**Tadpole:** Not described.

**Distribution.** *Hyalinobatrachium adespinosai* is only known from the type locality: San Jacinto River (1.3447 S, 78.1814 W; 1795 m asl), Tungurahua Province, Ecuador (Figure 100) [194].

**Figure 100.** Distribution of *Hyalinobatrachium adespinosai* in Ecuador (yellow spot).

**Evolutionary relationships**. *Hyalinobatrachium adespinosai* is sister to the Peruvian *H*. *anachoretus* [194]. Since the species was recently described, it is not included in the *Hyalinobatrachium*

tree shown in Figure 101. The most closely related species to *H*. *adespinosai* share several morphological traits, including a red heart exposed ventrally (*H*. *adespinosai* + *H*. *anachoretus* + *H*. *pellucidum* + *H*. *yaku)* [194].

**Figure 101.** Evolutionary relationships among species in the genus *Hyalinobatrachium*, inferred using maximum likelihood and Bayesian criteria.

**Conservation status:** *Hyalinobatrachium adespinosai* has not been evaluated by the IUCN. However, Guayasamin et al. [194] suggested that it should be considered as *Data Deficient*.

**Specimens examined.** *Hyalinobatrachium adespinosai:* Ecuador: *Provincia de Tungurahua:* San Jacinto River (1.3447 S, 78.1814 W; 1795 m asl), ZSFQ 1647–48, 1650–52 (type series).

	- latitud N y 76◦20- W, 1030 m.s.n.m.".

*Hyalinobatrachium aureoguttatum*—Ruiz-Carranza and Lynch, 1991 [6].

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

**Etymology:** The specific name *aureoguttatum* is derived from the Latin words *aureus* (gold) and *guttatus* (dappled, speckled, spotted), referring to the dorsal pattern of the species [195].

**Identification:** *Hyalinobatrachium aureoguttatum* is easily distinguished from most glassfrogs by having a completely transparent ventral parietal peritoneum, a white bulbous liver, white visceral peritoneum, and by the absence of humeral spines. It differs from all other *Hyalinobatrachium* by having, in life, large yellow spots on the dorsum (Figure 102). The most similar and related species is *H*. *valerioi*, from which is differentiated mainly by having large dorsal yellow spots produced by xanthophores and iridophores (absent in *H*. *valerioi;* see Figure 121).

**Figure 102.** *Hyalinobatrachium aureoguttatum* in life. (**Left and center**) Adult male, QCAZ 45365; photos by L. A. Coloma. (**Right**) Adult male, QCAZ 32068; photo by Martín Bustamante.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout truncated in dorsal and lateral profiles (Figure 103); (3) tympanum of moderate size (tympanum diameter 30%–34% of eye diameter), with dorsolateral orientation and posterior inclination, supratympanic fold low, tympanic membrane clearly differentiated from surrounding skin; (4) dorsal surfaces smooth to shagreen, lacking spicules; (5) venter smooth; lacking pair of enlarged subcloacal warts; (6) ventral parietal peritoneum transparent (condition P0); pericardium polymorphic, with or lacking iridophores; white peritonea covering intestines, stomach, and testes; transparent peritoneum covering kidneys, and urinary and gall bladders (Figure 102, Condition V5–V6); (7) liver bulbous, covered by white peritoneum (condition H2); (8) humeral spines absent; (9) basal webbing between Fingers I and II, extensive webbing between outer finger: I (2–2<sup>1</sup>/4)—(2–21/4) II (1−–1<sup>+</sup>)—(2–3<sup>+</sup>) III (1–2+)—(1–1+) IV; (10) foot webbing extensive: I (0+–1)—(1–1<sup>2</sup>/3) II (0+–1)—(1–1<sup>1</sup>/2) III (0−–1)—(1<sup>1</sup>/2–2) IV (1<sup>1</sup>/2–2<sup>+</sup>)—(1−–1) V; (11) ulnar and tarsal folds absent; (12) concealed prepollex; in males, nuptial pad Type III; (13) Finger I longer than Finger II (Finger II about 87%–95% length of Finger I); (14) disc of Finger III of moderate size, 35%–56% of eye diameter; (15) in life, dorsum greenish yellow with two to seven large yellow spots, and with or without brown flecks; ventral parietal peritoneum transparent, pericardium polymorphic (white or transparent), gastrointestinal peritoneum white; bones white; (16) in ethanol, dorsum cream with large white spots, and with or without brown flecks; (17) in life, iris white to yellow, with minute dark lavender flecks around the pupil or forming an horizontal stripe; (18) fingers and toes lacking melanophores, except for few melanophores on proximal portions of Toes IV and V; (19) males usually call from the underside of leaves; call undescribed; (20) fighting behavior unknown; (21) eggs deposited on the underside of leaves; prolonged parental care provided by males; maternal care absent; (22) oral apparatus of tadpoles with an emarginate disc; M-shaped upper jaw sheath; tooth row formula 2(2)/3; (23) minute body size; in males, SVL 20.4–24.0 mm (X = 21.8 ± 0.631, *n* = 36); in females, SVL 22.9–23.9 mm (X = 23.3, *n* = 3).

**Figure 103.** *Hyalinobatrachium aureoguttatum*, adult male, QCAZ 27429. (**A**) Head in lateral view. (**B**) Head in dorsal view. (**C**) Hand in ventral view. (**D**) Foot in ventral view. Illustrations by Juan M. Guayasamin

**Color in life** (Figure 102): Dorsum greenish yellow with two to seven large yellow spots (diameter 0.5–2.1 mm), and with or without black flecks; upper lip unpigmented; ventral parietal peritoneum transparent; pericardium polymorphic, transparent (red heart visible ventrally) or white (heart not visible ventrally); transparent peritonea covering the kidneys, and urinary and gall bladders; white peritonea covering the liver, intestines, stomach, and testes; bones white; iris white to yellow, with minute dark lavender flecks around the pupil or forming a horizontal stripe.

**Color in ethanol:** Dorsum cream with large white spots and with or without dark flecks; iris white with dark lavender pigment; white peritonea covering the liver, intestines, stomach, and testes; pericardium polymorphic (white, partially white, or transparent).

**Biology and Ecology:** According to Barrera-Rodríguez and Ruiz-Carranza [195], most individuals of *Hyalinobatrachium aureoguttatum* were found on the underside of leaves (*Heliconia* spp.) 100–700 cm above small streams during the night. Amplectant pairs and egg clutches were found during July and August 1987. Amplexus is axillary, and eggs are deposited on the underside of leaves. In Durango (Provincia de Esmeraldas, Ecuador), individuals were reproductively active (amplectant pairs and egg clutches) 24–26 May 2006. At Estero Piedras, the species was reproducing in August 2007; a clutch with 36 eggs was found on the underside of a leaf on a bush along a stream [167]. Valencia-Aguilar et al. [196] studied populations from the Pacific lowlands of Colombia. They found that males exhibit high fidelity to their territory; each male repeatedly uses the same leaf (*Heliconia* sp., *Anthurium* sp., *Philodendron* sp., *Cyclanthus* sp., *Calathea* sp., *Musa* sp.) for perching, calling, mating, and clutch attendance. Territoriality seems to be low, since several males where found in close proximity and fights or aggressive behaviors were not observed in intrusion events by co-specific males.

Female place green eggs on the underside of leaves; clutches 25–49 eggs [167,195–197]. Males provide parental care during the day and night until hatching (mean = 17.1 days ± 1.8). Males are polygynous and simultaneously attend up to five clutches [196].

**Call:** Most males call from the undersides of leaves, but some have been observed calling from leaf tops. The specific spectral and temporal characteristics of the call are unknown.

**Tadpole:** A description of the tadpole can be found in Ibáñez et al. [197] and Terán-Valdez et al. [167] (Figures 80 and 104).

**Figure 104.** *Hyalinobatrachium aureoguttatum*, ontogenetic variation of tadpoles. (**A**) Gosner Stage 25, QCAZ 37752. (**B**) Gosner Stage 27, QCAZ 32072. Photos by L. A. Coloma. Figure modified from Terán-Valdez et al. [167].

**Distribution** (Figure 105): *Hyalinobatrachium aureoguttatum* is known from extreme southwestern Panama and the Pacific lowlands and western slopes of the Cordillera Occidental of Colombia and Ecuador at elevations below 1340 m [101,167,195,197–199]. In Ecuador, this species has been reported from the provinces of Esmeraldas and Imbabura at elevations below 600 m (Specimens Examined). In Ecuador, the species has a potential distribution of 4,481 km<sup>2</sup> within the Chocoan Tropical Forest and Western Foothill Forest ecoregions.

**Conservation status:** Listed globally as *Near Threatened* by the IUCN [200]. Given that Ecuadorian populations of the species are fragmented because of agriculture, pasture lands, and mining, we sugges<sup>t</sup> placing the species in the *Endangered* category based on IUCN criteria B1, B2a, B2b(iii). The species is partially protected within the Mache-Chindul reserve.

**Evolutionary relationships** (Figure 101): *Hyalinobatrachium aureoguttatum* and *H*. *valerioi* are sister species.

**Remarks:** Some populations from Colombia usually have small (but visible) brown flecks on the dorsal surfaces of the head and body; Ecuadorian populations lack these flecks (Figure 102). Also, all Colombian specimens examined by us have a white pericardium, a character that is polymorphic in the Ecuadorian populations (white, partially white, or transparent pericardium). It is possible that these differences correspond to independent evolutionary lineages; however, we prefer to maintain these populations as one species until more data (e.g., acoustic, molecular) are available.

**Figure 105.** Distribution of *Hyalinobatrachium aureoguttatum* in Ecuador (yellow dots).

**Specimens examined:** *Hyalinobatrachium aureoguttatum:* Ecuador: *Provincia de Esmeraldas:* stream affluent of the Río Durango (1.05 N, 78.6167 W; 100–150 m), QCAZ 27429, 6302, 6303, 6441–42, 28802; 2 km E of San Francisco (1.0872 N, 78.6905 W; 60–80 m), on the San Francisco-Durango road, QCAZ 32101–02, 32105, 32129, 32132–33; Río Quingue, nearby Caimito (0.72096 S, 80.09117 W, 47 m), QCAZ 37306. *Provincia de Imbabura:* 6 km SE of Lita (0.79 N, 78.43 W; 600 m), QCAZ 4323. Colombia: *Departamento del Chocó:* Municipio El Carmen de Atrato, km 23 on road El Carmen–Quibdó (5◦47- N; 76◦20- W, 1030 m), ICN 17507, 17509–10, 17512; km 44 on road El Carmen–Quibdó, 630 m, ICN 17252–54, 17515–16; km 53 on road El Carmen–Quibdó, 420 m, ICN 17248, 17257, 17260, 17262, 17266–67, 17520–21, 17525, 17527–28, 17531–34, 17536–37.

## *Hyalinobatrachium chirripoi* (Taylor, 1958 [201]; Figures 106–108).

*Cochranella chirripoi* Taylor, 1958 [201]. Holotype: KU 36865.

Type locality: "Cocales Creek, Suretka, (Cantón de Talamanca,) Limón Province", Costa Rica. Savage [183] commented on the type locality.

*Centrolenella chirripoi*—Savage, 1967 [202].

*Hyalinobatrachium chirripoi*—Ruiz-Carranza and Lynch 1991 [6].

*Hyalinobatrachium cardiacalyptum* McCranie and Wilson, 1997 [203]. Holotype: USNM 342161.

Type locality: "Caño El Cajón (14◦21- N, 85◦29- W), at its junction with the Río Patuca, Departamento de Olancho, Honduras, elevation 200–225 m". Placed in synonymy by Cisneros-Heredia and McDiarmid, 2007 [17].

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

**Etymology:** The specific name *chirripoi* is named for the Chirripó Indians, local inhabitants of the area where the species was first found [201].

**Identification:** *Hyalinobatrachium chirripoi* is easily distinguished from most glassfrogs by having, in life, a lime green dorsum with small yellow spots, a completely transparent ventral parietal

peritoneum (red heart visible ventrally; Figure 106), and by lacking humeral spines. Additionally, it differs from most *Hyalinobatrachium* by having more webbing between Fingers II and III.

**Figure 106.** *Hyalinobatrachium chirripoi* in life. Reserva Canandé, 372 m, Esmeraldas province, Ecuador. Photos by Jaime Culebras.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout truncated in dorsal view, truncated to slightly protruding in lateral profile; (3) tympanum almost completely concealed, small when visible (tympanum diameter 2.3%–2.8% of SVL); tympanic membrane not differentiated from surrounding skin; (4) dorsal surfaces shagreen, lacking spicules; (5) venter areolate; lacking pair of enlarged subcloacal warts; (6) ventral parietal peritoneum transparent (condition P0); pericardium transparent; white peritonea covering the intestines and stomach; transparent peritoneum covering urinary and gall bladders; kidneys covered by a mostly translucent peritoneum, except for some iridophores on their ventral portions (condition V6); (7) liver bulbous, covered by white peritoneum (condition H2); (8) humeral spines absent; (9) basal to moderate webbing between Fingers I and II, extensive webbing between outer finger: I (2<sup>+</sup>–21/3)—(2–2−) II 1—(2<sup>2</sup>/3–3−) III (2−–2)—(1<sup>1</sup>/3–11/2) IV; (10) foot webbing extensive: I (0+–1)—(1<sup>1</sup>/2–12/3) II 1—(1<sup>1</sup>/2–12/3) III (0<sup>+</sup>–1)—2− IV (2−–2)—(1–1<sup>+</sup>) V; (11) ulnar and tarsal folds present, with minute iridophores; (12) concealed prepollex; in males, nuptial pad Type V; (13) Finger I longer than Finger II (Finger II about 85%–90% length of Finger I); (14) disc of Finger III of moderate size, 38%–47% of eye diameter; (15) in life, dorsum line green with small yellow spots, venter transparent (red heart visible; Figure 106); bones white; (16) in ethanol, dorsum cream with minute lavender spots and iridophores; (17) in life, iris yellowish white, with minute dark lavender flecks; (18) fingers and toes lacking melanophores, except for few on Toes IV and V; (19) males call from the upper and undersides of leaves; call undescribed; (20) fighting behavior unknown; (21) eggs deposited on the underside of leaves; maternal care absent; prolonged parental care provided by males; (22) elongate tadpole, with emarginate oral disc; tooth row formula 2/3; (23) small body size; in Ecuadorian males, SVL 25.0–25.5 mm (*n* = 2); Savage [148] reported that adult males have a SVL = 24–26 mm and that females are probably larger.

**Color in life** (Figure 106): Dorsum lime green with several small yellow spots; ventral parietal peritoneum transparent; pericardium transparent (red heart visible ventrally); transparent peritonea covering urinary bladder; white peritonea covering liver, intestines, and stomach; transparent peritoneum covering most of kidneys; bones white; iris yellowish white, with minute dark lavender flecks.

**Color in ethanol:** Dorsum cream with minute lavender spots and iridophores; ventral parietal pericardium translucent; white peritonea covering liver, intestines, and stomach; pericardium transparent; iris silvery white with dark lavender flecks.

**Biology and Ecology:** According to Kubicki [24,204], in southeastern Costa Rica, *Hyalinobatrachium chirripoi* deposits 65–80 greenish–white eggs in a single layer on the underside of leaves overhanging streams. Males were most often seen calling from below palm fronds or other smooth leaves between 1–4 m above the water. Males were also seen calling from the upper sides of vegetation, but much less frequently. One male was seen guarding eggs during the night, and another male was observed guarding eggs during daylight hours [24]. At Reserva Itapoa, Ecuador, in June 2014, a male was observed calling and guarding an egg clutch on the underside of a leaf. Maternal care is absent; prolonged parental care is provided by males [25].

**Call** (Figure 107): Most males call from the undersides of leaves, but some have been observed calling from the upper sides [24,204]. The call is a high-pitched insect-like buzz, very similar to that of *H*. *colymbiphyllum* [204]. We analyzed two notes from one individual (LBE-019). The typical advertisement call is composed by a single note. Note duration is 230–270 (mean = 250, SD = 28.3) ms. Notes are strongly pulsed and have 14–16 (mean = 15, SD = 1.4) amplitude peaks throughout the note, with pulses becoming difficult to distinguish near the end of the note. Pulses within a note have a rate of 59–61 (mean = 60, SD = 1) pulses per second. Notes have their peak amplitude in the first 50% of the note (relative peak time: Range = 0.0623–0.0638, mean = 0.063, SD = 0.001), where the peak amplitude occurs in the first several pulses. The dominant frequency of a note measured at peak amplitude is 4565 (mean = 4565, SD = 0) Hz and is contained within the fundamental frequency. The fundamental frequency has a lower limit of 1895–3273 (mean = 2584, SD = 974) Hz and a higher limit of 4651–5082 (mean = 4866, SD = 305) Hz.

**Figure 107.** Call of *Hyalinobatrachium chirripoi* recorded at Reserva Itapoa, 321 m, Esmeraldas province, Ecuador, LBE-C-019.

**Egg masses and tadpoles:** Greenish–white eggs (53–80) are deposited in a single layer on the underside of leaves; as development continues the embryos become pale tannish red [24,147,204]. Below, we present a summary of the tadpole description by Hoffmann [147]. Hatchlings (Gosner stages 25 and 26) have an unusual pattern on their dorsum: Two rows of brown-pigmented, spider-like dots of melanophores stretch longitudinally over the entire body and join in a semicircle at the tip of snout; another line of pigmented spots extends dorsally over the tail musculature. Because of the general lack of pigment and high transparency, the body coloration is partially determined by the red color of the gills and heart. As development continues (Gosner stages 25–41), the tadpoles become very elongated and are among the most elongated centrolenid larvae known (ratio of body length to body

width of 2.41 ± 0.22); they have a completely transparent skin and the dorsolateral lines are mostly lost; the body has a pale rose shine and the tail is cream or, sometimes, yellowish. The oral disc is emarginate and, except for a moderate dorsal gap, is bordered by a single row of about 45 marginal papillae; the ventral ones being flatter than the lateral ones. The LTRF is 2(2)/3; the tooth rows are about of equal width (P-3 is slightly narrower) and extend across most of the oral disc. The A-2 gap is about as wide as the upper jaw, which is broadly arched; the lower jaw is V-shaped; the edges of both jaw sheaths are edged by many narrow serrations [147].

**Distribution** (Figure 108): *Hyalinobatrachium chirripoi* is known from lowland forest localities in Central America in southeastern Costa Rica and eastern Panama, and south into South America in western Colombia and Ecuador at elevations below 600 m ([24,148,204], this work). In Ecuador, this species has been observed in few localities within the Esmeraldas province (Río Quingue, Río Bogotá, Reserva Itapoa, Tesoro Escondido, Reserva Canandé), in the Chocó ecoregion, at elevations below 320 m.

**Conservation status:** Listed globally as *Least Concern* by the IUCN [205]. In Ecuador, the species has been recently found in five localities in the Chocó Ecoregion, an area under constant deforestation pressure. Although the *Least Concern* category seems accurate at the global level, in Ecuador, the species is threatened by habitat destruction because of agriculture, pasture lands, and mining. Thus, we sugges<sup>t</sup> placing the species in the *Endangered* category at the local level, based on IUCN criteria B1, B2a, B2b(iii).

**Figure 108.** Distribution of *Hyalinobatrachium chirripoi* in Ecuador (yellow dots).

**Evolutionary relationships** (Figure 101): Molecular data support a sister relationship between *Hyalinobatrachium chirripoi* and *H*. *colymbiphyllum*.

**Specimens examined:** *Hyalinobatrachium chirripoi:* Ecuador: Provincia de Esmeraldas: Río Quingue (0.722◦ N, 80.08◦ W; 47 m), QCAZ 37309, 48271; Río Bogotá, DFCH-USFQ C1903; Reserva Itapoa (0.513◦ N, 79.134◦ W; 320 m), MZUTI 3609–10; Tesoro Escondido (0.542◦ N, 79.145◦ W; 225 m), MZUTI 3625; Reserva Canandé (0.526◦ N, 79.209◦ W; 310 m), MZUTI 4745. Costa Rica: Limón Province: Suretka, along Cocales Creek, KU 36862–64, 36866–70.

*Hyalinobatrachium fleischmanni* (Boettger, 1893 [206]; Figures 109–111).

*Hylella fleischmanni* Boettger, 1893 [206]. Lectotype: SMF 3760, designated by Mertens (1967) [207].

Type locality: "San José, Costa Rica".

*Hylella chrysops* Cope, 1894 [208]. Neotype: SMF 3760 (= holotype of *Hylella fleischmanni*), designated by Starrett and Savage (1973) [209]. Placed in synonymy by Boulenger (1895) [210].

*Centrolenella fleischmanni*—Noble (1924) [186].

*Centrolenella viridissima* Taylor, 1942 [211]. Holotype: EHT-HMS 27725 (now FMNH 100093). Type locality: "Agua de Obispo, Guerrero", Mexico. Placed in synonymy by Starrett and Savage (1973) [209].

*Cochranella fleischmanni*—Taylor (1951) [15].

*Cochranella decorata* Taylor, 1958. Holotype: KU 36896. Type locality: "Hda. La Florencia, about 3 miles west of Turrialba, Cartago Province, Costa Rica". Placed in synonymy by Starrett and Savage (1973) [209].

*Cochranella millepunctata* Taylor, 1958 [201]. Holotype: KU 36887. Type locality: "La Palma, San José Province, Costa Rica". Placed in synonymy by Starrett and Savage (1973) [209]. *Centrolenellafleischmanni*—Goin(1964)[187].

 *Hyalinobatrachium fleischmanni*—Ruiz-Carranza and Lynch (1991) [6].

**Common names:** English: Fleischmann's Glassfrog (Liner, 1994). Spanish: Rana de Cristal de Fleischmann.

**Etymology:** The specific name *fleischmanni* is a patronym for Carl Fleischmann, a German collector–naturalist who donated the specimens used by Boettger to describe the species [84].

**Identification:** *Hyalinobatrachium fleischmanni* is easily recognizable by having, in life, a lime green dorsum with small yellowish spots, a completely transparent ventral parietal peritoneum, white heart and digestive tract (pericardium and gastrointestinal peritoneum covered by iridophores), lacking humeral spines, and having reduced webbing between inner fingers (Figure 110). In the Ecuadorian Pacific lowlands, similar species include *H*. *aureoguttatum*, which has large yellow spots on the dorsum; *H*. *valerioi*, distinguished by having a green dorsal reticulum; and *H*. *chirripoi*, which, in life, has a transparent pericardium (red heart visible through ventral skin). We note that *H*. *fleischmanni* represents a species complex that requires taxonomic revision [212].

**Figure 109.** *Hyalinobatrachium fleischmanni* in life. Ecuador, near Durango, 77 m, QCAZ 32107. Photos by Martín Bustamante.

**Diagnosis:** (1) Vomerine teeth absent; (2) snout subacuminate in dorsal aspect and rounded in profile; (3) tympanum not visible; supratympanic fold absent; (4) dorsal skin shagreen; (5) venter smooth; lacking pair of enlarged subcloacal warts; (6) ventral parietal peritoneum transparent (condition P0); white peritoneum covering heart, intestines and stomach; transparent peritoneum on urinary bladder (condition V5); (7) liver bulbous, hepatic peritoneum white (condition H2); (8) humeral spines absent; (9) webbing reduced between Fingers I, II and III, moderate between outer fingers (Figure 110); webbing formula I (2–2+)—2 II (1–1+)—(3) III (2)—(1+–2) IV (1–2) IV; (10) webbing between toes moderate; webbing formula on feet I (1–1<sup>1</sup>/2)—(2) II (1)—(2–2+) III (1)—(2<sup>+</sup>–2−) IV (2–2+)—(1–1+) V; (11) ulnar and tarsal fold absent; (12) concealed prepollex; nuptial pad Type IV; (13) Finger II shorter than Finger I; (14) eye diameter larger than width of disc on Finger III (disc of Finger III width 33%–37% of eye diameter); (15) in life, lime green dorsum with yellow spots (Figure 109); venter transparent, pericardium usually white (but see Color in life); bones white; (16) in preservative, dorsum cream; (17) in life, iris yellowish to greyish white with some dark punctuations; (18) fingers and toes lacking melanophores; (19) males call from the underside of leaves; the call consists of a single "wheet" note, with duration of 150–300 ms, and a dominant frequency of 3800–4500 Hz at the beginning and 4800–5300 HZ at the end; (20) fighting behavior varies from vent to vent to amplexus-like; (21) egg clutches usually laid on the underside of leaves; maternal care absent; males provide prolonged parental care; (22) oral apparatus complete; oral disc with single row of marginal papillae laterally and ventrally, wide dorsal gap; jaw sheaths normal; 2(2)/3 labial tooth rows, A-2 with wide gap, tooth rows situated nearly lateral to mouth and upper jaw sheath; (23) minute body size; snout–vent length in adult males 19.3–26.8 mm (*n* = 13), and in adult females 22.4–31.1 mm (*n* = 7).

**Color in life** (Figure 109): Dorsum lime green with pale yellow or greenish spots. Venter transparent, pericardium usually white, visceral and hepatic peritonea white. Twomey et al. [19] report that within a single population of *H*. *fleischmanni* near San Gabriel Mixtepex (Oaxaca, Mexico), adults exhibited variation in the condition of the pericardium (from white to transparent).

**Figure 110.** *Hyalinobatrachium fleischmanni*, variation in hand webbing, ventral view. (**A**) KU 116447. (**B**) QCAZ 22301. Illustrations by Juan M. Guayasamin.

**Color in ethanol:** Cream dorsum with dark melanophores in the places where green coloration was in life. Venter translucent. Parietal peritoneum completely transparent, heart and all viscera covered by white lining.

**Biology and ecology:** The natural history of *Hyalinobatrachium fleischmanni* has been reviewed by several studies, including Greer and Wells [213], Villa [214], Jacobson [188], Hayes [189], Savage [148], Kubicki [24], and Delia et al. [215]. We summarize the essential information here but refer readers to those papers for more data. Individuals of *Hyalinobatrachium fleischmanni* have been found on vegetation up to 10 m above streams at night. Males are territorial and call from the lower (mainly) and upper surfaces of leaves. Females usually deposit eggs clutches on the lower surfaces of leaves, although some variation has been reported [215]; maternal care is absent [25]. Physical combat occurs when a male intrudes into an occupied territory. Initially the owner of the territory will call vigorously, but if that strategy fails, male–male combat starts usually with both males adopting an amplexus-like position. Both males may give quick calls. Although venter to venter combat was thought to be absent in *Hyalinobatrachium*, *H*. *fleischmanni* has been observed to adopt this combat position, with both males dangling upside down while holding vegetation with their hind limbs [215]. Males attend egg clutches at night, and may continue to advertise, sometimes ending with multiple clutches from different females. During the day, the male parent retreats to nearby vegetation. Only males are involved in parental care of the eggs. Delia et al. [25] observed that males of this species are attentive to individual embryo needs. Eggs are pale greenish white, and clutches are deposited on the underside of leaves. Each egg clutch has one layer of 10–50 eggs [189,216].

**Call:** The call consists of a single "wheet" note that rises at the end, with duration of 150–300 ms. The dominant frequency is 3800–4500 Hz at the beginning and 4800–5300 HZ at the end. Calls are emitted 4–19 per minute [148,188,209,213].

**Tadpole:** A description of the tadpole can be found in Starrett [146], Savage [148], and Hoffmann [147].

**Distribution** (Figure 111): *Hyalinobatrachium fleischmanni* is known from southern Mexico, across Mesoamerica, south to central Ecuador at elevations below 800 m [24,148,212,217,218]. In Ecuador, the species is known from localities in the Pacific lowlands (mostly below 300 m) in the provinces of Esmeraldas, Pichincha, Santo Domingo de los Tsáchilas, Los Ríos, and Guayas. In Ecuador, the potential distribution of the species is 39,738 km2. The distribution of *H*. *fleischmanni* is an overestimation because, as currently recognized, it is a species complex [212].

**Figure 111.** Distribution of *Hyalinobatrachium fleischmanni* in Ecuador (yellow dots).

**Conservation status:** Globally, *Hyalinobatrachium fleischmanni* is considered as *Least Concern* by the IUCN because of its wide distribution and tolerance of habitat modifications [219]. However, recent studies sugges<sup>t</sup> that this taxon represents a species complex that requires subdivision [212]. In Ecuador, about 55% of the potential distribution of the species is affected by human activities. Even though habitat destruction is considerable, *H*. *fleischmanni* is known to tolerate a degree of disturbance [24]. In Ecuador, we sugges<sup>t</sup> maintaining the category of *Least Concern* until new taxonomic studies are available.

**Evolutionary relationships** (Figure 101): Our tree shows *Hyalinobatrachium fleischmanni* as the sister species of *H*. *muiraquitan*, which is endemic to the Brazilian Amazon basin. Although the Venezuelan *H*. *tatayoi* falls within the genetic variation of *H*. *fleischmanni*, recent studies sugges<sup>t</sup> that *H*. *fleischmanni* represents a species complex that requires taxonomic revision [212].

**Specimens examined:** *Hyalinobatrachium fleischmanni:* Costa Rica: Alajuela, USNM 219249–61. San José, USNM 219263–80. Guanacaste, USNM 219282–303. Ecuador: *Provincia de Los Ríos:* Quevedo, USNM 60520; Río Palenque, USNM 286639–40; Patricia Pilar, USNM 286645; Hacienda Cerro Chico, USNM 286646. *Provincia de Esmeraldas*: 4 km W Durango, QCAZ 23549 km<sup>2</sup> in the San Francisco–Durango road, QCAZ 32073; Río Quingue, nearby Caimito (0.72096 S, 80.09117 W, 47 m), QCAZ 37308; Río Onzole, QCAZ 10433; Tesoro Escondido (0.542 N, 79.145 S, 225 m), MZUTI 3621–3625. *Provincia de Santo Domingo de los Tsáchilas*: Bosque Protector La Perla, QCAZ 12606. Honduras: Olancho: USNM 342162–342213. Nicaragua: Matagalpa: USNM 220013–18. Nueva Segovia: USNM 220019–36. México: Chiapas: USNM 115499.

**Localities from the literatura:** Ecuador: *Provincia de Guayas:* Cerro de Hayas (2.7299◦ S, 79.6297◦ W, 127 m), MZUA.AN.1693–1694; *Provincia de Los Ríos:* Macul (1.2279◦ S, 79.7531◦ W, 84 m), MZUA.AN.660–661 [218].

*Hyalinobatrachium iaspidiense* (Ayarzagüena, 1992 [220]; Figures 112 and 113).

*Centrolenella iaspidiensis* Ayarzagüena, 1992 [220]. Holotype: EBD 28803. Type locality: "Quebrada Jaspe. San Ignacio de Yuruaní. Edo. Bolivar. Venezuela." *Centrolene iaspidiensis*—Duellman, 1993 [221]. Unintended combination. *Hyalinobatrachium iaspidiense*—Myers and Donnelly, 1997 [104].

*Hyalinobatrachium nouraguensis*—Lescure and Marty, 2001 [222]. Holotype: MNHNP 1999.8604. Type locality: "Saut Arataye (environs du camp de base), Réserve des Nouragues (bassin de l'Approuague), Guyane française". Placed in synonymy by Yánez-Muñoz, Pérez-Peña, and Cisneros-Heredia, 2009 [223].

#### **Common names:** English: Jaspe Glassfrog. Spanish: Rana de Cristal de Jaspe.

**Etymology:** The specific name *iaspidiense* is derived from the Greek word *iaspis*, meaning jasper, in reference to the type locality of the species, Quebrada de Jaspe.

**Identification:** *Hyalinobatrachium iaspidiense* is one of the most easily recognizable centrolenid. In life, it has a dorsum with large, lime green, irregular blotches and small, black spots (Figure 112). The only species with a similar dorsal coloration is *H*. *mesai*, a species known only from the southern slope of Sarisariñama-tepui, Venezuela [224]. It has been suggested, however, that *H*. *mesai* and *H*. *iaspidiense* represent the same species [223].

**Figure 112.** *Hyalinobatrachium iaspidiense* in life. (**Left**): Adult female (QCAZ 38438) from Ecuador, Yachana Reserve, 300–350 m; photo by Susan North. (**Right**): Adult male (SMNS 12247) from Guyana, Mabura Hill Forest Reserve, 60 m; photo by Raffael Ernst.

**Diagnosis:** (1) Vomerine teeth absent; (2) snout truncated in dorsal aspect and slightly protruding in profile; (3) tympanum not visible; supratympanic fold absent; (4) dorsal skin shagreen; (5) venter smooth; pair of enlarged subcloacal warts, but low and difficult to see; (6) ventral parietal peritoneum transparent (condition P0); pericardium transparent; white peritoneum covering intestines and stomach; transparent peritoneum on urinary bladder (condition V6); (7) liver bulbous, hepatic peritoneum white (condition H2); (8) humeral spines absent; (9) webbing absent between Fingers I, II, and III, moderate between outer fingers; webbing formula III (2–2<sup>+</sup>)—(1<sup>+</sup>–2−) IV; (10) webbing between toes moderate; webbing formula on foot I (1–1<sup>+</sup>)—(2<sup>+</sup>–21/3) II (1<sup>+</sup>–11/3)—(2<sup>+</sup>–21/4) III (1–1+)—(2–2<sup>3</sup>/4) IV (2<sup>+</sup>–21/4)—(1<sup>+</sup>–11/4) V; (11) ulnar fold present, enameled; external tarsal fold present, enameled; internal tarsal fold short and low; (12) concealed prepollex; nuptial pad Type IV; (13) Finger II shorter than Finger I (Finger II about 93% of Finger I); (14) disc of Finger III width 39%–57% of eye diameter; (15) in life, dorsum translucent with a yellowish–green background coloration, large lime green blotches, and small black spots (Figure 112); bones white; (16) in preservative, dorsum cream with large, irregular white blotches and small, black spots; (17) in life, iris yellowish to greyish white; (18) fingers and toes

lacking melanophores, except few present on Finger IV and Toes IV and V; (19) males call from the underside of leaves; the call consists of a single pulsed note, with a dominant frequency at 4440–4710 Hz; (20) fighting behavior unknown; (21) egg deposition site unknown; parental care unknown; (22) tadpole unknown; (23) minute body size; in Venezuelan specimens: SVL in females 22.7 mm (*n* = 1), in males 19.8–21.8 mm (*n* = 8) [106]; in Ecuador, male SVL = 19.9 mm (*n* = 1), and female SVL = 21.9 mm (*n* = 1).

**Color in life** (Figure 112): Dorsum translucent, with a yellowish–green background coloration, large lime-green blotches, and small black spots. Venter transparent, pericardium transparent (red heart visible), visceral and hepatic peritonea white [223,225].

**Color in ethanol:** Dorsum cream lavender, with large irregular white marks and small black spots. Ulnar and tarsal folds enameled; some of the warts below cloaca enameled. Venter transparent, pericardium transparent, visceral and hepatic peritonea white; peritonea around urinary bladder transparent; gall bladder white (description based on two adults, MECN 4033, QCAZ 38438).

**Variation:** Ulnar and tarsal folds less evident and with few iridophores (QCAZ 38438).

**Biology and ecology:** Found on vegetation 4–7 m above streams [106]. In Ecuador (Yachana Reserve), an adult female was found on a long blade of tall grassy shrub overhanging water, located in the middle of a stream (4 m wide). Stream habitat is variable, but the female was found in a shallow area (~20 cm deep) where the stream becomes a riffle and is more fast-flowing. Stream bottom here is composed of about 80% pebbles and small rocks and 20% sand near edges. The forest adjacent to the stream is near pristine and about 25–30 m high [225]. Parental care unknown.

**Call:** The call consists of a single note, with 16 pulses and a duration of 62.7–75.1 ms. Dominant frequency at 4440–4710 Hz. The total frequency range for the call is between 3710 and 5850 Hz, with a secondary frequency at 8600–9200 Hz. The call was emitted frequently (7–10 calls per minute) [106].

**Tadpole:** Not described.

**Distribution** (Figure 113): *Hyalinobatrachium iaspidiense* is known from the Guiana region of Venezuela, central Guyana, Surinam (Sipaliwini District), and French Guiana, as well as from the Amazon Basin of Ecuador, Peru, and Brazil at elevations below 1000 m [106,222,225–229]. In Ecuador, the species is known from few localities in the Amazonian raiforest.

**Figure 113.** Distribution of *Hyalinobatrachium iaspidiense* in Ecuador (yellow dots).

**Conservation status:** *Hyalinobatrachium iaspidiense*is considered as *Data Deficient* by the IUCN [230]. However, given the large distribution of the species and the absence of immediate threats, we sugges<sup>t</sup> that it should be considered as *Least Concern,* both globally and in Ecuador.

**Evolutionary relationships** (Figure 101): *H. iaspidiense* is the sister species of *H. tricolor.*

**Specimens examined:** *H. iaspidiense:* Ecuador: *Provincia Napo:* Yachana Reserve (0◦52-21.71" S, 77◦14-13.43" W; 300–350 m), QCAZ 38438; Estación Biológica Jatun Sacha (1.066 S, 77.617 W, 405 m), QCAZ 53023. *Provincia Orellana:* Km 66 on the Pompella Sur–Iro road (0.8022 S, 76.398 W, 280 m), QCAZ 54947. *Provincia Sucumbíos:* Totoa Nai'qui (0.03442◦ S, 76.75278◦ W, ca. 280 m), MECN 4033.

*Hyalinobatrachium munozorum* (Lynch and Duellman, 1973 [22]; Figures 114–116).

*Centrolenella munozorum* Lynch and Duellman, 1973 [22]. Holotype: KU 118054.

Type locality: "Santa Cecilia, 340 m, Provincia Napo (Sucumbíos), Ecuador".

*Hyalinobatrachium munozorum*—Ruiz-Carranza and Lynch, 1991 [6].

*Hyalinobatrachium ruedai* Ruiz-Carranza and Lynch, 1998 [27]. Holotype: ICN 40409. Type

locality: "(Colombia,) Departamento de Caquetá, municipio de Miraflores, Parque Nacional Natural de Chiribiquete, campamento base, 530 m". **New synonymy.**

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

**Etymology:** The specific name *munozorum* is a patronym for Ildefonso Muñoz B. and Blanca Muñoz, hosts of the extensive herpetological work carried out in Santa Cecilia by W. E. Duellman and his students [22].

**Identification:** *Hyalinobatrachium munozorum* can be distinguished from most glassfrogs by having a green dorsum with pale yellow spots, a completely transparent ventral parietal peritoneum, white liver and digestive tract, and by lacking humeral spines (Figure 114). Similar species with a completely transparent venter that inhabit the Amazon basin include *H*. *bergeri*, *H*. *carlesvilai*, *H*. *iaspidiense*, *H*. *mondol*ffi, *Chimerella mariaelenae,* and *Teratohyla amelie*. *Hyalinobatrachium munozorum* differs from *C*. *mariaelenae* by having, in life, a green dorsum with pale yellow spots (dorsum green with dark grey spots in *C*. *mariaelenae*) and by lacking humeral spines (small humeral spines present in adult males of *C*. *mariaelenae*). *Teratohyla amelie* has a uniform green dorsum, lacking the pale-yellow spots visible in *H*. *munozorum*. *Hyalinobatrachium iaspidiense*, in life, has a dorsum with large, green, irregular marks and small, black spots. Morphological differentiation among *H*. *munozorum*, *H*. *mondolfii*, and *H*. *carlesvilai* is minimal, although molecular divergence is considerable [22,231,232]; the pupil of *H*. *bergeri* and *H*. *carlesvilai* is surrounded by a dark grey ring (absent in *H*. *munozorum*).

**Figure 114.** *Hyalinobatrachium munozorum* in life. Adult male from stream near Tena, 708 m, MZUTI 1616. Photos by Eduardo Toral.

**Diagnosis:** (1) Vomers lacking teeth; (2) snout varies from truncated to rounded in dorsal aspect and round to truncated in lateral view; (3) tympanum usually obscured by skin, small when evident, its diameter about 25% of eye diameter; supratympanic fold low; (4) dorsal skin shagreen; males lacking spicules; (5) ventral skin texture areolate; lacking pair of enlarged subcloacal warts; (6) ventral parietal peritoneum transparent, white pericardium, white peritonea covering intestines and stomach, transparent urinary bladder; (7) liver bulbous, covered by white peritoneum; (8) humeral spines absent; (9) webbing between inner fingers absent; webbing formula for outer fingers: III (1<sup>1</sup>/2–2−)—(1–2−) IV; (10) feet about three-fourths webbed; webbing formula: I (0<sup>+</sup>–1−)—(1<sup>1</sup>/3–2−) II (0+–1)—(1<sup>1</sup>/2–2) III (0<sup>+</sup>–11/3)—(2−–21/3) IV (2−–21/3)—(0<sup>+</sup>–1) V; (11) ulnar and tarsal folds variable, from inconspicuous to pronounced; (12) concealed prepollex; in males, nuptial pad not evident; (13) Finger I slightly longer than Finger II (Finger II length 93.7%–97.6% of Finger I); (14) disc of Finger III of moderate size, its width 37.9%–56.2% of eye diameter; (15) in life, dorsum green with diffuse yellow spots, venter transparent; bones white; (16) in preservative, dorsum cream lavender with small unpigmented spots; (17) in life, iris pale to bright gold with dark punctuations; (18) dorsal surfaces of fingers and toes lacking melanophores; (19) call composed by a single note with a duration of 145–178 (mean = 170, SD = 9.9) ms; each call is mostly tonal, with a dominant frequency at 5719–5906 (mean = 5812, SD = 64) Hz; (20) fighting behavior unknown; (21) egg clutches laid on the underside of leaves; parental care unknown; (22) tadpoles unknown; (23) minute body size; in males, SVL 19.9–21.9 mm ( X = 20.6 ± 0.691; *n* = 8); in females, SVL 20.9–23.6 mm (*n* = 3).

**Color in life** (Figure 114): Dorsal surfaces of head and body green with small, pale yellow spots and minute dark brown flecks. Parietal peritoneum transparent. White peritonea covering heart, liver, digestive tract, and testes; transparent urinary bladder. Iris pale gold with minute dark flecks. See Taxonomic Remarks.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs cream with numerous minute lavender flecks. White visceral and hepatic peritonea. Layer of iridophores covering heart.

**Variation:** Specimens in the type series have a round snout in lateral view, and low ulnar and tarsal folds that are inconspicuous and unpigmented. Additional specimens from the type locality (Santa Cecilia) show the following variation: Snout truncated in lateral view, ulnar and tarsal folds conspicuous and white (KU 155493–96); conspicuous tarsal fold and low ulnar fold (KU 152488, 152489, 175215). Specimens identified as *Hyalinobatrachium ruedai* (a synonym of *H*. *munozorum;* see below) by Ruiz-Carranza and Lynch [27] and Cisneros-Heredia and McDiarmid [233] show the following variation: Snout truncated to rounded in dorsal and lateral views, ulnar and tarsal folds vary from inconspicuous to low and white. A female (KU 154749) has a truncated snout and conspicuous white ulnar and tarsal folds. The pericardium of all specimens from Ecuador and Colombia (including the type series) is white. In some individuals (KU 123225, 152488–89, 155494, 155496, 175504), the layer of iridophores is thinner. Cisneros-Heredia and McDiarmid [17] reported that one specimen showed a bicolored iris in life, with a light grey circumpupilary zone. These authors also reported variation in the iris coloration in preservative, from lavender–cream background with dense dark punctuations and lavender mid-line to uniform white.

**Biology and ecology:** The type series was found on leaves of bushes and trees at night: One over a pond, one away from water in primary forest, one on a palm frond 2 m above a stream, and one on an herbaceous leaf more than 2 m above a stream. The specimen from Lago Agrio was obtained from the foliage of a large tree that was felled during the clearing of primary forest [22]. A specimen from Tena was collected at night on the leaf of a bush at a rivulet in secondary forest [17]. Males called from riverine vegetation at a locality nearby Tena on August 2012. Parental care is unknown.

**Call** (Figure 115): We analyzed 18 notes from 1 individual (LBE-C-020) recorded at Río Bigal, Orellana province, Ecuador. The call is composed by a single note. Note duration is 145–178 (mean = 170, SD = 9.9) ms. Notes are mostly tonal, but they exhibit one to seven (mean = 4.5, SD = 1.6) low-amplitude peaks. Notes usually have their peak amplitude in the first 50% of the note (relative peak time: Range = 0.1699–0.661, mean = 0.448, SD = 0.123). The dominant frequency of a note measured at peak amplitude is 5719–5906 (mean = 5812, SD = 64) Hz and is contained within the fundamental frequency. The fundamental frequency has a lower limit of 5625–5812 (mean = 5708, SD = 63) Hz and a higher limit of 5812–6000 (mean = 5906, SD = 64) Hz.

**Figure 115.** Vocalization of *Hyalinobatrachium munozorum* recorded at Río Bigal, 930 m, Orellana province, Ecuador, by Morley Read (LBE-C-020).
