**Tadpole:** Not described.

**Distribution** (Figure 173): *Nymphargus manduriacu* is known from a few streams within the Río Manduriacu Reserve (0.31◦ N, 78.85◦ W; 1215–1242 m), Imbabura province, Ecuador [21].

**Figure 173.** Distribution of *Nymphargus manduriacu* in Ecuador (yellow dot).

**Conservation status:** Guayasamin et al. [21] sugges<sup>t</sup> placing the species in the *Critically Endangered*. At Río Manduriacu Reserve (the only known locality of the species), mining has become one the most dangerous threat to biodiversity, especially to species with restricted distributions.

**Evolutionary relationships:** *N. manduriacu* was inferred as sister to *N*. *balionotus* [21].

**Specimens examined:** Ecuador: *Imbabura province:* Reserva Río Manduriacu (0.310◦ N, 78.857◦ W; 1215–1230 m), ZSFQ 0462–66 (type series).

*Nymphargus megacheirus* (Lynch and Duellman, 1973 [22]; Figures 174–176).

*Centrolenella megacheira* Lynch and Duellman, 1973 [22]. Holotype: KU 143245. Type locality: "16.5 km NNE of Santa Rosa, 1700 m, on Quito–Lago Agrio road, Provincia Napo, Ecuador".

*Cochranella megacheira*—Ruiz-Carranza and Lynch, 1991 [6]. *Nymphargus megacheirus—*Cisneros-Heredia and McDiarmid, 2007 [17].

**Common names:** English: Large-handed Glassfrog. Spanish: Rana de Cristal de manos grandes. **Etymology:** The specific epithet is from the Greek words *megas*, meaning large, and *cheiros*, meaning hand; the name is used to refer to the exceedingly large hands of the species [22].

**Identification:** *Nymphargus megacheirus* can be distinguished from other glassfrogs by its green dorsum with small blue to black spots (Figure 174), relatively large size (adult males, SVL 26.8–31.5 mm; adult females, SVL 31.2–32.9 mm), basal or no webbing among fingers (Figure 175), and lack of humeral spines. On the Amazonian slopes of the Ecuadorian Andes, only *N*. *cochranae* has similar characteristics; however, *N*. *cochranae* has small dark ocelli enclosing orange dots on the dorsum (Figure 140), whereas *N*. *megacheirus* has solid dark spots (Figure 174). *Nymphargus megacheirus* has a similar dorsal coloration and hand-webbing pattern as *N*. *garciae*, but the two species have allopatric distributions with *N*. *garciae* inhabiting higher elevations (1900–2700 m) than *N*. *megacheirus* (1300–1740 m). Additionally, *N*. *megacheirus* is slightly larger than *N*. *garciae* (*N*. *garciae*, SVL 25.1–29.9 mm in adult males: SVL 25.9–28.4 mm in adult females).

**Figure 174.** *Nymphargus megacheirus* in life. Adult male, holotype, KU 143245. Photo by W. E. Duellman.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout truncated in dorsal and lateral profiles (Figure 175); (3) tympanum relatively small, its diameter 18.8%–25.4% eye diameter, dorsal border of tympanic annulus covered by supratympanic fold, tympanic membrane pigmented as surrounding skin; (4) dorsal skin of males shagreen to pustular, shagreen in females; numerous spicules present in males; in females, spicules present only on head, tympanic region, and limbs; (5) skin of venter areolate; pair of enlarged subcloacal warts; (6) anterior two-thirds to three-fourths of venter covered by white parietal peritoneum, posterior portion transparent (condition P3); white pericardium; translucent peritoneum covering intestines, stomach, testes, kidneys, gall bladder, and urinary bladder (condition V1); (7) liver tetralobed, two large ventral lobes partially covering two smaller lobes; hepatic peritoneum transparent (condition H0); (8) humeral spines absent; (9) hand webbing absent between inner finger, absent or basal between outer fingers (Figure 175), webbing formula: III (2<sup>1</sup>/2–3)—(21/2–3) IV; (10) foot about one-half webbed: I

(2–2−)—(2<sup>+</sup>–21/2) II (1–1<sup>2</sup>/3)—(21/2–22/3) III (1<sup>+</sup>–13/4)—(21/2–3−) IV (2<sup>1</sup>/2–3−)—(1<sup>1</sup>/2–2−) V; (11) ulnar fold conspicuous; tarsal folds low; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger I usually slightly shorter than Finger II (Finger I 88.9%–101.3% length of Finger II); (14) disc of Finger III relatively large, 49.7%–59.0% eye diameter; (15) in life, dorsum green with blue to black dots (Figure 174); bones green; (16) in ethanol, dorsal surfaces lavender with small dark purple dots; (17) iris greyish bronze with thin black reticulation; (18) dorsal surfaces of fingers and toes lacking melanophores; (19) males call from upper side of leaves overhanging streams; calls unknown; (20) fighting behavior unknown; (21) eggs deposition site and parental care unknown; (22) tadpoles unknown; (23) medium body size; in adult males, SVL 26.8–31.5 mm (X = 28.3 ± 0.902, *n* = 29); in adult females, SVL 31.2–32.9 mm (X = 32.3 ± 0.7805, *n* = 4).

**Figure 175.** *Nymphargus megacheirus*, KU 143269. (**A**) Head in lateral view. (**B**) Head in dorsal view. (**C**) Hand in ventral view. Illustrations by Juan M. Guayasamin.

**Color in ethanol:** Dorsal surfaces of head, body, forelimbs, and hind limbs lavender with small, round, black spots [22]. White lining covering pericardium and nearly two-thirds of anterior ventral parietal peritoneum. Liver, intestines, stomach, testes, kidneys, gall bladder, and urinary bladder lack iridophores.

**Biology and ecology:** *Nymphargus megacheirus* is active at night. During the reproductive season, males were calling from the upper surfaces of leaves overhanging fast-moving streams in cloud forest at the type locality (16.5 km NNE of Santa Rosa) in October 1971; *Espadarana audax*, *Centrolene pipilata*, and *N*. *siren* were also found there. At the Río Azuela, *N*. *anomalus*, *N*. *siren*, *Hyalinobatrachium pellucidum*, and *C*. *pipilata* occurred along the same streams with *Nymphargus megacheirus* [22]. Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 176): *Nymphargus megacheirus* is endemic to the Amazonian slope of the Andes of Ecuador and Colombia at elevations between 1300 and 1750 m ([22,101], this work). In Ecuador, this species has been reported from the provinces of Napo and Sucumbíos (Specimens Examined). The habitat of the species in Ecuador is within the Eastern Montane Forest region.

**Figure 176.** Distribution of *Nymphargus megacheirus* in Ecuador (yellow dots).

**Color in life** (Figure 174): Dorsal surfaces green with small blue to black spots. Ventral surfaces mostly white, except for translucent posterior third of venter. Upper lip and ulnar folds white. Bones green. Iris greyish bronze [22].

**Conservation status:** Globally listed as *Endangered* by the IUCN [271]. The last records of *Nymphargus megacheirus* correspond to specimens collected at Río Azuela and Río Salado on 24 February 1979 (USNM 286700–01, RWM, pers. obs. [17]). Surveys at Río Azuela have failed to find additional individuals [91]. Then, considering its limited distribution and lack of recent records, we sugges<sup>t</sup> that the species should be considered as *Critically Endangered*.

**Evolutionary relationships** (Figure 136): With the current taxon and gene sampling, *Nymphargus megacheirus* is sister to *N*. *anomalus*.

**Specimens examined:** *Nymphargus megacheirus:* Ecuador: *Provincia de Napo:* 16.5 km NNE Santa Rosa (0.21861 S, 77.7319 W, 1700 m), KU 143245–72; 14.7 km (by road) NE of Río Salado (0.12889 S, 77.6083 W, 1300 m), USNM 286701; 2 km SSW Río Reventador (0.1 S, 77.6 W, 1700 m), KU 164614; *Provincia de Sucumbíos:* Río Azuela (0.1167 S, 77.6167 W, 1740 m), KU 143273–77, 166329; Rio Azuela, where river crosses Quito road (0.1166 S, 77.6166 W, 1700 m.), USNM 286700. Colombia: *Departamento de Putumayo:* 10.3 km W El Pepino (1.05 N, 76.9559 W, 1300 m), KU 169664–65.

*Nymphargus posadae* (Ruiz-Carranza and Lynch, 1995 [255]; Figures 177–179).

*Cochranella posadae* Ruiz-Carranza and Lynch, 1995 [255]. Holotype: ICN 11307. Type locality: "Departamento de Cauca, municipio de Inzá, Km 61 carretera Popayán a Inzá, vertiente oriental Cordillera Central, 2◦34- latitud, 76◦4- W de Greenwich, 2800 m", Colombia.

*Nymphargus posadae*—Cisneros-Heredia and McDiarmid 2007 [17].

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

**Etymology:** The specific epithet honors Dr. Andrés Posada Arango, for his work in the fields of zoology, botany, education, and conservation biology in Colombia [255].

**Identification:** Among glassfrogs that inhabit the Amazonian slopes of the Ecuadorian Andes, *Nymphargus posadae* is unique by having a green dorsum with small greenish–white warts (Figure 177), a slightly sloping snout in lateral profile, and by lacking webbing between fingers. The only species that could be confused with *N*. *posadae* is *Centrolene buckleyi*, which has humeral spines in adult males (humeral spines absent in *N*. *posadae*).

**Figure 177.** *Nymphargus posadae* in life. Adult male from Yanayacu Biological Station, 2100 m, Napo province, Ecuador, QCAZ 25090. Photos by Martín Bustamante.

**Diagnosis:** (1) Vomers with edentate dentigerous process; (2) snout truncated to round in dorsal aspect, and truncated to slightly sloping in lateral profile (Figure 178); (3) tympanum almost indistinguishable; small when visible, its diameter 21.2%–26.5% of eye diameter; tympanic membrane not differentiated from surrounding skin; supratympanic fold present; (4) dorsal skin covered with numerous small warts and some scattered larger warts; no spicules visible; (5) ventral skin areolate; pair of enlarged subcloacal warts; (6) white parietal peritoneum covering anterior 50%–60% of venter (condition P2); white pericardium; no iridophores in peritonea covering intestines, stomach, and kidneys; translucent peritoneum around gall and urinary bladders (condition V1); (7) liver lobate, covered by transparent peritoneum (condition H0); (8) humeral spines absent; (9) webbing between fingers absent or greatly reduced; webbing formula: III (2<sup>3</sup>/4–3−)—(2<sup>2</sup>/3–23/4) IV; (10) feet about two-thirds webbed; webbing formula: I (2−–2)—(2<sup>+</sup>–21/4) II (1<sup>1</sup>/3–11/2)—(21/3–23/4) III 11/2—(22/3–23/4) IV (2<sup>3</sup>/4–3)—2− V; (11) ulnar and tarsal folds low or absent; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger II longer than Finger I (Finger I length 92.2%–95.7% of Finger II); (14) disc width of Finger III about 49.3%–52.9% of eye diameter; (15) in life, dorsum green with small greenish–white warts; bones green; (16) in preservative, dorsum lavender with small white dots; (17) iris white with thin dark grey reticulations; (18) dorsal surfaces of fingers and toes lacking melanophores, except for proximal portion of Toes IV and V; (19) males call from upper surfaces of leaves; calls unknown; (20) fighting behavior unknown; (21) eggs deposition site and parental care unknown; (22) tadpole unknown; (23) large body size; male SVL 30.7–34.1 mm (X = 32.3, *n* = 6); female SVL 30.2–33.3 mm (X = 31.4, *n* = 4).

**Figure 178.** *Nymphargus posadae*, adult males from Yanayacu Biological Station, Napo province, Ecuador. (**A**) Head in lateral view, QCAZ 26023. (**B**) Head in dorsal view, QCAZ 26023. (**C**) Hand in ventral view, QCAZ 25090. (**D**) Finger I in dorsal view, QCAZ 26023. (**E**) Foot in ventral view, QCAZ 26023. Modified from Guayasamin et al. [20].

**Color in life** (Figure 177): The following description corresponds to individuals from Yanayacu Biological Station [20]. Dorsal surfaces of head, body, and limbs bright green with small, scattered, greenish–white warts. Upper lip white; lower lip with thin white border. Ventrolateral border of arm, Finger IV, tarsus, and Toe V white. Cloacal region with numerous small, white warts. White parietal peritoneum covering about anterior half of venter. Iris white with thin dark grey reticulations.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs lavender with some larger warts being bluish white. Upper lip white; thin white line evident on lower lip. Ventrolateral border of arm, Finger IV, tarsus, and Toe V white. Ventral surfaces of forearm and tarsus completely covered with white in two specimens (QCAZ 25090 and 26022). Dorsally, Fingers I and II and Toes I, II, and III unpigmented: Some pigmentation visible on Fingers III and IV and Toes IV and V. Cloacal region with several white warts. In adult males, nuptial pad cream (Type I). Parietal peritoneum white anteriorly, covering approximately 50%–60% of venter. Pericardium silver white. No iridophores on the hepatic peritoneum, digestive tract, or kidneys ([20], this work).

**Variation:** Males from Yanayacu Biological Station are smaller (SVL = 30.7–31.9, *n* = 3) than those from Colombian localities (SVL = 32.7–34.1 mm, *n* = 3) [20,255].

**Biology and ecology:** In Colombia, the species was observed on vegetation and rocks along a creek [255]. Carranza and Lynch [255] reported that the eggs have a pigmented animal pole (dark brown) and unpigmented vegetal pole (cream). Unfortunately, they did not mention where the eggs were deposited. In Ecuador, three individuals were collected during three years of inventory work at Yanayacu; all frogs were found calling on the same night (12 June 2003), on ferns 110–220 cm above a stream. Males call from the upper surfaces of leaves [20]. Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 179): In Colombia, *Nymphargus posadae* is known from the Caldas, Cauca, and Huila departments on the eastern flank of the Central Cordillera of the Andes, between 1100 and 2800 m [255]. In Ecuador, the species has been reported from localities on the Amazonian slopes of the Andes at elevations of 1750–2100 m ([17,20], this work). *Nymphargus posadae* is also present in Peru (Cordillera del Cóndor) [272].

**Figure 179.** Distribution of *Nymphargus posadae* in Ecuador (yellow dots).

**Conservation status:** Globally, *Nymphargus posadae* is considered as *Least Concern* by the IUCN [273]. The species has a relatively large distribution and it is found within several protected areas. Thus, we agree with the current conservation assessment.

**Evolutionary relationships** (Figure 136): Given the current taxon and gene sampling, *Nymphargus posadae* is sister to *N*. *pluvialis*.

**Specimens examined:** *Nymphargus posadae:* Colombia: *Departamento de Huila:* 6.2 km NW of San José de Isnos, 1940 m. Ecuador: *Provincia de Napo:* Yanayacu Biological Station (0◦41- S, 77◦53- W; 2100 m), QCAZ 25090, 26022–23. *Provincia de Zamora Chinchipe:* tributary of Río Jambue, ca. 15 km S from Zamora (ca. 4◦14- S, 78◦57- W; 1750 m). *Provincia de Sucumbíos:* Río Chingual, ca. 3 km N of Sebundoy, ca. 20 km N of La Bonita (ca. 0◦26- S, 77◦32- W; 1890 m), USNM 288464-65. Peru: Cordillera del Cóndor (ca. 05◦25-16.5" S, 78◦35-23.2" W; 1890 m).

*Nymphargus siren* (Lynch and Duellman, 1973 [22]; Figures 180–182).

*Centrolenella siren* Lynch and Duellman, 1973 [22]. Holotype: KU 146610.

Type locality: "small tributary of the Río Salado, about 1 km upstream from the Río Coca, 1410 m, Provincia Napo, Ecuador."

*Cochranella siren*—Ruiz-Carranza and Lynch, 1991 [6].

emphNymphargus siren—Cisneros-Heredia and McDiarmid, 2007 [17].

**Common names:** English: Siren Glassfrog. Spanish: Rana de Cristal sirena.

**Etymology:** In Greek mythology, sirens were bird-women, who by their sweet singing enticed seafarers to destruction; the name is used in allusion to the calls of these frogs that entice biologists to the nocturnal perils of streams [22].

**Identification:** Among glassfrogs that inhabit the Amazonian slopes of the Ecuadorian Andes, *Nymphargus siren* (Figure 180) is distinguished by having a green dorsum with small yellow spots, a partially white venter, small size (male SVL < 22.6 mm; female SVL < 23.3 mm), and lacking humeral spines. Similar species from eastern Ecuador include *Espadarana audax*, *Nymphargus humboldti* sp. nov., *Rulyrana flavopunctata*, and *Teratohyla midas*. Differences among these species include the presence of humeral spines in adult males of *E*. *audax* (humeral spines absent in *N*. *siren*), moderate webbing between Fingers III and IV in *E*. *audax* and *R*. *flavopunctata* (webbing absent in *N*. *siren* and basal in *T*. *midas*), and the white peritoneal covering of the digestive tract of *T*. *midas* (digestive tract opaque or translucent in *N*. *siren*, *E*. *audax*, and *R*. *flavopunctata*). *Nymphargus humboldti* sp. nov. and *N*. *siren* have non-overlapping body sizes (*N*. *humboldti* sp. nov., male SVL = 23.3–25.2 mm; female SVL = 24.3–25.9 mm).

**Figure 180.** *Nymphargus siren* in life. Individual (MZUTI 774) from creek on the Oyacachi–El Chaco trail, 1878 m, Napo province, Ecuador. Photos by Luis A. Coloma.

**Diagnosis:** (1) Vomers with edentate dentigerous process; (2) snout usually truncated in dorsal aspect; truncated to slightly protruding in lateral profile (Figure 181), (3) tympanum oriented almost vertically, with slight lateral and posterior inclinations, its diameter about 20.5%–30.5% of eye diameter; upper fourth of tympanic annulus obscured by supratympanic fold; tympanic membrane clearly differentiated from surrounding skin; (4) dorsal skin shagreen, usually with spicules in males; (5) pair of enlarged subcloacal warts; (6) white parietal peritoneum covering anterior half of venter (condition P2); white pericardium; translucent to opaque peritonea covering intestines, stomach, and kidneys; translucent peritoneum around gall and urinary bladders (condition V1); (7) liver lobate, covered by transparent peritoneum (condition H0); (8) humeral spines absent; (9) webbing absent between inner fingers, absent or basal between outer fingers (Figure 181); webbing formula IV (2<sup>2</sup>/3–3)—(21/4–23/4) V; (10) feet about two-thirds webbed; webbing formula: I (2−–2)—(2<sup>+</sup>–21/2) II (1<sup>1</sup>/4–11/2)—(21/2–3−) III (1<sup>+</sup>–2−)—(2<sup>3</sup>/4–3−) IV (3−–3)—(2−–2) V; (11) ulnar fold absent; external tarsal fold absent; low inner tarsal fold evident; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger II slightly longer than Finger I (Finger I length 91.5%–99.6% of Finger II); (14) disc of Finger III width about 43.3%–58.5% of eye diameter; (15) in life, dorsum green with small yellow spots (Figure 180); bones green; (16) in preservative, dorsum lavender with small white spots; (17) iris, in life, whitish cream, with a yellow hue around pupil and fine, dark grey reticulations; (18) melanophores absent from dorsal surfaces of fingers and toes, except for few on Toe V; (19) males call from upper surface of leaves, call unknown; (20) fighting behavior unknown; (21) egg deposition site and parental care unknown; (22) tadpoles unknown; (23) minute body size; males, SVL 19.8–22.6 mm (X = 20.9 ± 0.931, *n* = 24); females, SVL 22.5–23.3 mm (*n* = 2).

**Figure 181.** *Nymphargus siren*, KU 146610. (**A**) Head in lateral view. (**B**) Hand in ventral view. Illustrations by Juan M. Guayasamin.

**Color in life** (Figure 180): Green dorsum with small yellow spots, which are narrowly bordered by black in some individuals [22]. Anterior half of venter white, posterior half transparent. Bones green. Iris whitish cream, with yellow hue around pupil and fine, dark grey reticulations.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs light to dark lavender with small white spots. Anterior half of ventral parietal peritoneum white; posterior half translucent. Translucent peritonea covering gall and urinary bladders. Iridophores absent from digestive tract, liver, and kidneys.

**Variation:** Spicules are absent in a few males (KU 164636, 297290). As noted, the presence of spicules may be a reflection of reproductive activity. Two males (KU 178199, 297292) lack yellow spots on the dorsum.

**Biology and ecology:** Relatively large numbers of individuals of *Nymphargus siren*, including calling males, were observed at the type locality on 7 April 1972 (13 individuals) and again on 18 March 1975 (15 individuals; WED's field notes). Many specimens were also noted at 3.2 km NNE Oritoyacu on 15 July 1977 (21 individuals; John D. Lynch's field notes). Males call from the upper surfaces of leaves (WED field notes, 7 April 1972). Parental care is unknown.

**Call** (Figure 182): We analyzed 14 notes from two individuals (MZUTI 765, 775). The typical advertisement call is short and is composed by a single note. Note duration is 20–42 (mean = 24, SD = 7) ms. Notes are generally pulsed and have one to three (mean = 2.2, SD = 0.6) amplitude peaks, where the first peak is more pronounced that the others. Notes have their peak amplitude in the first 50% of the note (relative peak time: Range = 0.0595–0.1751, mean = 0.123, SD = 0.032). Pulses within a note have a rate of 24–130 (mean = 86, SD = 23) pulses per second. The dominant frequency of a note measured at peak amplitude is 4737–6029 (mean = 4977, SD = 323) Hz and is contained within the fundamental frequency. The fundamental frequency has a lower limit of 4651–5943 (mean = 4842, SD = 331) Hz and a higher limit of 4823–6115 (mean = 5088, SD = 318) Hz.

**Figure 182.** Call of *Nymphargus siren*, MZUTI 765, recorded at trail between Oyacachi and El Chaco, 1645 m, Napo province, Ecuador. Air temperature = 15 ◦C.

**Tadpole:** Not described.

**Distribution** (Figure 183): *Nymphargus siren* inhabits the cloud forests of the Amazonian slopes of the Ecuadorian and Colombian Andes at elevations between 1410 and 2000 m ([22,176], this work). In Ecuador, the species has been reported from the provinces of Napo and Sucumbíos. Rodríguez et al. [274] reported *N*. *siren* from Peru (Departamento Ayacucho), but we consider this report to be based on a misidentification and restrict the distribution of *N*. *siren* to Colombia and Ecuador. In Ecuador, the potential distribution of the species is 3881 km<sup>2</sup> within the Eastern Montane Forest region.

**Figure 183.** Distribution of *Nymphargus siren* in Ecuador (yellow dots).

**Conservation status:** *Nymphargus siren* is listed as *Vulnerable* by the IUCN because the extent of its occurrence is less than 20,000 km2, its distribution is severely fragmented, and the extent and quality of its habitat continues to decline [275]. No data are available on the population demography of the species nor on its susceptibility to chytridiomycosis, climate change, and/or changes on UV radiation. Recent records of *N*. *siren* are from the Río Azuela (March 2000) [91], Yanayacu Biological Station (April 2008) [125], and the Oyacachi–El Chaco trail (May 2012; JMG pers. obs.). The population status of the species at localities where it was historically abundant (e.g., Río Salado; 3.2 km NNE Oritoyacu) is unknown. We maintain its conservation category.

**Evolutionary relationships** (Figure 136): *Nymphargus siren* and *N*. *humboldti* sp. nov. are sister species.

**Remarks:** Although Lynch and Duellman [22] mentioned that some individuals have up to two teeth on the dentigerous process of the vomer, all the specimens examined by us had edentate vomers.

**Specimens examined:** *Nymphargus siren:* Ecuador: *Provincia de Napo:* Yanayacu Biological Station, 2100 m, QCAZ 37971, 37975; tributary of the Río Salado, about 1 km upstream from the Río Coca (0.19167 S, 77.6997 W; 1410 m), KU 146610 (holotype), KU 146611–23 (paratypes), 164635–49, 178191–206, 146610–23, QCAZ 14425, 30975, 30977–80; Baeza (0.46795 S, 77.567 W; 1650 m), KU 190020–21; 16.5 km NNE Santa Rosa (0.2186 S, 77.732 W; 1700 m), KU 143288–89, 143291, 143293–94; 3.2 km NEE Oritoyacu (0.4597 S, 77.867 W; 1910 m), KU 178170–90; Oyacachi-El Chaco trail at elevations between 1645–1800 m, MZUTI 765–776. *Provincia de Sucumbíos:* Río Azuela (0.11667 S; 77.6167 W, 1740 m), QCAZ 15263, 15266, KU 143295–97, 155499–501; Colombia: *Putumayo:* 35 km SE of San Francisco, 1950 m, KU 169668–69.

*Nymphargus sucre* Guayasamin, 2013 [276] (Figures 184–186).

*Nymphargus sucre* Guayasamin, 2013 [276]. Holotype: MZUTI 1421.

Type locality: "creek on the Plan de Milagro–Gualaceo road (3.0077◦ S, 78.53318◦ W; 2159 m),

Provincia Morona Santiago, Ecuador."

**Common names:** English: Sucre's glassfrog. Spanish: Rana de Cristal de Sucre.

**Etymology:** The specific epithet is a noun in apposition and honors Antonio José de Sucre, who, with Simón Bolívar, led the independence of most Andean countries (Bolivia, Colombia, Ecuador, Peru, and Venezuela) from Spain. The epithet also makes reference to the national currency of Ecuador between 1884 and 2000; the Sucre disappeared in the year 2000, when it was replaced by the US dollar after a disastrous economic policy that affected millions of Ecuadorians. As an analogy, the current destruction of habitats in southeastern Ecuador (as in many other regions) is likely to drive many species to extinction if activities such as mining, oil extraction, road building, cattle, and agriculture are promoted irresponsibly, and without assessing their effect on megadiverse areas and endangered species [276].

**Identification:** *Nymphargus sucre* is distinguished from most glassfrogs by having, in life, a brownish–yellow dorsal surface with yellow spots (Figure 184), and lacking webbing between the fingers. *Nymphargus sucre* is most similar to four other species that lack the typical green dorsal coloration of centrolenids (*N*. *anomalus*, *N*. *colomai* sp. nov., *N*. *ignotus*, and *N*. *rosada;* Figure 185). *Nymphargus anomalus* and *N*. *ignotus* differ from *N*. *sucre* mainly by having, in life, a pale tan to brown dorsum with black ocelli (ocelli absent in *N*. *sucre*) and lacking yellow spots. *Nymphargus rosada* is distinguished by its pink coloration in life with yellowish orange spots on the dorsum (brownish–yellow dorsum with yellow spots in *N*. *sucre)* and by being larger with non-overlapping SVL in adult males (SVL = 24.9–28.3 in male *N*. *rosada* [133]; SVL = 21.6–22.3 mm in male *N*. *sucre*). Finally, *N*. *colomai* sp. nov. differs by having numerous diffuse yellow spots on the dorsum (fewer and well-defined spots in *N*. *sucre*), and a white iris with a contrasting horizontal black stripe (iris lacking horizontal stripe and having a yellow hue around the pupil in *N*. *sucre* [276]; Figure 185).

Other species that may have yellowish–green dorsal patterns and could be confused with *N*. *sucre* are *N*. *armatus*, *N*. *oreonympha*, *N*. *nephelophila*, and *N*. *ruizi*. These four species are readily distinguished from *N*. *sucre* by having black dorsal spots and lacking yellow spots. Additionally, adult males of *N*. *armatus* have nuptial pads with a Type III morphology, whereas *N*. *sucre* has a Type I morphology. Adult males of *N*. *oreonympha*, *N*. *nephelophila*, *N*. *armatus*, and *N*. *ruizi* are larger (SVL 22.6–24.1 mm, SVL 24.0–26.3 mm, SVL 23.3–24.8 mm, SVL 24.3–26.4 mm, respectively [154,277,278] than males of *N*. *sucre* (SVL 21.6–22.3 mm). Finally, *N*. *spilotus*, an endemic to the eastern slope of the Cordillera Central of Colombia, has a dorsal pattern that resembles that of *N*. *sucre* (with yellow spots), but is a considerably larger species (adult male SVL = 25.3–26.4 mm; female SVL = 27.6–28.5 mm [133]) and has prominent vomerine teeth (absent in *N*. *sucre* [276]).

**Figure 184.** *Nymphargus sucre* in life. (**A**–**C**) Adult male, holotype, MZUTI 1421. (**D**) Adult female, MZUTI 1422. Photos by Alejandro Arteaga/Tropical Herping. Obtained from Guayasamin [276].

**Figure 185.** Glassfrog species similar to *Nymphargus sucre* [276]. (**A**) *Nymphargus anomalus*, QCAZ 47507; photo by Luis A. Coloma. (**B**) *N. colomai* sp. nov., QCAZ 41590; photo by Juan M. Guayasamin. (**C**) *N. rosada*, photo by M. Rivera. (**D**) *N. sucre*, MZUTI 1421; photo by A. Arteaga/Tropical Herping.

**Diagnosis:** (1) Dentigerous process of the vomer present, but lacking teeth; (2) snout truncated in lateral and dorsal views; (3) tympanum visible without magnification, oriented almost vertically, with slight lateral and posterior inclinations, its diameter about 23%–26% of eye diameter; upper fourth of tympanic annulus obscured by supratympanic fold; tympanic membrane translucent, but with melanophores like those on surrounding skin; (4) dorsal skin shagreen, with numerous minute spicules in males; (5) venter areolate; pair of enlarged subcloacal warts; (6) white parietal peritoneum covering about anterior half of venter (condition P2); white pericardium; translucent peritonea covering intestines, stomach, kidneys, gall and urinary bladders (condition V1); (7) liver trilobed, covered by transparent peritoneum (condition H0); (8) humeral spines absent; (9) webbing absent between inner fingers, basal between Fingers III and IV; webbing formula III (2<sup>3</sup>/4–3−)—(2<sup>1</sup>/2–23/4) IV; (10) feet about two-thirds webbed; webbing formula: I (2–2−)—2<sup>+</sup> II (1<sup>+</sup>–11/3)—(2<sup>+</sup>–21/3) III (1–1<sup>1</sup>/2)—(21/3–21/2) IV (2<sup>2</sup>/3–3−)—2− V; (11) ulnar and tarsal folds present, but low and inconspicuous, lacking pigmentation; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger I slightly shorter than Finger II (Finger I length 93%–96% of Finger II); (14) disc width of Finger III about 37%–53% of eye diameter; (15) in life, dorsum brownish yellow with yellow spots; color of bones unknown; (16) in preservative, dorsum grey lavender with small white or unpigmented spots; (17) iris white with minute dark spots, thin reticulation, and yellow hue around pupil; (18) dorsal surfaces of fingers and toes lacking melanophores, except for few on Fingers III and IV, and Toes III, IV, and V; (19) males call from the upper surfaces of leaves; call unknown; (20) fighting behavior unknown; (21) egg deposition site and parental care unknown; (22) tadpoles unknown; (23) medium body size; males, SVL 21.6–22.3 mm (*n* = 2); female, SVL 24.3 mm (*n* = 1).

**Color in life** (Figure 184): Brownish–yellow dorsum with small yellow spots; dorsum of the holotype with a greenish hue. Upper lip unpigmented. Anterior half of ventral parietal peritoneum white, posterior portion translucent. Color of bones unknown. Iris white with minute dark spots, thin reticulation, and yellow hue around pupil [276].

**Color in ethanol:** Dorsal surfaces of head, body, and limbs grey lavender with small white or unpigmented spots. Anterior half of ventral parietal peritoneum white. Heart covered by white pericardium; translucent peritonea covering gall and urinary bladders; iridophores absent from digestive tract, liver, and kidneys. Observations on internal anatomy were based on specimen MZUTI 1421 [276].

**Biology and ecology:** During the night, males of *Nymphargus sucre* were observed calling on leaves at 90–130 cm above a stream on 6 June 2012. Only one other species of glassfrog ( *N*. *cariticommatus)* was found at the same stream [276]. Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 186): *Nymphargus sucre* is only known from the type locality, a creek on the Plan de Milagro–Gualaceo road (3.0077◦ S, 78.53318◦ W; 2140–2160 m), Morona Santiago province, Ecuador [276].

**Conservation status:** *Nymphargus sucre* has not been evaluated by the IUCN. Because of habitat loss and mining, we sugges<sup>t</sup> that the species should be considered as *Critically Endangered*, following IUCN criteria B1, B2a,b(iii).

**Evolutionary relationships** (Figure 136): *Nymphargus sucre* is sister to *N*. *cariticommatus*.

**Figure 186.** Distribution of *Nymphargus sucre* in Ecuador (yellow dot).

**Specimens examined:** *Nymphargus sucre:* Ecuador: *Morona Santiago province:* Plan de Milagro-Gualaceo road (3.0077◦ S, 78.53318◦ W; 2100 m), MZUTI 1421 (holotype), MZUTI 1420, 1421 (paratypes).

*Nymphargus wileyi* (Guayasamin, Bustamante, Almeida-Reinoso, Funk, 2006 [20]; Figures 187–189).


*Nymphargus wileyi—*Cisneros-Heredia and McDiarmid, 2007 [17].

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

**Etymology:** The specific name is a noun in the genitive case and a patronym for Edward O. Wiley, for his influential work on the development of phylogenetic systematics and use of the evolutionary species concept [20].

**Identification:** *Nymphargus wileyi* differs from most species of glassfrogs by having a uniform green dorsum (Figure 187), white renal peritoneum, and by lacking any webbing between the fingers. On the Amazonian slopes of the Andes, only *N*. *cariticommatus* can be confused with *N*. *wileyi*. The differences between these two species are subtle; *N*. *wileyi* has a uniform dorsum (dorsum with small yellow dots in *N*. *cariticommatus*) and has a faint layer of iridophores covering the anterior portion of the esophagus (white peritoneum covering most of the esophagus in *N*. *cariticommatus*). The only other species with a uniform green dorsum on the Amazonian slopes of the Andes is *Espadarana durrellorum*, which differs by having webbing between Fingers III and IV, and a humeral spine in males.

**Figure 187.** *Nymphargus wileyi* in life. (**Left**): Adult male from Yanayacu Biological Station, Napo province, QCAZ 37972. (**Right**): Egg clutches of *N*. *wileyi* at Yanayacu Biological Station. Photos by Martín Bustamante.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout truncated in dorsal aspect, truncated to protruding in lateral profile (Figure 188); (3) tympanum oriented almost vertically, with slight posterior and lateral inclinations, its diameter 31.4%–37.8% of eye diameter; tympanic annulus visible anteroventrally, tympanic membrane unpigmented and clearly differentiated from surrounding skin; (4) dorsal surfaces of males and females shagreen, with small spicules evident in males; (5) venter areolate; pair of enlarged subcloacal warts (Figure 15); (6) anterior 40%–60% of ventral parietal peritoneum white, posterior portion transparent (condition P2); white pericardium; translucent to opaque peritoneum covering intestines, stomach, testes, gall bladder, and urinary bladder; peritoneum around kidneys white with unpigmented spots (condition V1); (7) liver tetralobed, covered by transparent peritoneum (condition H0); (8) humeral spines absent; (9) webbing absent between inner fingers; webbing reduced between outer fingers, III 3−—22/<sup>3</sup> IV; (10) concealed prepollex; in males, nuptial pad Type I; (11) Finger II slightly longer than Finger I (Finger I 92%–97% of Finger II); (12) ulnar fold low or absent; inner tarsal low and thin, outer tarsal fold absent; (13) webbing formula on foot I 2—21/<sup>3</sup> II (1<sup>1</sup>/3–12/3)—(21/2–3−) III (1<sup>+</sup>–2−)—(2<sup>2</sup>/3–3−) IV (3−–3)—(2−–2) V (Figure 188); (14) disc of Finger III of moderate size, about 50%–60% of eye diameter; (15) in life, dorsum pale green; bones green; (16) in preservative, dorsum uniform pale lavender; (17) iris coppery white with black reticulation; (18) fingers and toes lacking melanophores on dorsal surfaces; (19) males call from the upper sides of leaves; call unknown; (20) fighting behavior unknown; (21) females deposit eggs on upper surface of leaves near streams (Figure 187); short-term maternal care unknown; prolonged parental care absent; (22) tadpoles undescribed; (23) small body size; males, SVL 24.0–26.2 mm (X = 24.6; *n* = 5); 27.1 mm in one female.

**Figure 188.** *Nymphargus wileyi*. (**A**) Head in dorsal view, QCAZ 26029. (**B**) Head in lateral view, QCAZ 26029. (**C**) Hand in ventral view, QCAZ 26028. (**D**) Foot in ventral view, QCAZ 26028. Modified from Guayasamin et al. [20].

**Color in life** (Figure 187): Dorsal surfaces pale green, lacking spots; lower venter transparent; parietal peritoneum white, covering anterior part of abdomen (heart not visible); iris coppery white with black reticulation; bones green [20].

**Color in ethanol:** Dorsal surfaces of head, body, and limbs pale lavender; upper lip white; dorsally, all fingers, Toes I–III, and most of Toe IV unpigmented; cloacal region mostly unpigmented, except for few minute white flecks. Cream nuptial pad on Finger I. Ventral parietal peritoneum white anteriorly, pericardium white, hepatic peritoneum clear, visceral peritoneum opaque, peritoneum around kidneys white with unpigmented spots (dissected specimens: QCAZ 26029–30) [20].

**Biology and ecology:** *Nymphargus wileyi* is a nocturnal species that seems to be restricted to primary forest. Individuals have been found at night on leaves 120–220 cm above streams (five males) or above the ground (one female). Males call from the upper surfaces of leaves; males are territorial and are usually found near egg clutches, which are on the tip of leaves; distances between egg clutches and males can be as close as 20 cm (Figure 187). Males were never observed on the same leaf as the egg clutch; also, several egg clutches had no male nearby; these observations sugges<sup>t</sup> that prolonged parental care is absent [20]. Short-term maternal care is unknown. Clutches are deposited on the dorsal surface of leaves near their tips (Figure 187). The number of eggs per clutch varies from 19–28 (X = 22; *n* = 17); eggs and embryos in early developmental stages are whitish [20].

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 189): *Nymphargus wileyi* is known only from the type locality (Yanayacu Biological Station, 2100 m) on the Amazonian slopes of the Cordillera Oriental of the Ecuadorian Andes [20]. The habitat of the species is within the Eastern Montane Forest region.

**Figure 189.** Distribution of *Nymphargus wileyi* in Ecuador (yellow dot).

**Conservation status:** *Nymphargus wileyi* is listed as *Data Deficient* by the IUCN [279]. We consider this category appropriate.

**Evolutionary relationships** (Figure 136): *Nymphargus wileyi* is sister to a clade formed by *N*. *sucre* and *N*. *cariticommatus*. These three species are found on the Amazonian slopes of the Andes.

**Taxonomic Remarks:** Only one discrete morphological character separates *Nymphargus wileyi* from *N*. *cariticommatus*, the absence of pale spots in *N*. *wileyi*. Also, in preserved material, the esophageal peritoneum of *N*. *cariticommatus* is more pronounced and extensive than in *N*. *wileyi*, in which the iridophores are restricted to the anterior portion of the esophageal peritoneum. Surprisingly, the two species, although closely related, are not sister to each other (Figure 136).

**Specimens examined:** *Nymphargus wileyi:* Ecuador: *Provincia de Napo:* Yanayacu Biological Station (0◦41- S, 77◦53- W; 2100 m), QCAZ 26028 (holotype), 26024, 26029–30, 26057, 27435.

**Genus** *Rulyrana* Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, and Vilà, 2009 [1].

**Etymology:** The name *Rulyrana* honours Pedro Ruiz-Carranza† and John D. Lynch (Instituto de Ciencias Naturales, Universidad Nacional de Colombia), who have contributed enormously to the understanding of anuran diversity, biology, and evolution. Together, Pedro and John made an outstanding effort to describe the amphibian diversity of Colombia, producing a series of glassfrog papers [6,26,27, 101,110,113,129,133,154,246,255,277] that includes the description of the genus *Hyalinobatrachium* and numerous species. The name *Rulyrana* is femine in gender and comes from an arbitrary association of the two first letters of Ruiz and Lynch (Ruly) and the word rana (=frog). *Ruly* also is the nickname of JMG's good friend and colleague Martín Bustamante, whose work on biodiversity, conservation, and even sportive indoctrination, is remarkable ([1], this work).

*Rulyrana flavopunctata* (Lynch and Duellman, 1973 [22]; Figures 190–194).

*Centrolenella flavopunctata* Lynch and Duellman, 1973 [22]. Holotype: KU 121048. Type locality: "Mera, Provincia Pastaza, Ecuador." *Cochranella flavopunctata*—Ruiz-Carranza and Lynch, 1991 [6]. *Rulyrana flavopunctata*—Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, and Vilà,2009 [1].

**Common names:** English: Yellow-spotted Glassfrog. Spanish: Rana de Cristal de Puntos Amarillos.

**Etymology:** The specific name is a combination of the Latin *flavus*, meaning golden yellow, and *punctatus*, meaning dotted, and is used in reference to the dorsal coloration of the species [22].

**Identification:** Among glassfrogs that inhabit the Amazonian slopes of the Ecuadorian Andes, *Rulyrana flavopunctata* is distinguished by having a green to dark green dorsum with small yellow spots (Figure 190), a partially white venter, small to moderate body size (male SVL 21.0–23.9 mm; female SVL 24.0–27.4 mm), extensive webbing between Fingers III and IV, and lacking humeral spines. Species with a similar color pattern from eastern Ecuador include *Espadarana audax*, *Nymphargus siren*, *N*. *humboldti* sp. nov., *N*. *mariae*, *R*. *mcdiarmidi*, and *Teratohyla midas*. None of the species in the genus *Nymphargus* has webbing between Fingers III and IV; males of *Espadarana audax* have humeral spines; *T*. *midas* has very few yellow spots on the upper flanks and a white digestive tract; and *R*. *mcdiarmidi* has a slightly larger body size (male SVL 22.8–26.9 mm, female SVL 25.4–30.2 mm) and warts on its dorsum (absent in *R*. *flavopunctata*). The differences between *R*. *flavopunctata* and *R*. *mcdiarmidi* are subtle and more work is necessary to verify the validly of the specific status of the latter. See Taxonomic Remarks.

**Figure 190.** *Rulyrana flavopunctata* in life. Ecuador, Napo province, Cordillera de los Guacamayos, 1564 m, MZUTI 1250 (**left**), 1260 (**right**). Photos by Eduardo Toral.

**Diagnosis:** (1) Teeth on vomers present or absent; each vomer with zero to three teeth; (2) snout usually rounded in dorsal and lateral views; (3) tympanum visible, its diameter 27.9%–38.7% of eye diameter; supratympanic fold low; (4) dorsal skin shagreen; males with spicules; (5) skin of venter areolate; pair of slightly enlarged subcloacal warts; (6) anterior half of ventral parietal peritoneum white, posterior half translucent (condition P2); pericardium white; peritonea covering intestines, stomach, and kidneys lacking iridophores; urinary bladder transparent (condition V1); (7) lobed liver lacking iridophores (condition H0); (8) humeral spines absent; (9) webbing between Fingers I and II absent or basal, moderate between Fingers II and III, extensive between Fingers III and IV; webbing formula for outer fingers: II (1<sup>1</sup>/2–2−)—(3–3<sup>+</sup>) III (1<sup>1</sup>/3–2−)—(1<sup>+</sup>–11/2) IV; (10) feet about three-fourths webbed; webbing formula: I (0+–1)—(1–1<sup>3</sup>/4) II (0+–1)—(0+–2) III (0+–1)—(1<sup>1</sup>/3–2+) IV (2−–2<sup>+</sup>)—(0<sup>+</sup>–1) V (Figure 191); (11) ulnar fold low or absent; inner tarsal fold low, short; outer tarsal fold absent; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger I about same length as Finger II (Finger I length 95.5%–104.0% of Finger II); (14) disc of Finger III relatively large, its width 47.7%–67.0% of eye diameter; (15) in life, dorsum green to dark green with well-defined yellow spots; bones green; (16) in preservative, dorsum dark lavender to slate grey with small white to cream spots; (17) iris pale greyish white, with or without golden tint, with dark grey or brown flecks or fine reticulation; (18) dorsal surfaces of fingers and toes usually with melanophores, but some individuals lack melanophores on Finger I or on Fingers I and II; (19) males call from the upper side of leaves or rocks; call unknown; (20) fighting behavior unknown; (21) egg clutches placed on rock walls; parental unknown; (22) tadpoles unknown; (23) small body size in adult males, SVL 21.0–23.9 mm (X = 22.1 ± 0.834, *n* = 10); in adult females, SVL 24.0–27.4 mm (X = 25.3 ± 1.207, *n* = 6).

**Figure 191.** *Rulyrana flavopunctata*. (**A**) Head in lateral view, holotype, adult male, KU 121046. (**B**) Hand in ventral view, KU 121050. (**C**) Foot in ventral view, KU 121048. (**B**,**C**) Modified from Lynch and Duellman [22].

**Color in life** (Figure 190): Conspicuous variation has been observed in dorsal and ventral coloration. Dorsal surfaces of head and body vary from green to very dark green, with small yellow spots. Anterior half of ventral parietal peritoneum white; posterior half usually translucent, but some individuals have a milky colored peritoneum. Iris pale greyish white to brown, with yellow tint, with dark grey or brown flecks or fine reticulation ([22], this work). At one locality (Río Tayuntza), individuals showed considerable variation in dorsal coloration, from green to almost black, with yellow spots, indicating that intraspecific variation in this species is not necessarily associated with geography.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs dark lavender to slate grey, with small white or cream spots usually corresponding to warts. Internal organs lacking iridophores, except for heart that is covered by white pericardium.

**Biology and ecology:** Most individuals were found in a deep ravine where males were perched on small herbs in the spray-zone of a small waterfall. At several small streams between Mera and the Río Alpayacu, males were calling from the upper sides of leaves on 2, 14, and 24 July 1968 [22]. At Río Tayuntza (Morona Santiago province, Ecuador) egg clutches of *R*. *flavopunctata* were found on rock walls during the rainy season [280] (Figure 192). Parental care is unknown.

**Figure 192.** (**Left**): Habitat of a glassfrog tentatively assigned to *Rulyrana flavopunctata* or *R. mcdiarmidi*. (**Right**): Egg clutch of the rock walls of the river. Locality: Río Tayuntza, Cuevas de los Tayos, Río Tayuntza, Morona Santiago province, Ecuador. Photos by Octavio Jiménez-Robles (**Left**) and Ignacio de la Riva (**Right**). Modified from Jiménez-Robles et al. [280].

**Call** (Figure 193): We analyzed 85 notes contained within 13 calls from two individuals (MZUTI 1476, 1684). Each call is composed by a single short note that has a duration of 3–8 (mean = 4, SD = 1) ms. Notes are clearly pulsed; the first pulse of each note has the highest amplitude. Pulses within a note have a rate of 125–333 (mean = 262, SD = 55) pulses per second. The dominant frequency of a note measured at peak amplitude is 5857–7580 (mean = 6931, SD = 472) Hz and is contained within the fundamental frequency. The fundamental frequency has a lower limit of 5685–7149 (mean = 6648, SD = 447) Hz and a higher limit of 6115–7924 (mean = 7382, SD = 491) Hz.

**Figure 193.** Call of *Rulyrana flavopunctata*, MZUTI 1684, recorded at Quebrada Pangayaku, 1645 m, Napo province, Ecuador. Air temperature = 19.4 ◦C.

**Tadpole:** Not described.

**Distribution** (Figure 194): *Rulyrana flavopunctata* is known from the Amazonian slopes of the Andes of Colombia and Ecuador at elevations between 300 and 1715 m ([22,101,281], this work). In Ecuador, this species has been recorded from localities in the provinces of Napo, Morona Santiago, Pastaza, Sucumbíos, and Tungurahua, at elevations between 720 and 1715 m (Specimens examined). In Ecuador, the potential distribution of the species is 16,369 km<sup>2</sup> within the Eastern Foothill Forest and Eastern Montane Forest regions.

**Figure 194.** Distribution of *Rulyrana flavopunctata* in Ecuador (yellow dots).

**Conservation status:** Globally, *Rulyrana flavopunctata* is listed as *Least Concern* by the IUCN [281]. In Ecuador, there are several recent reports of the species, including: Parroquia Teniente Hugo Ortíz (30 March 2007), 7.6 km W of 9 de Octubre (August 2006), 6.8 km N of Limón (21 April 2004), and Río Tayuntza (April 2011), Cordillera de los Guacamayos (April 2012), Quebrada el Plancón (March 2012), Río Hollín (August 2012), Quebrada Pangayacu (August 2012), lower slopes of Volcán Sumaco (February 2012, 2018), Narupa (June 2018). We agree with its current conservation status.

**Evolutionary relationships** (Figure 195): According to our molecular phylogeny, *Rulyrana flavopunctata* and *R*. *mcdiarmidi* are genetically indistinguishable.

**Figure 195.** Evolutionary relationships of species in the glassfrog genus *Rulyrana*. The trees were inferred using maximum likelihood and Bayesian criteria.

**Taxonomic Remarks:** Morphological differences between *Rulyrana flavopunctata* and *R*. *mcdiarmidi* are subtle (body size, skin texture, color pattern), and the two species are genetically indistinguishable (Figure 195). It is possible that *R*. *mcdiarmidi* represents a geographic variation of *R*. *flavopunctata* or that the two species have recently diverged. Information on call variation and other traits are necessary to resolve the species status of *R*. *mcdiarmidi*.

**Specimens examined:** Ecuador: *Provincia Morona Santiago:* Río Tayuntza, Cuevas de los Tayos (02.1106◦ S, 77.75185◦ W; 700 m). *Provincia Napo:* Río Hollín (0.69583 S; 77.730277 W; 1190 m), QCAZ 22360–62; Reserva Narupa (0.6848 S, 77.742 W, 1170–1228 m), ZSFQ 0374–79. *Provincia Pastaza:* tributary of Río Rivadeneira (1.3604◦ S, 77.86534◦ W), SC 34963; 9.5 km NW Mera (1.4 S, 78.166667 W; 1270 m), KU 178094; Mera (1.466667 S, 78.13331 W; 1100 m), KU 121041–46, 121049–51, 178093; Río Alpayacu, 1 km E Mera (1.466667 S, 78.08333 W; 1080 m), KU 121047; near Río Rivadeneira (1.144307 S, 77.99667 W; 982 m), QCAZ 20734–35; km 6 on San Ramón-El Triunfo road (1.35998 S, 77.86564 W; 875 m), QCAZ 37911, 33269; Sacha Yacu (1.39519◦ S, 77.72946◦ W; 1078 m), MZUTI 176. *Provincia Sucumbíos:* Bermejo No. 4, 15 km ENE Umbaqui (0.1833 N, 77.366667 W; 720 m), KU 123224.

**Localities from the literature:** *Rulyrana flavopunctata:* Ecuador: *Provincia de Napo:* San José Abajo, on the eastern slope of Volcán Sumaco (00◦32- S, 77◦24- W; 700–1000 m), AMNH 22187. *Provincia de Pastaza:* 13 km WSW Puyo (ca. 1◦34- S, 78◦06- W; 1000 m), TCWC 24032 [22].


**Common names:** English: McDiarmid's Glassfrog. Spanish: Rana de Cristal de McDiarmid. **Etymology:** The specific epithet honors Dr. Roy McDiarmid, in recognition of his contributions to the understanding of Neotropical herpetology [272].

**Identification:** *Rulyrana mcdiarmidi* can be distinguished from all other glassfrogs by having a moderate-sized body (male SVL 22.8–26.9 mm; female SVL 25.4–30.2 mm), green dorsum with diverse darker shadows and pale yellow to green spots (Figure 170), dorsal skin with warts, ventral parietal peritoneum completely white or with posterior portion translucent, thick ulnar folds, and extensive hand and foot webbing. Among centrolenids from the eastern slopes of the Andes, species with a similar color pattern include *Espadarana audax*, *Nymphargus siren*, *N*. *humboldti* sp. nov., *N*. *mariae*, *R*. *flavopunctata*, and *Teratohyla midas*. None of the species in the genus *Nymphargus* has webbing between Fingers III and IV; males of *Espadarana audax* have humeral spines; *T*. *midas* has very few yellow spots on the upper flanks and a white digestive tract; *R*. *flavopunctata* differs by being slightly smaller (male SVL 21.0–23.9 mm; female SVL 24.0–27.4 mm), having yellow dorsal spots, and lacking dorsal warts. See Taxonomic Remarks.

**Figure 196.** *Rulyrana mcdiarmidi* in life. Adult (QCAZ 32265) from 7.6 km W of 9 de Octubre, 1715 m, Morona Santiago province, Ecuador. Photos by Martín Bustamante.

**Diagnosis:** (1) Vomerine teeth present; (2) snout rounded to subtruncated in dorsal and lateral views; (3) tympanic annulus evident, oriented dorsolaterally; weak supratympanic fold; (4) dorsal skin smooth to shagreen, with numerous warts; (5) ventral skin coarsely granular; subcloacal area coarsely granular, with abundant low, flat warts; other cloacal ornamentation absent; (6) ventral parietal peritoneum white, iridophores covering entirely or almost entirely abdomen to level of groin (condition P4); pericardium white, all other visceral peritonea clear (condition V1); (7) liver lobed, lacking iridophores (condition H0); (8) humeral spine absent; (9) webbing absent between Fingers I and II, basal between II and III, moderate to extensive between outer fingers; webbing formula: II 11/2—3<sup>+</sup> III (1<sup>1</sup>/2–2−)—(1− –1<sup>+</sup>) IV; (10) webbing on feet extensive, webbing formula: I (1−–1)—(1<sup>+</sup>–11/3) II (0+–1)—(1<sup>1</sup>/3–12/3) III (1–1−)—(1<sup>2</sup>/3–2<sup>+</sup>) IV 2−—(0<sup>+</sup>–1) V; (11) thick, non-enameled, non-crenulated ulnar fold; low, short inner tarsal fold; (12) nuptial excrescences Type I in adult males; concealed prepollex; (13) first finger slightly shorter than second; (14) eye diameter larger than width of disc on Finger III; (15) in life, dorsum dark olive green with creamy yellow or light yellowish orange or light green spots, bones green; (16) color in preservative, dorsal surfaces tan grey, pale brown, or greyish lavender with di ffuse light tan spots; (17) in life, iris olive brown to brown with thin dark reticulations; (18) abundant melanophores widespread on all fingers and toes; (19) males call from rocks along streams and waterfalls; call unknown; (20) fighting behavior unknown; (21) egg clutches unknown, parental care unknown; (22) tadpoles unknown; (23) small to medium body size; SVL in adult males 22.8–26.9 mm ( X = 24.5 ± 1.139; *n* = 11), and in adult females 25.4–30.2 mm ( X = 28.0 ± 1.384; *n* = 13).

**Color in life** (Figure 196): Olive green dorsum with darker su ffusions and yellowish cream, orange, or green spots; males with smaller dorsal spots than females. Venter yellowish cream. Iris olive brown to brown with fine dark reticulations. Green bones [272].

**Color in ethanol:** Dorsal surfaces grey or greyish lavender with light spots. Venter cream. Parietal peritoneum entirely covered by iridophores to level of groin; pericardium white, all other peritonea lack white lining [272].

**Biology and ecology:** Natural history information on *Rulyrana mcdiarmidi* is scarce. It is nocturnal and males call from rocks in the spray zone of small waterfalls in Foothill Evergreen forests. Parental care is unknown [272].

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 197): *Rulyrana mcdiarmidi* is known from localities in southeastern Ecuador (Morona Santiago and Zamora Chinchipe provinces) and one in northeastern Peru (Departamento de Cajamarca) at elevations between 1150 and 1500 m ([272], this work). In Ecuador, the potential distribution of the species is 19,002 km2.

**Figure 197.** Distribution of *Rulyrana mcdiarmidi* in Ecuador (yellow dots).

**Conservation status:** *Rulyrana mcdiarmidi* is considered as *Data Deficient* by the IUCN [282]. Given the distribution of the species and the current threats if faces (i.e., habitat fragmentation, contamination by mining), we sugges<sup>t</sup> that it should be considered as *Vulnerable*, following IUCN criteria following IUCN criteria B1, B2a, B2biii.

**Evolutionary relationships** (Figure 195): *Rulyrana mcdiarmidi* and *R*. *flavopunctata* are genetically indistinguishable. The clade *mcdiarmidi* + *flavopunctata* is sister to *R*. *saxiscandens*.

**Taxonomic remarks:** Morphological differences between *Rulyrana flavopunctata* and *R*. *mcdiarmidi* are subtle (body size, skin texture, color pattern), and the two species are not separated genetically (Figure 195). Therefore, it is possible that *R*. *mcdiarmidi*represents a geographic variant of *R*. *flavopunctata*, or that the two species have recently diverged. Information on call variation and ecological requirements are necessary to resolve the species status of *R*. *mcdiarmidi*.

**Specimens examined:** *Rulyrana mcdiarmidi:* Ecuador: *Provincia de Zamora Chinchipe*: small rivulet tributary of the Jambue River, ca. 6 km S from Zamora (ca. 04◦03- S, 78◦56- W, 1150 m), on the western slope of Contrafuerte de Tzunantza, Cordillera Oriental, eastern slopes of the Andes, DFCH-USFQ D132; km 90 of the Gualaceo–Indanza–Cochay road, ca. 1 km SW of Conchay (ca. 03◦06- S, 78◦25- W; 1100 m), DFCH-USFQ AL15. *Provincia de Morona Santiago:* Río Napinaza, 6.8 km N of Limón (2.92278 S, 78.407 W; 985 m), QCAZ 27356–58; 7.6 km W of 9 de Octubre on the 9 de Octubre–Guamote road (2.225 S, 78.2904 W; 1715 m), QCAZ 32265; 2.2 km S of San Juan Bosco (3.10612 S, 79.52515 W; 1013 m), QCAZ 26443; 4.8 km N of Rosario (2.8858 S, 78.38804 W; 841 m), QCAZ 26484; 3.1 km S of San Juan Bosco (3.1467 S, 78.53559 W; 1278 m), QCAZ 26426. Peru: *Departamento de Cajamarca:* Provincia de Jaen: stream tributary of the Río Chinchipe (ca. 05◦25-16.5" S, 78◦35-23.2" W; 1250 m), extreme southwestern slope of the Cordillera del Condor, MUSM 26322–4.

**Genus** *Sachatamia* Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, and Vilà, 2009 [1].

**Etymology:** The generic name *Sachatamia* comes from the Kichwa words *sacha*, meaning forest, and *tamia*, meaning rain, and refers to the tropical rainforest occupied by the species in this genus; *Sachatamia* is feminine in gender [1].

## *Sachatamia albomaculata* (Taylor, 1949 [145]; Figures 198–200).

*Centrolenella albomaculata* Taylor, 1949 [145]. Holotype: KU 23814.

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

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

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

**Common names:** English: Cascade Glassfrog [24], White-spotted glassfrog. Spanish: Rana de Cristal de Cascada [24], Rana de cristal de puntos blancos.

**Etymology:** The specific epithet *albomaculata* is a combination of the Latin words *albus* (=white) and *macula* (=spot, stain, mark) and refers to the dorsal color pattern of the species.

**Identification:** *Sachatamia albomaculata* is the only glassfrog on the Pacific versant of the Andes that has, in life, a green dorsum with minute and large yellow spots (Figure 198), relative extensive webbing between Fingers III and IV (Figure 199), and lacks humeral spines. *Sachatamia punctulata* seems to fall within the variation of *S*. *albomaculata* (see Taxonomic Remarks).

**Figure 198.** *Sachatamia albomaculata* in life, adult male, QCAZ 40816. Ecuador: Imbabura: near Lita, ca. 500 m. Photos by M. Bustamante (**left**) and Luis A. Coloma (**right**).

**Diagnosis:** (1) Teeth on dentigerous process of the vomer present, each process bearing four to six teeth; (2) snout round to truncated in dorsal profile, and mostly truncated in lateral profile (Figure 199); (3) tympanum evident, its diameter 25.7%–34.2% of eye diameter, dorsal border of tympanic annulus covered by supratympanic fold, tympanic membrane clearly differentiated from surrounding skin; (4) dorsal skin of males and females shagreen, spicules present in males; (5) skin on venter areolate; pair of enlarged subcloacal warts, which are more evident in females; (6) anterior half of ventral parietal peritoneum covered by white iridophores, posterior half transparent (condition P2); white pericardium; translucent peritoneum covering intestines, stomach, testes, kidneys, gall bladder, and urinary bladder (condition V1); (7) liver tetralobed, two large ventral lobes covering two smaller lobes;

hepatic peritoneum transparent (condition H0); (8) humeral spines absent; (9) no webbing between Fingers I and II; webbing between other fingers as follows: II (1<sup>1</sup>/3–2+)—(3–3<sup>1</sup>/2) III (1<sup>1</sup>/3–2)—(11/3–13/4) IV; (10) foot webbing extensive: I 1—(1<sup>3</sup>/4–2) II (0+–1)—(2–2+) III (1−–11/4)—(2−–2<sup>+</sup>) IV (2−–2<sup>+</sup>)—(1–1<sup>+</sup>) V; (11) ulnar and tarsal folds low; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger I about same length as Finger II (Finger I 96.5%–103.9% length of Finger II); (14) disc of Finger III of moderate size, 41.9%–45.0% of eye diameter; (15) in life, dorsum green with small and large yellow spots; bones green; (16) in ethanol, dorsal surfaces of head, body, and limbs dark lavender with small and large whitish cream spots; venter cream; (17) iris silvery white to yellow with black reticulation; (18) melanophores usually present on dorsal surfaces of outer fingers and toes; (19) males call from the upper surfaces of leaves and rocks along streams and in spray zone of waterfalls; calls consists of a rapid high-pitched single note, "tik", with a duration of 0.001–0.002 s; the dominant frequency is at 6.1–7.1 kHz; (20) fighting behavior unknown; (21) eggs deposited on the upper surfaces of leaves or on rocks; short-term maternal care present; prolonged parental care absent; (22) tadpoles with flattened body; dark brown with violet tint dorsally; venter translucent, with few pigment spots laterally; upper jaw smoothly curved, lower jaw V-shaped; labial tooth row formula 1-2/3; (23) small body size, SVL of adult males 22.1–24.7 mm ( X = 23.6 ± 0.920, *n* = 7); SVL of adult females 26.7–29.0 mm ( X = 27.9 ± 0.891, *n* = 6).

**Figure 199.** *Sachatamia albomaculata*, adult male, QCAZ 4325. ( **A**) Head in lateral view. (**B**) Head in dorsal view. ( **C**) Hand in ventral view. Illustrations Juan M. Guayasamin.

**Color in life** (Figure 198): In Central American, two clearly di fferent color patterns have been reported, one with a green dorsum and small yellow spots; see photographs in AmphibiaWeb [283], McCranie and Wilson (81:plate 10), Savage 2002 (142:plate 198), Kubicki (24:118), and the other with small and large yellow spots (24:114). The holotype of the species, as well as Ecuadorian populations of the species, exhibit the latter coloration pattern. The venter is white on the anterior half and transparent posteriorly. The bones are green. The iris varies from white to yellow with dark reticulations.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs lavender, with small whitish cream spots or with small and large spots. White lining covers pericardium and anterior half of ventral parietal peritoneum. Liver, intestines, stomach, testes, kidneys, and gall and urinary bladders covered by translucent peritonea.

**Biology and ecology:** On the night of 1 March 1994, a male and a female (QCAZ 4324–25) were found in amplexus on a leaf about 180 cm above a stream (6 km E of Lita). At a Reserve of the Universidad de la Paz (Costa Rica), *Sachatamia albomaculata* was abundant in most of the smaller creeks that feed into the upper Río Jaris basin [284]. During the wet season, males and egg clutches were found throughout the stream channel and females deposited their eggs on the upper surfaces of leaves hanging over the water. At the beginning of the dry seasons (December 2001 and 2002), they noted a shift in the reproductive behavior and oviposition sites of this species. A large aggregation of males and egg clutches (*n* = 53; December 2002) were found on rocks in the splash area of two waterfalls; calling males and two egg clutches were also seen on a large boulder in the Jaris river. These observations suggested that in some areas *S*. *albomaculata* uses wet splash zones to prolong breeding during part of the dry season. The karyotype of *Sachatamia albomaculata* is 2N = 20 [153]. Females provide short-term parental care; prolonged parental care absent [25].

**Call:** Males typically call from the upper surfaces of leaves but have also been seen calling from plants growing on the trunks of fallen trees crossing streams, from vegetation and branches growing along rocky riparian walls and from rocky surfaces along the streams ([24,148], RWM, pers. obs.). The call consists of a rapid high-pitched single note, "tik"; the average note duration of the call is 0.001–0.002 s; the dominant frequency is 6.1–7.1 kHz [24].

**Egg masses and tadpoles:** In the oviduct and shortly after laying, the eggs are black-and-white [24,148]. In Costa Rica, during the wet season, females deposit eggs on the upper surface of riparian vegetation; during the dry season, egg clutches are deposited in the splash area of waterfalls [284]. The eggs are distinctive in that a space without eggs often occurs in the center of the mass ([24], RWM, pers. obs.). Egg clutches have 37–81 eggs (mean = 57 ± 18) [147]. The tadpole of *S*. *albomaculata* was described in detail by Hoffmann [147] and we present a summary of his description below. Embryos in stage 22 are tan in color, whereas hatchlings at stage 25 are dark greyish brown, a coloration that extends to the flanks of the body and onto the venter. Ventral coloration is lighter because of the yellow yolk visible through the skin. The larvae of this species are among the most densely pigmented centrolenids in Costa Rica. While hatchlings are consuming their yolk reserve, the oral disc remains incomplete. More developed tadpoles (stages 25–37) have the typical dorsoventrally compressed body shape, which, together with the larvae of *S*. *ilex* and *T*. *spinosa*, are the most compressed of Costa Rican centrolenids. The oral disc is very similar to that of *S*. *ilex;* it is not emarginate and has a single row of 24–40 marginal papillae with a distinctly broad anterior labium (dorsal gap), a trait that is also present in *S*. *ilex*; broad dorsal gap; papillae are often flattened, irregular, and sometimes absent. The upper jaw is smoothly curved, and the lower jaw is broad and V-shaped. The LTRF probably is 2(2)/3 but most tadpoles lack the A-1 tooth row; if present, the A-2 is short and consists mostly of a few teeth in short rows along the lateral margins of the upper jaw. Three ridges are evident on the lower labium and all or some of them bear teeth [147]. Whether these irregularities in the labial tooth rows result from laboratory rearing or are typical of the species remains to be demonstrated.

**Distribution** (Figure 200): *Sachatamia albomaculata* is known from humid lowlands and premontane slopes from north-central Honduras to western Colombia and Ecuador at elevations between 20 and 1500 m ([130,148,160], this work). In Ecuador, this species has been reported from the provinces of Esmeraldas and Imbabura at elevations below 700 m. In Ecuador, the potential distribution of the species is 22,608 km<sup>2</sup> within the Chocoan Tropical Forest and the Western Foothill Forest regions.

**Figure 200.** Distribution of *Sachatamia albomaculata* in Ecuador (yellow dots).

**Conservation status:** Listed as *Least Concern* by the IUCN in view of its wide distribution and presumed large population [285]. In Ecuador, the species is known mainly from a few reports in the Chocó, an area with the highest deforestation rate (because of oil palm and wood extraction) and spreading mining. We sugges<sup>t</sup> that the species should be considered as *Endangered* in Ecuador, following IUCN criteria A2, B2a, B2(iii).

**Evolutionary relationships** (Figure 195): *Sachatamia albomaculata* is sister to a clade formed by *S*. *punctulata* + *S*. *electrops*.

**Taxonomic Remarks:** *S*. *albomaculata*, as currently defined, has two clear dorsal color patterns: (a) Dorsum with small yellow spots, (b) dorsum with small and large yellow spots (see *Color in life* section). Two scenarios can explain the observed polymorphism: Two or more species occur within what is now recognized as *S*. *albomaculata*, or *S*. *albomaculata* has a natural polymorphic coloration. To test the two hypotheses, it is necessary to plot the occurrence of the patterns, document variation and possible sympatry, and obtain acoustic data and molecular samples of the different color patterns for comparative analyses. It would also be worth obtaining samples from the Colombian species *S*. *punctulata*, which has a color pattern identical to pattern (a) of *S*. *albomaculata*.

**Specimens examined:** *Sachatamia albomaculata:* Colombia: *Departamento de Antioquia:* Dabeiba, Río Amparradó, Quebrada Iotó, 805 m, ICN 10685–87. Costa Rica: Limón: Los Diamantes, one mile south of Guápiles, KU 23814 (holotype). Ecuador: *Provincia de Imbabura:* 6 km SE of Lita (0◦47-41" N, 78◦25-43" W), QCAZ 4324–25. *Provincia de Esmeraldas:* Estero Vicente, an affluent of the Río San Miguel 0◦47-32" N, 79◦11-52" W, 225–275 m), QCAZ 11369–70; Reserva Biológica Canandé (0◦27-4" N, 79◦08-45" W, 700 m), MECN 2618–19; 2.1 km E of Durango (1.02477 N, 78.61746 W; 284 m), QCAZ 32172–73; 5 km E of Lita, on the Lita–Ibarra road (0.84773 N, 78.42175 W), QCAZ 40816; Reserva Itapoa (0.513◦ N, 79.134◦ W, 321 m), MZUTI 3013.

**Localities from the literature:** *Sachatamia albomaculata:* Ecuador: *Provincia de Esmeraldas:* Estación Biológica Bilsa (0.359 N, 79.701 W) [160].

*Sachatamia ilex* (Savage, 1967 [202]; Figures 201–203).

*Centrolenella ilex* Savage, 1967 [202]. Holotype: LACM 25205.

Type locality: "Costa Rica: Provincia de Limón: Cantón de Limón: Alta Talamanca: 16 km SW Amubri, on Río Lari, 300 m"; corrected to 14 km SW Amubri, Río Lari, Cantón de

Talamanca, Provincia de Limón, 300 m by Savage, 1974 [183].

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

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

**Common names:** English: Holly's Glassfrog. Spanish: Rana de Cristal de Holly, Rana de Cristal Fantasma.

**Etymology:** The specific name *ilex* honors Priscilla Hollister "Holly" Starrett (*Ilex* is the genus of the holly trees), for her contributions to centrolenid systematics [202].

**Identification:** *Sachatamia ilex* is distinguished from other glassfrogs by having, in life, a uniform dark green dorsum, a white to pale yellow or greenish–yellow iris with thick and contrasting black reticulations (Figure 201), and a moderate body size (male SVL 27.0–29.0 mm; female SVL 28.0–34.0 mm). Adult males have a pointed humeral spine (Figure 201) that is embedded in the arm musculature (not easily visible externally). *Sachatamia ilex* closely resembles spotless populations of *Espadarana prosoblepon* (Figure 95) and *E*. *callistomma* (Figure 89), the males of which have externally visible humeral spines; most individuals of *E*. *prosoblepon* lack the thick black reticulation of the iris that is evident in *S*. *ilex*. Other species with uniform green dorsal coloration that could be confused with *S*. *ilex* are *"Cochranella" prasina*, which lacks webbing between Fingers II, III, and IV (present in *S*. *ilex*), and *Teratohyla spinosa*, which is much smaller in body size (SVL < 21 mm) and lacks humeral spines.

**Figure 201.** *Sachatamia ilex* in life. ( **A**,**B**) Adult male, Alto Tambo, Esmeraldas province, 620 m, QCAZ 47193. ( **C**) Adult male, Alto Tambo–El Placer, Esmeraldas province, 706 m, QCAZ 48338. ( **D**) Adult male, Alto Tambo-El Placer, Esmeraldas province, not collected. Photos by L. A. Coloma.

**Diagnosis:** (1) Each dentigerous process of the vomer bearing three to five teeth; (2) snout truncated in dorsal and lateral profiles; (3) tympanum oriented almost vertically, relatively small, its diameter 26.1%–28.7% of eye diameter; supratympanic fold conspicuous, covering dorsal margin of tympanic annulus; tympanic membrane pigmented, but clearly di fferentiated from surrounding skin; (4) dorsal surfaces of males and females smooth to shagreen, lacking spicules; (5) venter areolate; lacking pair of enlarged subcloacal warts; (6) anterior two-thirds of ventral parietal peritoneum white, posterior third transparent (condition P3); white pericardium; translucent peritoneum covering intestines, stomach, kidneys, testes, gall bladder, and urinary bladder (condition V1); (7) liver tetralobed, covered by transparent peritoneum (condition H0); (8) in males, humeral spines present, but embedded in arm

musculature; (9) webbing absent between Fingers I and II, reduced between Fingers II and III, extensive between III and IV; webbing formula II (1<sup>1</sup>/2–2−)—(3<sup>+</sup>–31/4) III (1<sup>1</sup>/3–2)—(1<sup>+</sup>–11/3) IV; (10) webbing formula on foot I (1–1<sup>1</sup>/4)—(11/2–2−) II (1–1+)—(1<sup>3</sup>/4–2) III (1–1<sup>+</sup>)—(2−–2<sup>+</sup>) IV (1<sup>1</sup>/2–2+)—(1–1+) V; (11) ulnar fold absent or low and inconspicuous; inner tarsal fold low, short; outer tarsal fold absent; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger I slightly shorter or about same length as Finger II (Finger I 97.2%–100.0% of Finger II); (14) disc of Finger III relatively narrow, its width about 34.0%–45.9% of eye diameter; (15) in life, dorsum uniform green; upper lip white; bones green; (16) in preservative, dorsum uniform lavender; (17) iris, in life, silvery white to greenish yellow with thick, contrasting black reticulation; (18) melanophores usually present on fingers and toes, except sometimes absent from Toes IV and V; (19) males call from upper sides of leaves near streams; each call is frequency modulated and consists of a single note with a duration of 0.077–0.086 s; dominant frequency starts at 6562–6937 Hz, rises to 7406–7500 Hz, and drops to 6562–6843 Hz; (20) males fight upside down, grasping one another venter to venter; (21) brown eggs deposited on upper side of leaves; parental care unknown; (22) tadpoles with flattened body shape; blackish brown dorsal coloration, venter translucent; upper jaw in form of broad arch; lower jaw V-shaped; labial tooth row formula 0/2-3; (23) medium body size; SVL of adult males 27.0–29.0 mm; SVL of adult females 28.0–35.0 mm.

**Figure 202.** *Sachatamia ilex*. ( **A**) Humeral spine of adult male, LACM 72910, not to scale. (**B**) Head in lateral, paratype. ( **C**) Hand (left) and foot (right) in ventral view, paratype, not to scale. ( **D**) Dorsal view of paratype. ( **A**) Modified from Guayasamin et al. [1]. (**B**–**D**) Modified from Savage [202].

**Color in life** (Figure 201): Dorsum uniform dark green; throat and venter cream white; dull yellow hands and feet; upper lip with thin white line. White parietal peritoneum covers anterior two-thirds of venter; clear visceral and hepatic peritonea. Bones green. Iris silvery white to greenish yellow with thick, contrasting black reticulations.

**Color in ethanol:** Dorsal surfaces of head and body uniform lavender, although faint cream spots might be visible in some individuals [202]; limbs creamy lavender dorsally; margin of upper lip white or cream. White parietal peritoneum covering anterior two-thirds of venter; pericardium white; clear peritoneum covering liver, intestines, stomach, kidneys, and urinary bladder. In males, nuptial pad creamy white.

**Biology and ecology:** Nocturnal and arboreal. On 10 November 1999, at Hacienda La Joya (northwestern Ecuador), one male was calling, and two females were gravid, suggesting reproductive activity [130]. Kubicki [24] described and illustrated the combat behavior of *Sachatamia ilex* as follows: Males hang vertically from vegetation by the tips of their toes and grasp each other chest to chest in a type of arm lock; males may hang in this position for several minutes to hours, and try to dislodge the other male's feet, so that he loses grip of the vegetation. During this behavior, the two males produce softer *preep* calls. Parental care is unknown.

**Call:** The following call description is based on a recording of one male of *Sachatamia ilex* made by Elicio Tapia on 17 May 2010 at Reserva Otokiki (0.91104◦ N, 78.57369◦ W; 706 m), Provincia de Esmeraldas, Ecuador. Males call from the upper surfaces of leaves. Ten calls where emitted during a five-minute interval; the call consists of a single, pulsed note; the note ends with a series of pulses the amplitude of which is conspicuously lower than the pulses at the beginning of the note. Each note has a duration of 0.077–0.086 s ( X = 0.08 ± 0.033, *n* = 6). The note is frequency modulated; the dominant frequency at the beginning of the note (pulsed section) is at 6562–6937 Hz ( X = 6640 ± 150.2, *n* = 6); during the tonal section of the note, the dominant frequency increases to 7406–7500 Hz ( X = 7422 ± 38.4, *n* = 6); finally, it drops to 6562–6843 Hz ( X = 6718 ± 128.0, *n* = 6). The maximum value of the dominant frequency is always during the first pulsed section of the note.

**Egg masses and tadpoles:** Brown eggs are deposited as a single layer on the upper leaf surface of plants overhanging streams; sometimes a mass has a slight hole in the center, resembling a doughnut, because they might rotate around from the middle while laying eggs; egg clutches have 12–25 eggs [24]. Species of *Drosophila* have been observed to parasitize egg clutches of *S*. *ilex* [147]. The tadpole description presented below is a summary obtained from Hoffmann [147]. When hatching, larvae have a dark greyish–brown dorsal coloration; the body is nearly circular in cross-section with a length of about 12 mm; it is longer than other Costa Rican centrolenid hatchlings. In tadpoles that are living off their yolk reserves, oral discs are incomplete. As development continues (Gosner stages 28–40) the body becomes flattened and a dense dorsal pigmentation develops, giving them a blackish–brown ground tone; there is a slightly violet coloration because of the vascularized underlying organs. Lateral body pigmentation decreases ventrally, and the tail coloration is the same as the dorsum, decreasing gradually towards its ventral side. The snout is rounded in dorsal view and acutely angled in lateral view. The anteroventral oral disc has marginal papillae only on the lateral and ventral margins; the anterior labium forms the upper lip and is uniformly broad; the disc is not emarginate. Because of the lack of an anterior disc area, the labial tooth rows A-1 and A-2 are absent. P-1 is usually completely developed, P-2 is shorter not always completely developed, and P-3 lacks teeth or has very few of them [147].

**Distribution** (Figure 203): *Sachatamia ilex* is known from localities in eastern Nicaragua, Costa Rica, western Panama, western Colombia, and western Ecuador at elevations from sea level up to 1420 m ([130,148,217,284,285], this work). In Ecuador, *Sachatamia ilex* has been found in Esmeraldas and Pichincha provinces at elevations between 150 and 800 m ([130], this work). In Ecuador, the potential distribution of the species is 12,669 km<sup>2</sup> within the Chocoan Tropical Forest and the Western Foothill Forest regions.

**Figure 203.** Distribution of *Sachatamia ilex* in Ecuador (yellow dots).

**Conservation status:** *Sachatamia ilex* is globally listed as *Least Concern* [285]. Kubicki (2007) mentioned that *S*. *ilex* is fairly common in much of its range along the Caribbean drainage in Costa Rica. In Ecuador, habitat reduction is a serious threat for this and other species that inhabit the Chocó Ecoregion; thus, at the local level, we sugges<sup>t</sup> that the species should be considered as *Endangered*, following IUCN criteria B2a, B2(iii).

**Evolutionary relationships** (Figure 204): Molecular evidence places *Sachatamia ilex* and *S*. *orejuela* as sister species.

**Figure 204.** Evolutionary relationships of species in the glassfrog genus *Sachatamia*. The trees were inferred using maximum likelihood and Bayesian criteria.

**Specimens examined:** *Sachatamia ilex:* Colombia: *Departamento de Antioquia:* Dabeiba, Río Amparradó, Quebrada Iotó, 805 m, ICN 10625–29, 10630 (C&S), 10631–32. Costa Rica: *Provincia de Limón:* Costa Rican Amphibian Research Center, UCR 16861. Ecuador: *Provincia de Esmeraldas:* Reserva Biológica Canandé (0.45◦ N, 79.14◦ W, 700 m), MECN 2620–26; Río Tululbí (1.0634◦ N, 78.59397◦ W, 189 m), MECN 3199–03; Río Verde (0.92◦ N, 78.63◦ W; 300 m), MECN 3204; Río Verde (1.183◦ N, 78.7166◦ W; 300 m), MECN 3199; recinto Ventanas (0.89816◦ N, 78.6175◦ W. 200 m), MECN 3204; Reserva Otokiki (0.91104◦ N, 78.57369◦ W; 706 m), QCAZ 48338; 4 km N of Durango (1.44307◦ N, 77.99667◦ W; 253 m), QCAZ 33325; Río La Carolina (0.70449◦ N, 78.20115◦ W; 500 m), QCAZ 35363; 2.1 km E Durango (1.02477◦ N, 78.61746◦ W; 284 m), QCAZ 32158; 4 km W Durango (1.02348◦ N, 78.19296◦ W; 238 m), QCAZ 33057. *Provincia de Santo Domingo de los Tsáchilas:* 4 km NW La Florida, Finca Gloria (0.25694◦ N, 79.0538◦ W; 896 m), QCAZ 19881; La Florida, near Alluriquín (0.2836◦ N, 79.0188◦ W), QCAZ 13055. *Provincia de Pichincha:* Puerto Quito (0.1167◦ N, 79.2667◦ W; 150 m), KU 221613; Hacienda La Joya, km 109 of the Calacalí–Nanegalito–P.V. Maldonado Road, next to the town of San Vicente de Andoas (0.083◦ N, 78.983◦ W; 750–800 m), DFCH-USFQ D260–61; near Pedro Vicente Maldonado (0.10421◦ N, 79.10279◦ W; 544 m), QCAZ 35429. Panama: *Comarca San Blas:* Camp Summit, 400 m, KU 116464.

*Sachatamia orejuela* (Duellman and Burrowes, 1989 [86]; Figures 205 and 206).

*Centrolenella orejuela* Duellman and Burrowes, 1989 [86]. Holotype: KU 145081. Type locality: "between El Tambo and La Costa, 800 m, Departamento de Cauca, Colombia." *Cochranella orejuela*—Ruiz-Carranza and Lynch, 1991 [6].

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

*Sachatamia orejuela*—Twomey, Delia, and Castroviejo-Fisher, 2014 [19].

*Teratohyla sornozai*—Cisneros-Heredia, Yánez-Muñoz, and Ortega-Andrade, 2009 [286].

Synonymy by Cisneros-Heredia, Yánez-Muñoz, and Ortega-Andrade, 2010 [287].

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

**Etymology:** The specific name *orejuela* is a patronym for the Orejuela family (Jorge, Anamaría, and Tomás), who, at the time when the study was conducted, resided, and administered the Reserva La Planada, Colombia [86].

**Identification:** *Sachatamia orejuela* is mainly recognized by having a uniformly olive-green dorsum (Figure 205). In the Pacific versant of the Andes, only the following species have a uniformly green dorsum: *Nymphargus prasinus*, *Sachatamia ilex*, *S*. *orejuela*, *Teratohyla spinosa*, and some populations of *E*. *prosoblepon*. From these species, *Sachatamia orejuela* is distinguished by the absence of humeral spines (spines present in males of *E*. *prosoblepon* and *S*. *ilex*), relatively large body size (SVL 27.3–33.8 mm in *S*. *orejuela;* SVL < 22.0 mm in *T*. *spinosa*), and extensive webbing between Fingers III and IV (webbing absent or basal in *Nymphargus prasinus*). Additionally, *S*. *orejuela* is found in a very specific microhabitat, on rocks nearby waterfalls.

**Figure 205.** *Sachatamia orejuela* in life. Adult male, QCAZ 45993. Locality near Río Aguas Verdes, 670 m, Imbabura province, Ecuador. Photos by Luis A. Coloma.

**Diagnosis:** (1) Each vomer with two to five teeth; (2) snout truncated in dorsal aspect, slightly protruding in lateral profile; (3) tympanum small, visible, its diameter 18.5%–25.6% of eye diameter; tympanic membrane pigmented as surrounding skin; supratympanic fold conspicuous; (4) dorsal surfaces finely shagreen, with small spicules evident in sexually active males; (5) ventral surfaces of body areolate; pair of enlarged subcloacal warts absent; in males, cloacal region with minute spinules; (6) anterior half of the parietal peritoneum white (condition P2); white pericardium, translucent visceral peritoneum (condition V1); (7) tetralobed liver covered by transparent peritoneum (condition H0); (8) humeral spines absent; (9) webbing between Fingers I and II basal, moderate between Fingers II and III, and extensive between Fingers III and IV; webbing formula for fingers: II (1–1<sup>1</sup>/2)—(3−–3<sup>+</sup>) III (1<sup>+</sup>–2−)—(1–1<sup>+</sup>) IV; (10) webbing on foot extensive: I 0<sup>+</sup>—(1−–1) II 0<sup>+</sup>—(0<sup>+</sup>–1) III 0<sup>+</sup>—(0<sup>+</sup>–1) IV (1<sup>1</sup>/3–11/2)—0 V; (11) ulnar and tarsal folds absent; (12) nuptial pad Type I, concealed prepollex; (13) Finger I slightly longer than Finger II (Finger II about 95%–98% length of Finger I); (14) disc of Finger III large, its width about 62%–70% of eye diameter; (15) in life, dorsum uniform dark olive green; (16) in preservative, dorsum uniform dull grey; (17) iris, in life, dark grey with yellow ring around pupil; (18) melanophores on dorsal surfaces of all fingers and toes; (19) males call from rocks along or within streams; call unknown; (20) fighting behavior unknown; (21) eggs deposition site unknown; parental care unknown; (22) tadpoles unknown; (23) medium body size; SVL 27.3–28.3 mm (*n* = 3) in males; 29.6–33.8 mm (*n* = 6) in females.

**Color in life** (Figure 205): Uniform dark olive-green dorsum, greyish green flanks; throat translucent, with light green hue; venter whitish cream. Bones green. Iris dark grey to brown ([86], this work).

**Color in ethanol:** Dorsum, including hands, feet, and webbing dull grey [86]. Anterior half of venter cream white, posterior half translucent; white pericardium, translucent visceral peritonea, and transparent hepatic peritoneum (liver brown).

**Biology and ecology:** During the night, *Sachatamia orejuela* is mostly found on rocks along steep stream banks or within the stream in the spray zone of cascades ([86,288], this work). As other species adapted to spray zones (e.g., *Centrolene geckoidea*, *C*. *paezorum*, *"Centrolene" petrophilum*, *"Centrolene" medemi*), hand and foot webbing in *Sachatamia orejuela* is relatively extensive when compared to glassfrogs not found in this microhabitat. Spiders of the genus *Clubiona* have been observed to prey on juveniles of *S*. *orejuela* [289]. Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 206): *Sachatamia orejuela* is known from four localities in southern Colombia and four localities in northern Ecuador ([86,101,288,290], this work). All localities are on the Pacific flank of the Andean Cordillera Occidental at elevations between 500 and 1250 m, within the Eastern Foothill Forest region. In Ecuador, the potential distribution of the species is 27,514 km2.

**Conservation status:** *Sachatamia orejuela* is classified as *Least Concern* by the IUCN [291]. However, in Ecuador, the habitat of the species is fragmented and threatened by logging and mining. We sugges<sup>t</sup> that the species should be considered as *Endangered*, following IUCN criteria B2a, B2(iii).

**Evolutionary relationships** (Figure 204): *Sachatamia orejuela* and *S*. *ilex* are inferred as sister species. Herein, we modify the phylogenetic definition of the genus *Sachatamia* [1] as follows: *Sachatamia* is the clade stemming from the most recent common ancestor of *Centrolenella albomaculata* Taylor 1949 [145], and *Cochranella orejuela* Duellman and Burrowes 1989 [86].

**Taxonomic Remarks:** *Teratohyla sornozai* was synonymized with *Sachatamia orejuela* by Cisneros-Heredia et al. [287]. Examination of the type series of *T*. *sornozai* showed that all differences that separated it from *S*. *orejuela* were expressions of ontogenetic and intraspecific variation of the later.

**Figure 206.** Distribution of *Sachatamia orejuela* in Ecuador (yellow dots).

**Specimens examined:** *Sachatamia orejuela:* Colombia: *Departamento de Cauca:* between El Tambo and La Costa, 800 m, KU 145081 (holotype), 145080 (paratype). Ecuador: *Provincia de Esmeraldas:* Reserva Biológica Canandé (0.5299 N, 79.0354 S; 550 m), DHMECN 2634. *Provincia de Imbabura:* Zona de amortiguamiento de Reserva Cotacachi Cayapas, near Río Aguas Verdes (0.331010◦ N, 78.93152◦ W; 670 m), QCAZ 45993; Reserva Río Manduriacu (0.3108 N, 78.8576 S; 1230 m), JMG 1581. *Provincia de Pichincha:* Bosque Protector Mashpi (00◦10-2.34" N 78◦52-2.32" W; 1200 m), DHMECN 04309; Río Chalpi (00◦13-32.38" N 78◦51-28.87" W; 615 m), DHMECN 04551; Río Anope (00◦12-45.54" N 78◦48-58.34" W; 1080 m), in the surroundings of the town of Saguangal, DHMECN 04552. *Provincia de Santo Domingo de los Tsáchilas:* Trail within Hotel Tinalandia (0.2727 S, 79.079 W), QCAZ 45452.

**Localities from the Literature:** Colombia: *Departamento de Nariño:* Pialapí, 1250 m, IND-AN 1520–21, LP248 [86]; Departamento Valle del Cauca: Campo Alegre (IUCN, 2010). Ecuador: *Provincia de Esmeraldas:* Reserva Biológica Canandé (0.306 N, 79.138 W; 550 m), DHMECN 2634. *Provincia de Pichincha:* Bosque Protector Mashpi (0.167 N, 78.867 W; 550 m), DHMECN 4308. *Provincia de Imbabura:* Stream tributary of Río Naranjal (0.351 N, 78.917 W; 750 m), DHMECN 3522 ([286] as *Teratohyla sornozai*).

## **Genus** *Teratohyla* Taylor 1951 [15].

**Etymology:** The generic name *Teratohyla* is derived from the Greek *teras*, meaning monster, marvel or wonder, and the word *Hyla*, traditionally associated with treefrogs. The origin of the frog name *Hyla* is based on the mythological Greek boy *Hylas* and, although the boy's name is masculine, it has been unambiguously treated as feminine by amphibian systematists [292].

### *Teratohyla amelie* (Cisneros-Heredia and Meza-Ramos, 2007 [253]; Figures 207–209).

*Cochranella amelie* Cisneros-Heredia and Meza-Ramos, 2007 [253]. Holotype: DHMECN 3066. Type locality: "Comunidad de Oglán, Cantón Arajuno, Provincia de Pastaza, República del Ecuador(01◦18-65"S,77◦42-41"W,600melevation)."

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

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

**Etymology:** The specific name is for *Amelie*, protagonist of the movie "Le Fabuleux Destin d'Amélie Poulain", a movie where little details play an important role in the achievement of *joie de vivre*; like the important role that glassfrogs and all amphibians and reptiles play in the health of our planet. [253].

**Identification:** *Teratohyla amelie* differs from most species inhabiting the Amazonian lowlands and Amazonian slopes of the Andes by having a completely transparent ventral parietal peritoneum (Figure 207). In Ecuador, only species in the genus *Hyalinobatrachium* and *Chimerella mariaelenae* share the transparent parietal peritoneum character. *Chimerella mariaelenae* differs from *T*. *amelie* by having a green dorsum with many dark lavender punctuations and scattered larger dark spots in life (dorsum uniform green in *T*. *amelie*). Also, adult males of *C*. *mariaelenae* have a small humeral spine (absent in *T*. *amelie*). *Teratohyla amelie* differs from all species in the genus *Hyalinobatrachium* (characteristics in parentheses) by having a uniform green dorsum in life that turns lavender in preservative (green dorsum with small yellow spots in life; cream in preservative), by depositing its egg clutches on the upper surfaces of leaves (underside of leaves), and by having a Type I nuptial pad (TypeV pad in most *Hyalinobatrachium*). Two other species that are not found in Ecuador, *T*. *pulverata* and *Vitreorana antisthenesi*, also have a transparent ventral peritoneum and deposit eggs on the upper surfaces of leaves. *Teratohyla pulverata*, however, is restricted to Central America and areas west of the Andes (*T*. *amelie* is endemic to Amazonian lowlands of Ecuador and Peru), and differs from *T*. *amelie* by having a sloping snout in lateral view and a green dorsum with white spots in life. The Venezuelan *V*. *antisthenesi* can be differentiated from *T*. *amelie* by the presence of vomerine teeth, a green dorsum with light spots, and being larger (21.4–26.2 mm SVL in adult males of *V*. *antisthenesi* [106]). *Teratohyla midas* is also found on the Amazonian lowlands, but it has yellow dorsal spots and a venter that is white anteriorly and transparent posteriorly.

**Diagnosis:** (1) Vomerine teeth absent; (2) snout truncated in dorsal view and rounded in lateral view; (3) tympanic annulus evident, oriented dorsolaterally with posterior inclination; very weak supratympanic fold above the tympanum; (4) dorsal skin shagreen; (5) ventral and subcloacal surfaces granular, pair of enlarged subcloacal warts; (6) parietal peritoneum completely transparent (lacking iridophores, condition P0), all visceral peritonea covered with iridophores (condition V5); (7) liver white, bulbous (condition H2); (8) humeral spine absent; (9) webbing on hand absent between Fingers I and II, moderate to extensive between outer fingers: II (1–1+)—(3–3+) III (1<sup>1</sup>/2–2−)—(1–1<sup>1</sup>/2) IV; (10) webbing on feet extensive: I (0–1)—(1<sup>1</sup>/2–2−) II (1−–1)—2− III (1−–1<sup>+</sup>)—(2−–21/4) IV (2−–21/3)—(0−–1−) V; (11) low non-enameled ulnar fold; low non-enameled inner tarsal fold; (12) nuptial excrescences present, Type I; concealed prepollex; (13) first finger longer than second, (14) eye diameter larger than width of disc on Finger III; (15) color in life, uniform green to bluish–green dorsally; (16) in preservative, dorsal

surfaces uniform lavender; (17) in life, iris cream brown; (18) numerous melanophores on all fingers and toes; (19) males call from upper sides of leaves; each call consists of single note with three or four pulses; dominant frequency at 4984–7085 Hz, reaching maximum frequency at 5906–6023 Hz; (20) fighting behavior unknown; (21) egg clutches deposited in single layer on upper surfaces of leaves next to streams; males call from tops of leaves next to their clutches; parental care unknown; (22) tadpoles unknown; (23) minute body size; SVL in adult males 18.1–19.3 mm (X = 18.8 ± 0.47; *n* = 6); females unknown.

**Figure 207.** *Teratohyla amelie* in life. (**Left**): Adult male, QCAZ 37920, Ecuador, Pastaza province, stream tributary of Río Lliquino, 350 m (photo by Martín Bustamante). (**Right**): Adult male, QCAZ 47204, Ecuador, Pastaza province, Reserva Ecológica Río Anzu, 1200 m. Photo by Lucas Bustamante/Tropical Herping.

**Color in life** (Figure 207): Uniform green to bluish–green dorsum, lacking flecks or spots. Greenish throat, other ventral surfaces transparent. White heart and viscera visible ventrally. Green bones. Iris cream brown.

**Color in ethanol:** Dorsal surfaces uniformly lavender. Ventral surfaces cream. Ventral parietal peritoneum without iridophores. Iridophores covering heart, digestive tract, and liver.

**Biology and ecology:** *Teratohyla amelie* is a recently described species and little is known about its natural history. It is nocturnal and males call from and pairs deposit single layer clutches on the upper surfaces of *Heliconia* leaves about 80 cm above small streams in pristine Lowland Evergreen forests. Parental care is unknown.

**Call** (Figure 208): The description provided below is based on a recording obtained by Diego Paucar at a tributary of Río Lliquino (1.45295◦ S; 77.443◦ W, 350 m; QCAZ 38779), Provincia Pastaza, Ecuador, on 15 May 2008. Calls are emitted every 10–30 s (X = 17.8 s ± 7.55, *n* = 8). Each call consists of a single pulsed note that lasts 0.011–0.014 s (X = 0.013 ± 0.001, *n* = 10); each call has three to four pulses (X = 3.5 ± 0.515, *n* = 10). The dominant frequency is between 4984–7085 Hz (*n* = 10), with its maximum frequency at 5906–6023 Hz; a first harmonic is visible at 10,923–12,474 Hz (*n* = 10); a second harmonic is visible at 16,815–19,120 Hz (*n* = 10).

**Figure 208.** Call of *Teratohyla amelie*(QCAZ 38779) from a tributary of Río Lliquino (1.45295◦ S; 77.443◦ W, 350 m; QCAZ 38779), Provincia Pastaza, Ecuador.
