be locally listed as *Critically Endangered*.

**Evolutionary relationships** (Figure 24): Based on morphological traits, *"Centrolene" medemi* was placed in the *Centrolene geckoidea* species group by Ruiz-Carranza and Lynch [6]. Guayasamin et al. [1] considered the generic placement of *"Centrolene" medemi* as *incertae sedis* within the subfamily Centroleninae. No genetic data are available for this species.

**Specimens examined:** *"Centrolene" medemi:* Ecuador: *Provincia de Napo:* Stream 2 km SSW of junction between Río Reventador and Baeza-Lumbaqui road (0.1 S; 77.6 W, 1490 m), KU 164493–94.

*Centrolene peristicta* (Lynch and Duellman, 1973 [22]; Figures 50–53).

*Centrolenella peristicta* Lynch and Duellman, 1973 [22]. Holotype: KU 118051.

Type locality: "Tandapi, 1460 m, Provincia Pichincha, Ecuador".

*Centrolene peristictum*—Ruiz-Carranza and Lynch, 1991 [6]. Guayasamin, Castroviejo-Fisher,

Trueb, Ayarzagüena, Rada, and Vilà, 2009 [1].

*Centrolene peristicta*—Barrio-Amorós, Rojas-Runjaic, and Señaris, 2019 [85].

**Common names:** English: Dappled Glassfrog. Spanish: Rana de Cristal Punteada.

**Etymology:** The specific epithet is derived from the Greek *peristiktos*, meaning dappled, and refers to the spotted color pattern of the species [22].

**Identification:** *Centrolene peristicta* is distinguished from most glassfrogs by its minute body size (SVL < 21.3 mm) and having, in life, a yellowish–green dorsum with minute white flecks and small diffuse black spots (Figure 50). Additionally, adult males of *C*. *peristicta* have a small, curved humeral spine that is morphologically different from the spines present in most glassfrogs (i.e., not curved). Similar species include *C*. *antioquiensis*, *C*. *lynchi*, *C*. *pipilata*, and *Nymphargus truebae*. *Centrolene antioquiensis*, an endemic species from Colombia, is remarkably similar to *C*. *peristicta* (see Taxonomic Remarks), but *C*. *antioquiensis* has less hand webbing: hand webbing formulae in *C*. *antioquiensis*, III (2<sup>+</sup>–2−)—(2–2−) IV; in *C*. *peristicta*, III (2<sup>+</sup>–2−)—(1<sup>1</sup>/2–1<sup>+</sup>) IV. *Centrolene lynchi* is larger than *C*. *peristicta* (in males, SVL 23.3–26.5 mm in *Centrolene lynchi*) and has a very different call. *Centrolene pipilata* occurs on the Amazonian slopes of the Andes, whereas *C*. *peristicta* is restricted to the Pacific slopes of the Andes. *Nymphargus truebae*, a species only known from the Andes of southern Peru, has the same color pattern as *C*. *peristicta*, but *N*. *truebae* lacks humeral spines and webbing between Fingers III and IV (present in *C*. *peristicta*).

**Diagnosis:** (1) Vomers lacking teeth; (2) snout round in dorsal aspect, round or truncated in lateral profile (Figure 51); (3) tympanum large, oriented almost vertically, with slight lateral and posterior inclinations, its diameter 44.0%–51.9% of eye diameter; tympanic annulus visible, supratympanic fold evident; tympanic membrane translucent, partially pigmented, differentiated from surrounding skin; (4) dorsal skin shagreen with small warts corresponding to yellowish–white spots, males and females lack spicules; (5) pair of enlarged subcloacal warts (Figure 15); (6) anterior half of the ventral parietal peritoneum white, posterior half transparent (condition P2); white pericardium; iridophores partially or completely covering stomach and colon; no iridophores in peritonea covering kidneys, gall bladder, and urinary bladder (condition V2); (7) liver with four clearly defined lobes, lacking iridophores (condition H0); (8) males with conspicuous humeral spines; (9) webbing absent between Fingers I and II, reduced between Fingers II and III, and moderate to extensive between outer fingers (Figure 51); webbing formula: II (1<sup>3</sup>/4–2−)—(3<sup>+</sup>–31/4) III (2<sup>+</sup>–2−)—(1<sup>1</sup>/2–1<sup>+</sup>) IV; (10) webbing between toes extensive; webbing formula on foot I 1—(1<sup>1</sup>/2–2) II 1—(2−–2<sup>+</sup>) III (1−–1<sup>+</sup>)—(2−–2<sup>+</sup>) IV (2−–2<sup>+</sup>)—(1–1<sup>1</sup>/3) V; (11) ulnar fold present, with low white tubercles; outer tarsal fold present, with low white tubercles; internal tarsal fold low and short; (12) prepollex usually exposed; nuptial pad Type I or Type III; (13) Finger I slightly shorter or as long as Finger II (Finger I length 90.9%–102.7% Finger II); (14) disc of Finger III width about 45.6%–67.3% of eye diameter; (15) in life, dorsum green with minute yellowish–white spots and larger black spots (Figure 50); green bones; (16) in preservative, dorsum lavender with numerous minute white spots and larger diffuse dark lavender spots; (17) in life, iris white grey with a yellow hue and thin black reticulation; thin yellow line surrounds the pupil; (18) melanophores mostly absent from fingers and toes, except for a few on dorsal surfaces of Finger IV and proximal portion of Finger III; (19) males call from mostly from the lower sides of leaves; each call has one pulsed note with a duration of 0.036–0.087 s and a dominant frequency of 6471–7278 Hz; (20) fighting behavior unknown; (21) egg clutches usually deposited on the underside of leaves; short-term maternal care absent; prolonged parental care provided by males; (22) tadpoles undescribed; (23) minute body size; in adult males, SVL 17.9–21.2 mm (X = 19.9, *n* = 14); in two adult females SVL 20.8–20.9 mm.

**Figure 50.** *Centrolene peristicta* in life. (**Top row**): Male guarding two egg clutches; Reserva Las Gralarias. (**Bottom left**): Adult male guarding eggs, from Mindo Biological Reserve, QCAZ 22313; photo by Martín Bustamante. (**Bottom right**): Individual from Reserva Las Gralarias; photo by Marco Rada.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout round in dorsal aspect, round or truncated in lateral profile (Figure 51); (3) tympanum large, oriented almost vertically, with slight lateral and posterior inclinations, its diameter 44.0%–51.9% of eye diameter; tympanic annulus visible, supratympanic fold evident; tympanic membrane translucent, partially pigmented, differentiated from surrounding skin; (4) dorsal skin shagreen with small warts corresponding to yellowish–white spots, males and females lack spicules; (5) pair of enlarged subcloacal warts (Figure 15); (6) anterior half of the ventral parietal peritoneum white, posterior half transparent (condition P2); white pericardium; iridophores partially or completely covering stomach and colon; no iridophores in peritonea covering kidneys, gall bladder, and urinary bladder (condition V2); (7) liver with four clearly defined lobes, lacking iridophores (condition H0); (8) males with conspicuous humeral spines; (9) webbing absent between Fingers I and II, reduced between Fingers II and III, and moderate to extensive between outer fingers (Figure 51); webbing formula: II (1<sup>3</sup>/4–2−)—(3<sup>+</sup>–31/4) III (2<sup>+</sup>–2−)—(1<sup>1</sup>/2–1<sup>+</sup>) IV; (10) webbing between toes extensive; webbing formula on foot I 1—(1<sup>1</sup>/2–2) II 1—(2−–2<sup>+</sup>) III (1−–1<sup>+</sup>)—(2−–2<sup>+</sup>) IV (2−–2<sup>+</sup>)—(1–1<sup>1</sup>/3) V; (11) ulnar fold present, with low white tubercles; outer tarsal fold present, with low white tubercles; internal tarsal fold low and short; (12) prepollex usually exposed; nuptial pad Type I or Type III; (13) Finger I slightly shorter or as long as Finger II (Finger I length 90.%–102.7% Finger II); (14) disc of Finger III width about 45.6%–67.3% of eye diameter; (15) in life, dorsum green with minute yellowish–white spots and larger black spots (Figure 50); green bones; (16) in preservative, dorsum lavender with numerous minute white spots and larger di ffuse dark lavender spots; (17) in life, iris white grey with a yellow hue and thin black reticulation; thin yellow line surrounds the pupil; (18) melanophores mostly absent from fingers and toes, except for a few on dorsal surfaces of Finger IV and proximal portion of Finger III; (19) males call from mostly from the lower sides of leaves; each call has one pulsed note with a duration of 0.036–0.087 s and a dominant frequency of 6471–7278 Hz; (20) fighting behavior unknown; (21) egg clutches usually deposited on the underside of leaves; short-term maternal care absent; prolonged parental care provided by males; (22) tadpoles undescribed; (23) minute body size; in adult males, SVL 17.9–21.2 mm ( X = 19.9, *n* = 14); in two adult females SVL 20.8–20.9 mm.

**Figure 51.** *Centrolene peristicta*, KU 178148. ( **A**) Head in lateral view. (**B**) Hand in ventral view. (**C**) Dorsal view of Finger I, showing nuptial pad. Illustrations by Juan M. Guayasamin.

**Color in life** (Figure 50): Dorsum green with minute yellowish–white flecks and small dark grey spots; upper lip white; region below eye with small white warts; bones green; vocal sac green; upper flanks green with minute white spots; lower flanks cream with minute white spots; venter yellowish cream; fingers and toes dull yellow green; ulnar fold with thin white line or with low white ulnar tubercles; outer tarsal fold with low white tubercles; small white tubercles just posterior to cloaca.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs lavender with minute white spots and small black spots; margin of upper lip white; region below eye with small white warts; white tubercles just posterior to cloaca. White parietal peritoneum covers anterior half of venter; white pericardium; clear peritoneum on liver and kidneys; digestive tract partially or completely covered with white lining.

**Variation:** One individual (QCAZ 16313) has considerably less webbing on the hands (III 21/2—2<sup>+</sup> IV). Individuals from streams nearby the town of Mindo lack iridophores on the gastrointestinal peritonea.

**Biology and ecology:** The information shown below is from Salgado and Guayasamin [132]. At Reserva Las Gralarias, during the night, *Centrolene peristicta* is active on vegetation 40–600 cm above permanent streams in primary evergreen lower-montane forests and cloudforests. Its breeding season is in the rainy (December–April), but peaks in February–April. Males vocalize to advertise themselves and defend territories in which females place clutches containing 6–41 eggs. Most of the time, egg clutches are placed on the underside of leaves and ferns 70–500 cm directly above streams, or under dead leaves nearby streams. Males are polygynous and exhibit high site fidelity; some males were observed simultaneously guarding egg clutches at di fferent stages of development. Parental care by males was demonstrated experimentally; unattended clutches have a significantly lower eclosion rate than attended clutches, and prolonged hatching time. Clutch mortality was mainly because of desiccation, predation, and parasitism. Embryos develop for 17–27 days, then hatching as free-swimming larvae. At Reserva Las Gralarias, the species is abundant at Lucy's Creek, but it is also found at Kathy's Creek and Santa Rosa River [88].

**Call** (Figure 52): The information shown below is from Salgado and Guayasamin [132]. Advertisement call of males of *Centrolene peristicta* consists of one short and pulsed note that resembles the sound of a cricket chirp. The note has a duration of 0.036–0.087 s ( *X* + *SE* = 0.062 + 1.07, *N* = 35) and each call has one note. The dominant frequency is at 6471–7278 Hz ( *X* + *SE* = 6878 + 177 Hz., *N* = 35). The fundamental frequency is the same as the dominant frequency; there is one (*range* = 3235–6198 Hz) or two harmonics (*range* = 6471–7278 Hz). There is no conspicuous frequency modulation during the call. The repetition rate of the advertisement call varies from two to 20 calls per minute.

**Tadpole:** Not described.

**Distribution** (Figure 53): *Centrolene peristicta* is known from the Pacific slope of the Cordillera Occidental of the Andes in Ecuador and Colombia at elevations between 1380 and 1900 m ([22,87,133,134], this work). In Ecuador, *C*. *peristicta* is known from localities in the provinces of Carchi, Pichincha, and Santo Domingo de los Tsáchilas, at elevations of 1400–1852 m, with a potential distribution of 12,603 km2. All localities are within the Western Montane Forest ecoregion.

**Conservation status:** Globally, *Centrolene peristicta* is currently listed as *Least Concern* by the IUCN [135]. In Ecuador, Arteaga et al. [87] suggested the category of *Near Threatened*. Several recent observations show that the species has a wider distribution than previously thought. Reproductive populations have been observed at Reserva Las Gralarias (December 2009–May 2018; JMG, pers. obs.), Mindo Biology Station (February 2002; December 2014; M. R. Bustamante and L. Bustamante, pers. com.), and at Río Pachijal on August 2001 (I. Tapia, pers. com.). The potential distribution of the species covers an area of 12,603 km2, 44.6% of which is a ffected by human activities. At Reserva Las Gralarias, the species is infected by the amphibian chytrid fungus *Batrachochytrium dendrobatidis* [92], but populations look healthy and there is no evidence of declines. The conservation category of *Least Concern* seems too optimistic, given the fragmented distribution of the species. We sugges<sup>t</sup> that the species should be considered as *Near Threatened*, in agreemen<sup>t</sup> with Arteaga et al. (2013).

**Figure 52.** Call of *Centrolene peristicta* (LBE-C-047), recorded at Reserva Las Gralarias, 1800 m, Pichincha province, Ecuador. (**A**) Series of calls. (**B**) Single, pulsed call.

**Figure 53.** Distribution of *Centrolene peristicta* in Ecuador (yellow dots).

**Evolutionary relationships** (Figure 24): Molecular evidence places *Centrolene peristicta* and *C*. *antioquiensis* as sister species.

**Taxonomic Remarks:** *Centrolene antioquiensis* and *C*. *peristicta* are almost identical in morphology and could represent a single species. The subtle differences found between these two species might be a consequence of incomplete sampling or intraspecific geographic variation. The analysis of calls from *C*. *antioquiensis* is critical to assess the validity of the species status of *C*. *peristicta*.

**Specimens examined:** *Centrolene peristicta:* Ecuador: *Provincia de Carchi:* Maldonado (0.9 N, 78.1 W, 1410 m), KU 178137–44, 178145–51, 180325; *Provincia de Pichincha:* Bosque Protector Río Guajalito (0.233 S, 78.817 W; 1900 m), QCAZ 6446; stream near Mindo Biology Station (0.07805 S, 78.7319 W; 1600 m), QCAZ 22757–59, 22312–14; Tandapi (0.416389 S, 78.7989 W, 1520 m), KU 118051–52, 121053; 1.6 km W of Tandapi (0.40472 S, 78.8058 W; 1400 m), KU 178152; 5 km W of Tandapi on the Tandapi–Atenas road (0.3954 S, 78.8326 W; 1670 m), QCAZ 15901, 15922; Río Pachijal (0.2 S, 78.75 W; 1740 m), QCAZ 16316; Reserva Las Gralarias (0.00806 S, 78.72443 W; 1852 m), QCAZ 47298. *Provincia de Santo Domingo de los Tsáchilas:* Río Faisanes, ca. 15 km NE of La Palma, on the Quito–Chiriboga–Santo Domingo road (0.3167 S, 78.817 W; 1380 m), USNM 286714. Colombia: *Departamento de Nariño*, Reserva La Planada, ICN 12114 (PR 7871).

*Centrolene pipilata* (Lynch and Duellman 1973 [22]; Figures 54–56).

*Centrolenella pipilata* Lynch and Duellman, 1973 [22]. Holotype: KU 143278. Type locality: "16.5 km NNE of Santa Rosa, 1700 m, on Quito-Lago Agrio road, Provincia de Napo, Ecuador". *Centrolene pipilatum*—Ruiz-Carranza and Lynch, 1991 [6]. *Centrolenepipilata*—Barrio-Amorós,Rojas-Runjaic,andSeñaris,2019[85].

**Commonnames:**English:PeepingGlassfrog.Spanish:RanadeCristalPiadora.

 **Etymology:** The specific epithet *pipilata* is an adjectival derivative of the Latin verb *pipila*, meaning to peep, and refers to the call of this centrolenid frog [22].

**Identification:** *Centrolene pipilata* is distinguished from most glassfrogs by having, in life, a green dorsum with yellowish–white flecks and di ffuse dark green marks, and a distinct prepollex (Figures 54 and 55). On the Amazonian slopes of Ecuador, the only species that have a similar dorsal color pattern are *C*. *sanchezi* and *C*. *huilensis*. *Centrolene sanchezi* has a short, thin humeral spine (large, wide, and usually with the tip projected anteriorly in *C*. *pipilata*) and lacks a distinct prepollex. *Centrolene huilensis*is conspicuously larger than *C*. *pipilata* (in *C*. *huilensis*, male SVL = 23.6–26.7 mm; SVL = 28.7 mm in 1 female; in *C*. *pipilata*, male SVL 19.7–22.6 mm, female SVL 22.6–23.6 mm). *Nymphargus truebae*, a species only known from the Andes of southern Peru, has the same color pattern as *C*. *pipilata*, however, males in *N*. *truebae* lack humeral spines (present in males of *C*. *pipilata*). The Colombian *Centrolene hybrida* has a white gastrointestinal peritoneum (iridophores absent in *C*. *pipilata*), and lacks a distinct prepollex.

**Figure 54.** *Centrolene pipilata*. (**Top row**): Coloration in life, adult male, holotype, KU 143278; photos by William E. Duellman. (**Bottom row**): Coloration in preservative of KU 178155 (**left**) and KU 143287 (**right**); photos by Juan M. Guayasamin.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout round in dorsal aspect, round or truncated in lateral profile (Figure 55); (3) tympanum moderate, oriented almost vertically, with slight lateral and posterior inclinations, its diameter 31.0%–39.4% of eye diameter; tympanic annulus completely visible; supratympanic fold evident; tympanic membrane translucent, partially pigmented, clearly di fferentiated from surrounding skin; (4) dorsal skin shagreen, males with spicules coinciding with white spots; (5) pair of enlarged subcloacal warts (Figure 15); (6) anterior 50%–60% of the ventral parietal peritoneum white, posterior portion transparent (condition P2–P3); white pericardium; no iridophores in peritonea covering intestines, stomach, kidneys, gall bladder, and urinary bladder (condition V1); (7) liver with four clearly defined lobes, lacking iridophores (condition H0); (8) males with conspicuous humeral spines; (9) webbing absent between Fingers I, II, and III; moderate between outer fingers (Figure 55); webbing formula: III (2<sup>+</sup>–21/2)—(2–2<sup>+</sup>) IV; (10) webbing between toes extensive; webbing formula on foot: I (1<sup>+</sup>–2−)—(2–2<sup>+</sup>) II (1–1<sup>1</sup>/3)—(2–21/3) III (1–1<sup>1</sup>/2)—2+ IV (2–2<sup>1</sup>/2)—(1–11/2) V; (11) white ulnar fold present; outer tarsal margin with low white tubercles: Internal tarsal fold low; (12) exposed prepollex; nuptial pad Type II; (13) Finger I as long as Finger II or slightly shorter (Finger I 90.0%–100% of Finger II); (14) disc of Finger III width 44.8%–56.5% of eye diameter; (15) in life, dorsum green with small di ffuse black spots and yellowish–white flecks (Figure 54); green bones; (16) in preservative, dorsum lavender with numerous minute white spots and larger dark lavender spots (Figure 54); (17) in life, iris greyish–white with thin black reticulations and a yellow hue around pupil; (18) melanophores mostly absent from fingers and toes, except for a few on proximal half of Finger IV, and along Toes IV and V; (19) males call from upper side of leaves; call undescribed; (20) fighting behavior unknown; (21) eggs placed on upper surface of leaves; parental care unknown; (22) tadpoles unknown; (23) minute body size; in adult males, SVL 19.7–22.6 mm ( X = 21.5, *n* = 9); in two adult females SVL 22.6–23.6 mm.

**Color in life** (Figure 54): Dorsum green with minute yellowish–white flecks and larger, di ffuse black spots; upper lip white; region below eye with small white warts that are spiculated in males; bones green; upper flanks green with minute white spots; lower flanks whitish cream; ulnar fold with thin white line; outer tarsal fold with low white tubercles; small white tubercles just posterior to cloaca; iris greyish–white with thin black reticulations and a yellow hue around pupil.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs lavender with small white spots and larger dark lavender spots (Figure 54); margin of upper lip white; region below eye with small, spiculated white warts; white tubercles just posterior to cloaca. White parietal peritoneum covers anterior 50%–60% of venter; white pericardium; no iridophores in peritonea covering digestive tract, liver, kidneys, gall bladder, and urinary bladder.

**Variation:** In preservative, some individuals lack dorsal dark spots.

**Figure 55.** *Centrolene pipilata*, KU 143278, holotype. ( **A**) Head in lateral view. (**B**) Hand in ventral view. Illustrations by Juan M. Guayasamin.

**Biology and ecology:** All individuals have been found at night on vegetation along cascading mountain streams. In captivity, a female deposited a clutch of 18 eggs. Males call from the upper side of leaves. At Río Azuela, *Centrolene pipilata* was found in sympatry with *Nymphargus anomalus*, *N*. *megacheirus*, *N*. *siren*, and *Hyalinobatrachium pellucidum*. At 16.5 NNE of Santa Rosa, the species was found with *Espadarana audax*, *N*. *megacheirus*, and *N*. *siren* [22]. Eggs have clear jelly and pale green yolks [22]. Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 56): *Centrolene pipilata* is endemic to the cloud forest on the Amazonian slope of the Ecuadorian Andes at elevations between 1300–1910 m ([17,22], this work). The species has been recorded from four localities in the provinces of Napo and Sucumbíos. The habitat of the species is within the Eastern Montane Forest region.

**Figure 56.** Distribution of *Centrolene pipilata* in Ecuador (yellow dots).

**Conservation status:** Globally, *Centrolene pipilata* is listed by the IUCN as *Endangered* [136] because of its limited distribution and the continuing decline in the extent and quality of suitable habitat. The last confirmed report of *C*. *pipilata* was at 14.7 km NE of Río Salado on February 1979 [17]. Recent surveys at Río Azuela [91] and the type locality (JMG, pers. obs.) have failed to find the species. In Ecuador, based on the IUCN criteria A2c,e, we sugges<sup>t</sup> that the species should be considered as *Critically Endangered*.

**Evolutionary relationships** (Figure 24): Given the current taxon and gene sampling, *Centrolene pipilata* and *C*. *hybrida* are sister species.

**Specimens examined:** *Centrolene pipilata:* Ecuador: *Provincia de Napo:* 16.5 km NNE Santa Rosa (0.2186 S, 77.732 W, 1700 m), KU 143278–83, 143554; 3.2 km NNE Oritoyacu (0.4597 S, 77.8672 W; 1910 m), KU 178153; Río Azuela (0.11667 S, 77.61667 W; 1740 m), KU 143284–87, 155498, 166331; Río Salado, 1 km upstream from Río Coca (0.19167 S, 77.6997 W; 1420 m), KU 178154–55.

### *Centrolene sanchezi* Ruiz-Carranza and Lynch, 1991 [129] (Figures 57–61).

*Centrolene sanchezi* Ruiz-Carranza and Lynch, 1991 [129]. Holotype: ICN 24293.


*Centrolene bacata*—Barrio-Amorós, Rojas-Runjaic, and Señaris, 2019 [85].

**Common names:** English: Sánchez's Glassfrog. Spanish: Rana de cristal de Sánchez.

**Etymology:** The specific epithet *sanchezi* honors Ricardo Sánchez, who, with John D. Lynch, discovered the species.

**Identification:** *Centrolene sanchezi* is easily distinguished from other glassfrogs by its minute body size (SVL 18.9–22.3 mm), green dorsum with small white spots, and the presence of white warts in an area that extends from below the eye to the insertion of the arm (Figures 57 and 58A). Additionally, adult males have conspicuous humeral spines. The only other centrolenid from eastern Ecuador that can be confused with *Centrolene sanchezi* is *Centrolene pipilata*, which has a green dorsum with small, diffuse black spots and yellowish–white flecks (Figure 54), and a distinct prepollex. Also, the two species occupy different elevations on the Amazonian slope of the Andes; *C*. *sanchezi* is found at elevations of 1950–2350 m, whereas *C*. *pipilata* occurs at 1420–1910 m.

**Diagnosis:** (1) Vomerine teeth absent; (2) snout rounded in dorsal aspect, bluntly rounded or truncated in lateral profile (Figure 58); (3) tympanum oriented almost vertically, its diameter 31.4%–37.8% of eye diameter; tympanic annulus visible except for dorsal border covered by supratympanic fold; tympanic membrane partially pigmented, but differentiated from surrounding skin; (4) dorsal surfaces shagreen, with small spicules evident in most males; (5) pair of enlarged subcloacal warts (Figure 15); (6) anterior half of ventral parietal peritoneum white, posterior half translucent (condition P2); white pericardium, translucent visceral peritoneum (condition V1); (7) liver tetralobed, lacking iridophores (condition H0); (8) humeral spines present in adult males; (9) no webbing between Fingers I and II; webbing formula on hand: II 2—31/<sup>3</sup> III 21/2—21/<sup>4</sup> IV (Figure 58); (10) webbing formula on foot: I 11/2—2<sup>+</sup> II (1–1<sup>1</sup>/2)—(2–21/3) III 1<sup>+</sup>—(2<sup>+</sup>–21/4) IV 21/2—(1–1<sup>+</sup>) V (Figure 58); (11) ulnar and inner tarsal folds low or absent; outer tarsal fold absent; (12) nuptial pad Type I, concealed prepollex; (13) Fingers I and II about equal in length (FII/FI = 91.3%–104.5%); (14) disc of Finger III of moderate size, about 40.3%–50.6% of eye diameter; (15) in life, dorsum dark green with small white spots (Figure 57); conspicuous white warts below the eye and tympanum; upper lip white; bones green; (16) in preservative, dorsum lavender with small white spots; (17) iris pale bronze with black reticulation; (18) melanophores mostly absent from fingers and toes, except for a few on Toes IV and V and on base of outer fingers; (19) males call from the upper side of leaves; calls are produced in series; each call has one or two notes, with a duration of 6–21 ms (mean = 11, SD = 2.8); dominant frequency at peak amplitude is 5719–6188 Hz (mean = 5996, SD = 131); (20) fighting behavior unknown; (21) eggs deposited on upper side of leaves; short-term maternal care unknown; parental care by males absent; (22) tadpoles unknown; (23) minute body size, SVL 18.9–22.3 mm in males (*n* = 15); 20.9 mm in one female.

**Figure 57.** *Centrolene sanchezi* in life. (**A**,**B**) Male and female from Yanayacu Biological Station, Ecuador. (**C**) Holotype of *C*. *sanchezi*, ICN 24293. (**D**) Paratype of *C*. *guanacarum*, ICN 11685. Photos by A. Arteaga (**A**,**B**) and P. Ruiz-Carranza (**C**,**D**).

**Figure 58.** *Centrolene sanchezi*, KU 202803 (holotype of *Centrolene bacata)*. (**A**) Head in lateral view. (**B**) Head in dorsal view. (**C**) Hand in ventral view. (**D**) Foot in ventral view. Drawings not to scale. Modified from Wild [137].

**Color in life** (Figure 57): Based on field notes by W. E. Duellman (4 March 1984), reported in Wild [137], and observation by authors. Dorsum dark green; series of white–cream tubercles under eye; throat and ventral surfaces of limbs green; digits pale green; cloacal region with white warts; ventral parietal peritoneum white anteriorly and translucent posteriorly; visceral peritoneum translucent; bones green; iris pale bronze with thin black reticulation.

**Color in ethanol** (Figure 59): Dorsal surfaces lavender with small, unpigmented spots and white warts; limbs cream lavender with numerous small, unpigmented spots and few white warts; white warts on lateral surface of head; upper lip white; tympanum pigmented with purple specks; cloacal region with white or cream warts; iris silvery white with dark purple reticulation. Fingers I and II and Toes I–III dorsally unpigmented; some pigmentation visible on Fingers III and IV, and Toes IV and V. White parietal peritoneum covering anterior half of venter, posterior half translucent; silvery-white pericardium; translucent peritonea covering liver, gastrointestinal track, and renal capsules (dissected male: QCAZ 22386).

**Figure 59.** Types of *Centrolene sanchezi* and *C*. *bacata* in preservative. (**A**,**B**) *Centrolene bacata*, holotype KU 202803. (**C**,**D**) *C*. *sanchezi*, paratype ICN 24294. Photos by Juan M. Guayasamin.

**Biology and ecology:** During the night, *Centrolene sanchezi* has been found on leaves approximately 130–300 cm above streams in primary and secondary forest. Males call from the upper sides of leaves. A male (QCAZ 22728) was found nearby two egg clutches, one on the underside and the other on the upper side of a single leaf; egg clutches have 14–18 pale yellowish–green eggs. During February 2013, males have been heard calling at Yanayacu Biological Station. At this locality, *Centrolene sanchezi* (reported as *C*. *bacata*) is the most abundant centrolenid; other sympatric species at Yanayacu include *Centrolene* <sup>a</sup>ff. *buckleyi*, *C*. *huilensis*, *Nymphargus posadae*, *N*. *siren*, and *N*. *wileyi* [20,125]. Short-term maternal care unknown; parental care by males absent ([25], as *C*. *bacata*).

**Call** (Figure 60): We analyzed 63 notes contained within 11 calls from 1 individual (LBE-C-023). Calls are produced in series, which can be relatively long (range = 618–3085 ms, mean = 1462.6 ms, SD = 639.5 ms); each can have one or two notes; note repetition rate is 3–9 (mean = 5.7, SD = 1.6) notes per call. Notes sometimes occur in pairs (i.e., much shorter note interval than compared to the rest of the call). Each call is very short, with a duration of 6–21 (mean = 11, SD = 2.8) ms. Notes are strongly pulsed and have one or two (mean = 1.4, SD = 0.5) amplitude peaks throughout the note, where the second amplitude peak is generally weaker than the first. Pulses within a note have a rate of 83–200 (mean = 122, SD = 29) pulses per second. Notes have their peak amplitude in the first 50% of the note (relative peak time: Range = 0.0987–0.5497, mean = 0.216, SD = 0.076). The dominant frequency measured at peak amplitude is 5719–6188 (mean = 5996, SD = 131) Hz and is contained within the fundamental frequency. The fundamental frequency has a lower limit of 4688–6000 (mean = 5714, SD = 222) Hz and a higher limit of 5906–6656 (mean = 6268, SD = 144) Hz.

**Figure 60.** Call of *Centrolene sanchezi* (LBE-C-023), recorded at Reserva Yanayacu, 2150 m, Napo province, Ecuador. (**A**) Series of calls. (**B**) Single, pulsed call.

**Tadpole:** Not described.

**Distribution** (Figure 61): *Centrolene sanchezi* is known from a few localities on the Amazonian slopes of the Andes of Ecuador and Colombia and from one locality on the eastern slope of the Colombian Cordillera Central (see Specimens Examined), at elevations between 1800 and 2350 m [20,101,129,137]. In Ecuador, the habitat of the species is within the Eastern Montane Forest region.

**Figure 61.** Distribution of *Centrolene sanchezi* in Ecuador (yellow dots).

**Conservation status:** Globally, *Centrolene sanchezi* is listed as *Data Deficient* by the IUCN [138], but the evaluation does not account for the synonym presented herein. The habitat of the species is fragmented by agriculture and pastureland and threatened by mining (mainly in southern Ecuador). We sugges<sup>t</sup> that the species should be considered as *Endangered*, following IUCN criteria B2, Ba, Bb(iii).

**Evolutionary relationships** (Figure 24): *Centrolene sanchezi* is the sister species to a clade formed by *C*. *pipilata* plus *C*. *hybrida*. These three species are found on the Amazonian slopes of the Andes.

**Taxonomic Remarks:** Examination of the type material of *Centrolene sanchezi*, *C*. *bacata*, and *C*. *guanacarum* reveals no morphological differences among them. Moreover, all specimens share two distinctive traits; a small, laminar humeral spine in males and the presence of a lateral row of enameled warts that extends from below the eye to just posterior to the insertion of the arm (Figure 51). Therefore, we place *Centrolene bacatum* Wild 1994 [137] and *Centrolene guanacarum* Ruiz-Carranza and Lynch 1995 [26] under the synonymy of *Centrolene sanchezi* Ruiz-Carranza and Lynch, 1991 [129].

**Specimens examined:** *Centrolene sanchezi:* Ecuador: *Provincia de Morona Santiago:* 11.2 km WSW Plan de Milagro (03◦07- S, 78◦30- W; 2350 m), KU 202803 (holotype of *Centrolene bacata*), 202804, 202807–12 (paratypes of *C*. *bacata*); *Provincia de Napo:* Yanayacu Biological Station (0◦41- S, 77◦53- W; 2100 m), QCAZ 16212, 17807, 22386–87, 22728, 26025–27, 26056, 27438. Colombia: *Departamento de Caquetá:* Municipio de Florencia, Vereda Gabinete, 3.1 Km por carretera abajo del Alto Gabinete, vertiente oriental, Cordillera Oriental, 1◦4- latitud N, 75◦4- W de Greenwich, 2190 m, ICN 24293 (holotype of *C*. *sanchezi*); *Departamento del Cauca:* Municipio de Inzá, Km. 84 carretera Popayán a Inzá, Río Guanacas, quebrada afluente, Internado Indígena Río Guanacas, vertiente oriental Cordillera Central, 2◦34- Latitud N, 76◦05- W de Greenwich, 1800–1900 m, ICN 11685 (paratype of *C*. *guanacarum*). **Genus** *Chimerella* Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, & Vilà, 2009 [1].

**Etymology:** The name *Chimerella* comes from the Greek *Khímaira* and the suffix–*ella* diminutive. In Greek mythology, the Chimera is a creature composed of parts of multiple animals; the name refers to the peculiar combination of morphological characteristics of *Chimerella mariaelenae* [1].

*Chimerella mariaelenae* (Cisneros-Heredia and McDiarmid, 2006 [139]; Figures 62–65).

	- Podocarpus National Park (ca. 04◦15- S, 78◦56- W, 1820 m), on the western slope of Contrafuerte de Tzunantza, Cordillera Oriental, eastern slopes of the Andes, Provincia de Zamora Chinchipe, Republic of Ecuador".

*Cochranella parabambae* (in part)—Goin, 1961 [97].

*Chimerella mariaelenae*—Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, and Vilà, 2009 [1].

**Common names:** English: María Elena's Glassfrog. Spanish: Rana de Cristal de María Elena.

**Etymology:** The specific epithet *mariaelenae* is a noun in the genitive case and a patronym for María Elena Heredia, Diego F. Cisneros-Heredia's mother [139].

**Identification:** *Chimerella mariaelenae* (Figures 62 and 63), *Chimerella corleone*, and *Vitreorana gorzulae* are the only known glassfrog that have the following combination of traits: Humeral spine in adult males, transparent ventral parietal peritoneum, and white pericardial, hepatic, and visceral peritonea. *Vitreorana gorzulae*, an endemic to the Guiana Shield, is distinguished by lacking dark spots on the dorsum (present in *C*. *mariaelenae*). *Chimerella mariaelenae* is easily differentiated from the Peruvian *C*. *corleone* by having an orange to reddish iris (silvery white in *C*. *corleone*) and a dorsum with small black dots (dorsum with yellow dots in *C*. *corleone* [19]). On the Amazonian slopes of Ecuador, only species in the genus *Hyalinobatrachium* and *Teratohyla amelie* could be confused with *C*. *mariaelenae*. However, these species lack the small dark spots that characterize the dorsum of *C*. *mariaelenae*. Additionally, adult males of *C*. *mariaelenae* have small humeral spines, which are absent in all *Hyalinobatrachium* species and *Teratohyla amelie*.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout truncated in dorsal aspect, truncated to slightly protruding in lateral profile (Figure 63); (3) tympanum oriented posterolaterally with slight dorsal inclination, its diameter 23.0%–27.4% of eye diameter; tympanic membrane translucent, clearly differentiated from surrounding skin; (4) dorsal skin shagreen; males with minute dorsal spicules (only visible under magnification ×250); (5) pair of enlarged subcloacal warts; (6) ventral parietal peritoneum transparent (condition P0); iridophores in pericardium and peritonea covering digestive tract and testes; kidneys and urinary bladder lacking iridophores (condition V5); (7) liver bulbous and covered by iridophores (condition H2); (8) males with small humeral spines; (9) webbing absent or basal between inner fingers, moderate between outer fingers (Figure 63); webbing formula IV (2<sup>+</sup>–21/2)—(2<sup>+</sup>–21/2) V; (10) webbing between toes extensive; webbing formula on foot I (1<sup>1</sup>/2–2−)—(2–2<sup>1</sup>/2) II 1—2<sup>+</sup> III (1<sup>+</sup>–11/2)—(3−–3<sup>+</sup>) IV (2<sup>1</sup>/2–31/4)—(1<sup>+</sup>–11/4) V; (11) ulnar and tarsal folds absent; (12) concealed prepollex; nuptial pad Type I; (13) Finger I slightly longer than Finger II (Finger II length 95.6%–97.3% of Finger I); (14) disc of Finger III relatively narrow, its width about 22.3%–25.7% of eye diameter; (15) in life, dorsum yellowish green with small grey to black spots (Figure 62); bones light green; (16) in preservative, dorsum pale lavender with small, dark, lavender spots; (17) in life, iris white with abundant dark flecks, pupil surrounded by a rufous to orange ring, outlined by a dark grey to blue ring (Figure 62); (18) dorsal surfaces of fingers and toes lacking melanophores, except for some on base of Toe V; (19) males

call from the upper surface of leaves; typical call with two notes; each note extremely short at 4–7 (mean = 6, SD = 0.9) ms; dominant frequency measured at peak amplitude is 6718–8010 (mean = 7510, SD = 408) Hz; (20) fighting behavior unknown; (21) eggs laid on the upper surfaces of leaves; short-term maternal care present; parental care by males absent; (22) tadpoles with elongated body, head with dorsoventral compression; labial tooth row formula 2(1)(2)/3; (23) minute body size; adult males, SVL 17.9–19.7 mm (*n* = 3); one adult female SVL 20.8 mm.

**Color in life** (Figure 62): Dorsal surfaces yellowish green, with several grey to black spots. No white coloration is evident on upper lip or ulnar and tarsal regions. Iris bicolored, with dark brown circumpupilar area separated from pupil by orange to red–brown ring; whitish external background with abundant dark flecks on uppermost and lowermost portions of iris. Venter completely transparent (clear ventral parietal peritoneum), showing white heart (covered by white pericardium); white digestive tract, liver, testes, and gall bladder. Transparent peritoneum covering urinary bladder. Bones white to light green.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs pale lavender, with small dark lavender spots. Ventral parietal peritoneum and kidneys lack iridophores. White pericardial, hepatic, and visceral peritonea.

**Figure 62.** *Chimerella mariaelenae* in life. (**Top row**): Adult male from Bigal River Biological Station, 931 m, Ecuador; photos by Ross Maynard. (**Bottom row**): Amplectant pair, Quebrada Pangayaku, 930 m, Napo province, Ecuador, MZUTI 1680–81; photo by Eduardo Toral.

**Figure 63.** *Chimerella mariaelenae*. (**A**) Head in lateral view, QCAZ 31729. (**B**) Head in dorsal view, QCAZ 21252. (**C**) Hand in ventral view, QCAZ 22363. Illustrations by Juan M. Guayasamin.

**Biology and ecology:** The biology of *Chimerella mariaelenae* is poorly known. All individuals have been found on the upper surfaces of leaves along small streams and ditches in cloud forest. At the type locality, *C*. *mariaelenae* is sympatric with *Nymphargus cochranae* and *Hyloscirtus phyllognathus* [139]. Reproductively active individuals have been observed nearby Cascada San Rafael in Quebrada Pangayacu in August 2012. Short-term maternal care present; parental care by males absent [25].

**Tadpole:** The following information is summarized from Terán-Valdez and Guayasamin [140] and is based on a specimen at Gosner Stage 39 raised in laboratory conditions. Body elongated and oval-depressed, wider (body width = 5.1 mm) than higher (body height = 3.8 mm). Chondrictial elements not visible. Snout rounded in dorsal and lateral views. Lateral line system visible, formed by several stitches parallel or perpendicular to longitudinal axis of body. Short, single, sinistral spiracle located at posterolateral region of body; spiracular aperture with dorsoposterior orientation, with inner wall present as a low ridge. Vent tube short and abdominal, free posteriorly, opening directed posteriorly. Tail long, with subacute tip. Myotomes visible throughout length of tail; straight medial line visible, separating dorsal and ventral myotomes. Dorsal fin originating at about mid-length of tail; height relatively uniform until distal end, where it decreases abruptly. Proximally, ventral fin originating at base of tail muscle, reaching its maximum height after mid-length of tail. Oral disc non-emarginated and surrounded by 49 marginal uniserial papillae. Only ventral and lateral papillae present, lacking dorsal papillae. Upper and lower jaw sheaths nearly straight and fully keratinized, with serrated edge. Labial tooth row formula 2(1)(2)/3; gap in tooth row A-1 could be artificial (because of teeth loss). In life, dorsally, tadpoles at Stage 39 brownish with two areas without pigmentation at anterolateral border of eye. Reddish coloration visible on anterior half of dorsum because of its transparency. Anterior-most part of head is grey. Iridophore aggregations present on dorsum, along vertebral column. Iris bronze. Ontogenetic variation is provided in Terán-Valdez and Guayasamin [140].

**Call** (Figure 64): We analyzed 26 notes contained within six calls from one individual (LBE-C-021). The typical advertisement call is variable from being short to a moderate length call (range=231–1761 ms, mean = 679 ms, SD = 623.3 ms). The typical call has two notes, but some call can have up to 10 (mean = 4.3, SD = 3.4) notes. Each note is extremely short at 4–7 (mean = 6, SD = 0.9) ms. Notes have a single amplitude peak. The dominant frequency measured at peak amplitude is 6718–8010 (mean = 7510, SD = 408) Hz and is contained within the fundamental frequency. The fundamental frequency has a lower limit of 6460–7752 (mean = 7222, SD = 387) Hz and a higher limit of 7063–8441 (mean = 7828, SD = 398) Hz.

**Figure 64.** Call of *Chimerella mariaelenae* (LBE-C-021), recorded at Pangayaku creek, 929 m, Napo province, Ecuador. (**A**) Series of calls. (**B**) Single, pulsed note.

**Distribution** (Figure 65): *Chimerella mariaelenae* occurs on the Amazonian slopes of the Ecuadorian Andes at elevations between 813 and 1820 m ([139,141,142], this work), and was recently reported from Peru [143]. The species is known from a few localities in the provinces of Napo, Morona Santiago, Orellana, Tungurahua, and Zamora Chinchipe (Specimens Examined). The potential distribution of the species is 25,472 km<sup>2</sup> within the Eastern Foothill and Montane Forest regions.

**Figure 65.** Distribution of *Chimerella mariaelenae* in Ecuador (yellow dots).

**Conservation status:** Globally, *Chimerella mariaelenae* is currently listed as *Least Concern* by the IUCN [144]. The species is threatened by human activities, although it tolerates some habitat disturbance and has a relatively large distribution, including populations within protected areas (i.e., Bigal River Biological Station, Parque Nacional Podocarpus, Parque Nacional Sangay, Reserva Ecológica Cayambe-Coca, Reserva Narupa). In Ecuador, major threats for the species include deforestation and mining; thus, we sugges<sup>t</sup> that the species should be considered as *Near Threatened* at the local level.

**Evolutionary relationships** (Figures 19 and 66): In the original description and based on morphological data, *Chimerella mariaelenae* was assigned to the *Centrolene gorzulai* group, which, otherwise, consisted of species from the Guiana Shield. Under this hypothesis, the distribution of *C*. *mariaelenae* was thought to support a biogeographical connection between the Andes and the Guiana Shield [139]. Subsequently, Cisneros-Heredia and McDiarmid [17], based on differences in the chromatophore organization, considered that the relationships of *C*. *mariaelenae* were uncertain. Phylogenetic trees based on molecular data show that *Chimerella mariaelenae* and *C*. *corleone* are sister species [19] (Figure 66); both species are restricted to the Amazonian versant of the Andes, with no close relationship with species from the Guiana Shield. The genus *Chimerella* is herein inferred as sister to *Espadarana* (Figure 19), but with low nodal support.

**Figure 66.** Evolutionary relationships between species in the genus *Chimerella*, inferred using maximum likelihood and Bayesian criteria.

**Remarks:** One of the specimens reported by Goin [97] as *Centrolenella parabambae* from the Topo River is an adult female *Chimerella mariaelenae*.

**Specimens examined:** *Chimerella mariaelenae:* Ecuador: *Provincia de Napo:* Río Hollín (ca. 00◦58- S, 77◦45- W, ca. 1400 m), QCAZ 18618–19, 22363. Quebrada Pangayaku (0.78255 S, 77.791 W, 929 m), MZUTI 1680–81; Reserva Narupa (0.684 S, 77.741 W, 1179–1208 m), ZSFQ 0437–38; Reserva Narupa (0.671 S, 77.774 W, 1502 m), ZSFQ 0439–41. *Provincia de Morona Santiago:* 6.7 km W of 9 de Octubre (02◦13-30.5" S, 78◦17-25.6" W, 1715 m), QCAZ 32643. *Provincia de Tungurahua:* near Río Negro (01◦24- S, 78◦15- W, 1423 m), on the Río Negro–Río Verde road, QCAZ 21252, 31729; Río Topo, BM 1912.11.1.69. *Provincia de Zamora Chinchipe:* stream tributary of the Río Jambue (ca. 4◦15- S, 78◦56- W, ca. 1820 m), ca. 16 km S from Zamora, Parque Nacional Podocarpus, DFCH-USFQ D125; gravel road E to Sarsa (3.80783976 S, 78.60593076 W; 1500 m), QCAZ 47053.

**Photographic records:** *Provincia de Orellana:* Reserva Río Bigal (0.532913◦ S, 77.423228◦ W; 981 m). Photo by Morley Read.

## **Genus** *Cochranella* Taylor, 1951 [15].

**Etymology:** Named in honor of Doris M. Cochran, herpetologist and curator of the Smithsonian Institution, Washington, D.C.

*Cochranella granulosa* (Taylor, 1949 [145]; Figures 67 and 68).

*Centrolenella granulosa* Taylor, 1949 [145]. Holotype: FMNH 178269.

Type locality: "Los Diamantes, one mile south of Guápiles, (Cantón de Pococí, Provincia Limón), Costa Rica".

*Cochranella granulosa*—Taylor, 1951 [15].

**Common names:** English: Granular Glassfrog. Spanish: Rana de Cristal Granulosa.

**Etymology:** The specific epithet *granulosa* comes from the Latin word *granum* (grain), in allusion to the granular dorsal skin texture of the species.

**Identification:** *Cochranella granulosa* can be distinguished from most glassfrogs by having a dark green to bluish–green dorsum with whitish granules and faint to well-defined dark spots (Figure 67), yellowish–green hands and feet, a reduced white ventral parietal peritoneum (which covers less than 40% of the venter), white digestive tract, and by lacking humeral spines. Another distinctive trait is the granular texture of its dorsal skin. Species that might have a sympatric distribution with *C*. *granulosa* and that present a similar dorsal coloration include *Cochranella litoralis* and some populations of *Espadarana prosoblepon*. However, *C*. *granulosa* has a granular skin (shagreen in *E*. *prosoblepon* and *C*. *litoralis*) and lacks humeral spines (present in males of *E*. *prosoblepon* and *C*. *litoralis*). Also, *C*. *litoralis* has an orange to red iris (yellow to golden in *C*. *granulosa*) and *E*. *prosoblepon* has a digestive tract that lacks white iridophores (present in *C*. *granulosa*).

**Figure 67.** *Cochranella granulosa* in life. (**Top row**): Reserva Guayacán, Costa Rica; photos by Brian Kubicki. (**Bottom row**): Reserva El Jardín de los Sueños, 461 m, Cotopaxi province, Ecuador, MZUTI 4811; photos by Jaime Culebras (dorsolateral) and Javier Aznar (ventral).

**Diagnosis:** (1) Vomerine teeth present, each vomer with one to three teeth; (2) snout truncated in dorsal aspect, slightly sloping in lateral profile; (3) tympanum oriented dorsolaterally, with slight posterior inclination, its diameter about 35% of eye diameter; tympanic annulus visible, low supratympanic fold evident, tympanic membrane partially pigmented and clearly differentiated from surrounding skin; (4) dorsum granular, lacking spicules; (5) venter areolate, lacking pair of enlarged subcloacal warts; (6) white lining on the anterior 20%–40% of the ventral parietal peritoneum (condition P1); peritoneum with white iridophores covering digestive tract (condition V2); (7) liver tetralobed; hepatic peritoneum lacking iridophores (condition H0); (8) humeral spines absent; (9) webbing absent between Fingers I–II, vestigial to moderate between Fingers II–III, extensive between III–IV; webbing formula on hand: II (1+–2)—(3–3+) III (1<sup>1</sup>/3–21/2)—(1–2+) IV; (10) webbing on foot extensive, formula: I 0<sup>+</sup>—11/<sup>2</sup> II 0<sup>+</sup>—12/<sup>3</sup> III (1–1<sup>+</sup>)—2− IV 2—1− V; (11) ulnar fold low; inner and outer tarsal folds low and thin; (12) concealed prepollex; nuptial pad Type I; (13) Finger I about same length as Finger II; (14) disc of Finger III narrow, about 40% of eye diameter; (15) in life, dorsum dark green to bluish green with fine, yellowish–white granules and faint to clearly marked dark spots (Figure 67); bones green; (16) in preservative, dorsum cream to light lavender with faint to clearly defined dark spots; (17) iris yellow to golden with few, faint brownish streaks; (18) melanophores absent from dorsal surfaces of hands and toes; (19) males call from the upper side of leaves; each call consists of three to five notes, the first note being slightly longer that the others; average duration for a three-note call is 0.8–0.9 s; dominant frequency at 3700 Hz; (20) males fight venter to venter, hanging from the vegetation by the tips of their toes; (21) dark egg are placed on the upper sides of leaves; short-term maternal care present; parental care by males absent; (22) tadpoles with a flattened body and slender tail; tadpole in early stages have a brown pigmentations that is lost in older, red larvae; jaw sheaths are strongly serrated, the upper jaw forms a smooth M-shape; the lower sheath is V-shaped; labial tooth row formula 2/3; (23) medium body size; in males, SVL 22.5–29.0 mm; in females, SVL 29–32 mm.

**Color in life** (Figure 67): Dorsal surfaces green to bluish green, with yellowish or bluish granules and faint to clearly marked blue to black spots. Anterior 20%–40% of venter white, posterior portion translucent. Iris yellow to golden. Pale or whitish lip in populations from Central America (Savage, 2002; Kubicki, 2007). One individual from Ecuador (MZUTI 4811) lacks dark dorsal spots.

**Color in ethanol:** Dorsum cream to light lavender to dark lavender with faint to clearly marked dark spots (the Ecuadorian specimen lacks dark spots, MZUTI 4811); iridophores on the anterior 20%–40% of the ventral parietal peritoneum; pericardium and gastrointestinal peritonea with white iridophores; hepatic peritoneum lacking iridophores.

**Biology and ecology:** The information below is from Kubicki [24] and RWM (pers. obs.) *Cochranella granulosa* is active at night in riparian habitats of streams through pristine tropical forest, but also sometimes along streams in disturbed areas. Males fight venter to venter, hanging from the vegetation by the tips of their toes. Reproduction usually occurs in riparian habitats, but Kubicki [24] also reported reproductive activity in a small forest pond about 10 m away from the nearest stream. Egg clutches consist of 40 to 104 eggs [24,146,147] that are attached on masses on the upper distal half of a leaf overhanging the stream. Following hydration, the eggs form a pendent mass (i.e., drip tip) that hangs o ff the leaf and thereby ensures a constant flow of water over the developing larvae [4,146]. There is no long-term parental care in this species [148]. Delia et al. [25] experimentally demonstrated that females provide short-term parental care (brooding) to their egg clutches just after oviposition; this behavior, although brief (i.e., few hours), reduces embryo mortality from dehydration.

**Call:** Males often call in a chorus, with one male initiating calling and then being answered by several males. The call consists of a rapid high-pitched pulsed trill, usually composed by three notes (range three to five notes; [24]). Usually, the first note is longer (average duration = 0.3 s) than the subsequent notes (average duration = 0.15 s). The mean duration for a three-note call is 0.8–0.9 s [24]. The dominant frequency is 3700–4500 Hz [24,149,150]. In Ecuador (Los Laureles, Cotopaxi province) males were actively calling in February 2016 (Jaime Culebras, pers. obs.).

**Tadpoles:** The tadpole of *Cochranella granulosa* was first described by Starrett [146]. Ho ffmann [147] recently provided an additional and detailed description; below, we present a summary based on Ho ffmann's description. Tadpoles of *C*. *granulosa* and *C*. *euknemos* share a particular slim shape. Recently hatched tadpoles (one to five days old) are markedly slender and dark brown. In the laboratory, older tadpoles are slender with a very long tail; they lack much body pigment and their bright red coloration is from the highly vascularized internal anatomy anterior to the body cavity and visible through the body wall. The oral disc is not emarginate and is bordered laterally and along the posterior margin with 28–41 marginal papillae (mean = 33 ± 4 papillae). Jaw sheaths are strongly serrate; the upper jaw forms a smooth M-shape, and the lower sheath is V-shaped. Generally, all five tooth rows are well developed; LTRF is 2(2)/3; the A2 row has a large central gap with the two halves extending as short wings on each side of the lateral descendent ends of the upper jaw sheath.

**Distribution** (Figure 68): The distribution of *Cochranella granulosa* includes the humid lowland and premontane slopes from the Atlantic drainage of eastern Honduras to central Panama and on the Pacific versant in humid upland or gallery forests from northern Costa Rica to northern Ecuador, 40–1500 m elevation ([148,151], this work). Records of *Cochranella granulosa* in Ecuador are from El Jardín de los Sueños reserve, Cotopaxi province [151], and Durango, 238 m, Esmeraldas province.

**Conservation status:** Globally, *Cochranella granulosa* is listed as *Least Concern* by the IUCN [152]. We sugges<sup>t</sup> *Data Deficient* in Ecuador, since it is known from two recent records (Figure 68).

**Evolutionary relationships** (Figure 69): *Cochranella granulosa* is sister to *C*. *resplendens*, being an example of allopatric speciation mediated by the uplift of the Andes.

**Figure 68.** Distribution of *Cochranella granulosa* in Ecuador (yellow dot).

**Figure 69.** Evolutionary relationships among species in the genus *Cochranella*, inferred using maximum likelihood and Bayesian criteria.

**Remarks:** Karyotype of *Cochranella granulosa* is 2N = 20 [153].

**Specimens examined:** *Cochranella granulosa*: Ecuador: *Provincia de Esmeraldas:* 4 km W Durango (1.042◦ N, 78.1081◦ W, 253 m), QCAZ 32769. *Provincia de Cotopaxi:* Reserva El Jardín de los Sueños (0.8416◦ S, 79.2006◦ W, 461 m), MZUTI 4811.

*Cochranella litoralis* (Ruiz-Carranza and Lynch, 1996 [154]; Figures 70–73).

*Centrolene litoralis* Ruiz-Carranza and Lynch, 1996 [154]. Holotype: ICN 13821.

Type locality: "Departamento de Nariño, municipio de Tumaco, La Guayacana, Litoral Pacífico, 1◦49.8- Latitud N, 78◦46.2- W de Greenwich, 100 m, (Colombia)".

*Centrolene litorale*—Frost, 2004 [155].

*Cochranella litoralis*—Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, and Vilà, 2009 [1].

**Common names:** English: Litoral Glassfrog. Spanish: Rana de Cristal del Litoral.

**Etymology:** The specific epithet *litoralis* makes reference to the proximity of the type locality to the ocean [154].

**Identification:** *Cochranella litoralis* is the only species in Centrolenidae that has a bright orange to red iris (Figures 70 and 71) and adult males with humeral spines and a nuptial pad that partially covers the two innermost fingers (Figure 72). Additionally, its dorsum is pale yellow green with small grey to black dots and faint yellow–cream dorsolateral stripes. Males are small (SVL<20 mm) and have humeral and distinct prepollex. In the Chocoan lowlands, only some populations of *Espadarana prosoblepon* and *Nymphargus gri*ffi*thsi* have a green dorsum with dark dots and humeral spines. Both *E*. *prosoblepon* and *N*. *gri*ffi*thsi* are considerably larger, have a cream to white iris, and lack prepollical spines. The Peruvian *Cochranella guayasamini* also has a red iris and humeral spines, but lacks dark dorsal spots [19].

**Figure 70.** *Cochranella litoralis* in life. Adult male, QCAZ 27693, from near Durango, 220 m, Esmeraldas province, Ecuador. Photos by Luis A. Coloma.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout truncated in dorsal and lateral profiles (Figure 72); (3) tympanum of moderate size, oriented almost vertically, with slight lateral and posterior inclinations, its diameter about 30% of eye diameter; tympanic annulus mostly visible, with supratympanic fold covering its posterodorsal margin; tympanic membrane translucent, partially pigmented, clearly differentiated from surrounding skin; (4) dorsal skin smooth anteriorly and shagreen in sacral region, lacking spicules; (5) lacking pair of enlarged subcloacal warts; (6) anterior half of ventral parietal peritoneum with iridophores, posterior half translucent (condition P2); iridophores in pericardium and gastrointestinal peritoneum; no iridophores on testes, gall bladder, urinary bladder, and kidneys (condition V2); (7) liver with clearly defined lobes covered by transparent peritoneum (condition H0); (8) males with small and pointy humeral spines; (9) webbing absent between Fingers I and II, absent or vestigial between Fingers II and III, moderate between outer fingers; webbing formula IV 2—2− V; (10) webbing between toes moderate; webbing formula on foot I 1—2 II 1—2<sup>+</sup> III 1—21/<sup>2</sup> IV 21/2—1 V; (11) ulnar and tarsal folds absent or low and inconspicuous, lacking white coloration; (12) distinct prepollex; nuptial pad Type V; (13) Finger I as long as Finger II; (14) disc of Finger III width about 33% of eye diameter; (15) in life, dorsum green usually with small dark spots and faint yellow–cream dorsolateral stripes (Figures 70 and 71); pale green bones; (16) in preservative, dorsum lavender with dark spots; (17) in life, iris orange to red; (18) dorsal surfaces of fingers and toes lacking melanophores, except for some on base of Toe V; (19) calling behavior unknown; call undescribed; (20) fighting behavior unknown; (21) egg deposition site unknown; parental care unknown; (22) tadpoles unknown; (23) minute body size; in two adult males, SVL 19.4–20.0 mm; females unknown.

**Figure 71.** *Cochranella litoralis*in life. Male from Tundaloma Lodge (1.182 N, 78.749 W; 74 m), Esmeraldas province, Ecuador (3 January, 2014; not collected). Photo by Lucas Bustamante/Tropical Herping.

**Color in life** (Figures 70 and 71): *Cochranella litoralis* has a pale yellowish–green dorsum with dark spots that vary in size and conspicuousness; the anterior half of the venter is white, turning transparent posteriorly; the iris is bright orange to red.

**Color in ethanol:** Dorsal surfaces cream lavender with small dark spots, and two faint white dorsolateral lines. White parietal peritoneum covering anterior half of venter. White pericardium; white peritonea covering stomach and lower colon. Liver, testes, kidneys, and urinary and gall bladders covered by translucent peritonea. Iris silvery white, with black punctuations.

**Biology and ecology:** A male of *Cochranella litoralis* and an egg clutch presumed to be from the same species were found on riverine vegetation (L. Bustamante, pers. comm.). The egg clutch was placed on the upper side of a leaf and contained 25 embryos (Figure 71). Parental care unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Figure 72.** *Cochranella litoralis*, holotype, adult male, ICN 13821. (**A**) Head in lateral view. (**B**) Head in dorsal view. (**C**) Fingers I and II in dorsal view; note that nuptial pad extends on the two fingers; illustrated by Juan M. Guayasamin. (**A**,**B**) Modified from Ruiz-Carranza and Lynch [154].

**Distribution** (Figure 73): *Cochranella litoralis* has been reported from the type locality in Colombia and four localities in Ecuador at elevations below 260 m ([130,154], this work). In Ecuador, this species has a potential distribution of 5784 km<sup>2</sup> within the Chocoan Tropical Forest region.

**Figure 73.** Distribution of *Cochranella litoralis* in Ecuador (yellow dots).

**Conservation status:** Globally, *Cochranella litoralis* is currently listed as *Vulnerable* by the IUCN [156]. In Ecuador, it has a distribution restricted to the provinces of Esmeraldas and Cotopaxi (Chocó ecoregion), where logging is causing continuous habitat reduction and fragmentation. The most recent reports of this species are from Río Cachabí (September 2005), Tundaloma (January 2014), and Jardín de los Sueños (2018). In Ecuador, because of habitat loss and mining, we sugges<sup>t</sup> that the species should be considered as *Endangered*, following IUCN criteria B2a, B2(iii).

**Evolutionary relationships** (Figure 69): *Cochranella litoralis* is sister to all other member of the genus *Cochranella*, but with low nodal support. Twomey et al. [19] recovered a sister relationship between *C*. *litoralis* and *C*. *nola*.

**Specimens examined:** *Cochranella litoralis:* Colombia: *Departamento de Nariño*: Municipio de Tumaco, La Guayacana (1.83 N, 78.77 W; 100 m), ICN 13821. Ecuador: *Provincia de Esmeraldas:* stream near Durango (1.047 N, 78.618 W; 220 m), QCAZ 27693; Pichiyacu, Comunidad Chachi, Río Cayapas (0.9397◦ N, 79.005◦ W; 260 m), QCAZ 31705; Río Cachabí (1.03 N, 78.77 W, 200 m), 2 km NE Urbina on the San Lorenzo—Lita road, DHMECN 3198; Tundaloma Lodge (1.17868◦ N, 78.7497◦ W; 74 m), MZUTI 3481.

**Localities from the literature:** *Cochranella litoralis:* Ecuador: *Provincia de Esmeraldas:* Tsejpu, Río Zapallo (0.7 N, 78.9 W, 150 m) [156].

*Cochranella mache* Guayasamin and Bonaccorso, 2004 [157] (Figures 74–76).

*Cochranella mache* Guayasamin and Bonaccorso, 2004 [157]. Holotype: QCAZ 22412.

Type locality: "Riachuelo La Ducha (0◦20-41" N, 79◦42-36" W; 510 m), tributary of Río Aguacatal, Reserva Biológica Bilsa, 27.4 km W (airline distance) of the town of Quinindé, Montañas del Mache, Provincia Esmeraldas, Ecuador".

**Common names:** English: Mache Glassfrog. Spanish: Rana de Cristal de Mache.

**Etymology:** The specific name *mache* is a noun in apposition and refers to the Montañas de Mache, the type locality of the species [157].

**Identification:** *Cochranella mache*is differentiated from most glassfrogs by having a snout gradually inclined in profile and dermal folds with white tubercles on the ventrolateral edges of Finger V, forearm, elbow, Toe V, tarsus, and heel (Figures 74 and 75). Species sharing similar characteristics are: *Centrolene daidalea*, *C*. *savagei*, *C*. *solitaria*, *Cochranella resplendens*, and *C*. *euknemos*. Three of these species (*C*. *daidalea*, *C*. *savagei*, and *C*. *solitaria*) are restricted to the Andes, and *Cochranella resplendens* is found only in the Amazon Basin. The closely related *C*. *euknemos* lacks the tubercles that are characteristic in *C*. *mache* [157].

**Figure 74.** *Cochranella mache* in life. Adult male, QCAZ 27764, from Río La Carolina, 500 m, Esmeraldas province, Ecuador. Photos by Luis A. Coloma.

**Diagnosis:** (1) Each vomer with three or four teeth on dentigerous process; (2) snout subacuminate in dorsal aspect and gradually inclined in lateral profile (Figure 75); (3) tympanum oriented almost vertically, with slight lateral and posterior inclinations, its diameter about 33%–37% of eye diameter; tympanic annulus visible, low supratympanic fold evident, tympanic membrane translucent and pigmented as surrounding skin; (4) dorsal skin shagreen with warts that usually correspond to light spots; males with numerous minute spicules; (5) ventral skin granular; several round, enameled warts around cloaca; cloacal fold present; pair of enlarged subcloacal warts; (6) anterior one-third of parietal peritoneum covered by iridophores (condition P1); iridophores over pericardium and visceral peritonea (digestive tract, gonads); renal capsules covered by iridophores in some individuals (condition V2); (7) liver trior tetralobed, hepatic peritoneum clear or with small, isolated patches of iridophores (condition H0); (8) humeral spines absent; (9) webbing absent between Fingers I and II, reduced between Fingers II and III, and extensive between outer fingers (Figure 75); webbing formula II (1−–1<sup>+</sup>)—(3−–3<sup>+</sup>) III (2−–2)—(1–1<sup>+</sup>) IV; (10) webbing between toes extensive (Figure 75); webbing formula on foot I (1−–1)—(2–2−) II 1—2 III (1–1−)—2− IV (2−–2)—(1–1−) V; (11) ventrolateral edges of Finger IV, forearm, elbow, Toe V, tarsus, and heel with dermal enameled folds and tubercles; (12) concealed prepollex; nuptial pad large; (13) Finger I about same length as Finger II (Finger I about 94%–98% of Finger II); (14) disc of Finger III width about 44%–54% of eye diameter; (15) color in life, dark olive-green to lavender–blue or pale blue dorsum with numerous small yellow to orange spots, large dull to bright yellow patch on top of head (Figure 74); (16) color in preservative, dorsal surfaces pale lavender with small white or cream spots; tubercles on dermal folds of limbs, fingers, and toes cream-white; inner fingers and toes white or unpigmented; (17) in life, iris whitish cream to beige with thin brown reticulation; white to golden circumpupilary ring; (18) melanophores covering dorsal surfaces of Fingers III and IV and Toes IV and V; (19) males call from the upper sides of leaves; each call formed by two notes; dominant frequency at 5383–5426 Hz; (20) fighting behavior unknown; (21) brown eggs placed on upper sides of leaves; parental care unknown; (22) tadpoles unknown; (23) small body size; in adult males, SVL 22.0–27.0 mm (*n* = 43); in adult females, SVL 28.0–33.4 mm (*n* = 4).

**Figure 75.** *Cochranella mache*. ( **A**) Head in lateral view, paratype, adult male, KU 291176. (**B**) Nuptial excrescences on dorsal surface of Finger I, paratype, adult male, QCAZ 22413. ( **C**) Foot in ventral view, KU 291176. ( **D**) Hand in ventral view, KU 291176. Hand and foot not drawn to scale. Modified from Guayasamin and Bonaccorso [157].

**Color in life** (Figure 74): Dorsal surfaces vary from green to lavender blue or pale blue, with small yellow to orange spots. Upper lip with a thin, white margin. White tubercles visible on dermal folds of hind- and forelimbs. Innermost fingers and toes (Fingers I and II, Toes I and II) white in males and partially white in females. Throat and ventral surfaces of limbs blue green. White parietal peritoneum covering anterior half of venter, posterior portion translucent; white pericardial and visceral peritonea. Transparent hepatic peritoneum, but in some individuals, a small patch of white iridophores covers part of liver. White warts surround cloaca. Iris whitish cream to beige with thin brown reticulation; white to golden ring surrounding pupil. Bones green ([157,158], this work).

**Color in ethanol:** Dorsum of head, body, and limbs lavender with small white or cream spots. Anterior one-third of parietal peritoneum covered by iridophores; iridophores also cover pericardium and visceral peritonea (digestive tract and gonads); hepatic peritoneum lacking iridophores or with small, isolated patches of iridophores; renal capsules covered by iridophores in some specimens ([157,158], this work).

**Biology and ecology:** The information shown below was obtained mainly from Ortega-Andrade et al. [159] and, to a lesser extent, from Guayasamin and Bonaccorso [157]. *Cochranella mache* seems to be restricted to small streams and rivulets in primary and secondary forests in lowlands and piedmont forest. Abundance of the species is correlated with rainfall, being higher during rainy season. Males call from the upper surface of leaves or branches on trees and bushes; reproduction seems to be restricted to the rainy season. Although individuals have been observed on di fferent strata of the forest up to 6 m high, *C*. *mache* prefers the midstory vegetation. Amplexus is axillary and a gravid female was observed to have about 30 eggs. The diversity of amphibians and reptiles at the type locality (Reserva Biológica Bilsa) of *C*. *mache* is described by Ortega-Andrade et al. [160]. Parental care unknown.

**Call:** The advertisement call of *Cochranella mache* was described by Ortega-Andrade et al. [159]. The recorded male was calling from the upper side of a dead *Heliconia* leaf, about 2.5 m above ground, horizontally separated from the stream by about 3m. A call consists of two notes, separated by an interval of 0.107–0.130 s. Each note has a duration of 0.038 ± 0.008 (0.029–0.049) s. The dominant frequency is at 5410.2 ± 17.9 (5383–5426) Hz. The harmonics are not visible. The call rate is 1.46 calls per minute.
