**Tadpole:** Not described.

**Distribution** (Figure 214): *T. midas* has been reported from the Amazonian lowlands (<940 m) of Ecuador, Peru, and Brazil, and is likely to be present in Colombia and Bolivia ([17,19,22,139,229,296], this work). In Ecuador, *T*. *midas* localities are at elevation of 190–930 m, with a potential distribution of 51,794 km<sup>2</sup> within the Amazonian Tropical Rainforest and the Eastern Foothill Forest regions.

**Figure 214.** Distribution of *Teratohyla midas* in Ecuador (yellow dots).

**Conservation status:** Globally, *Teratohyla midas* is listed as *Least Concern* by the IUCN [297]. In Ecuador, the species has a broad distribution; thus, we agree with the current conservation status.

**Evolutionary relationships** (Figure 210): *Teratohyla midas*, as currently recognized, is the sister species of *T*. *adenocheira*. However, there is considerable genetic structure within *Teratohyla midas* and it might represent a species a complex.

**Specimens examined:** *Teratohyla midas:* Ecuador: *Provincia de Morona Santiago:* Río Kusutka (02.111◦ S, 77.73897◦ W; 650 m), nearby Wisui Biological Station, MZUTI 031. *Provincia de Napo:* Puerto Misahualli (1.033 S, 77.667 W; 400 m), QCAZ 20001–02; S side of Río Napo, 6.5 km ESE Puerto Misahualli at La Cruz Blanca in Jatun Sacha Biological Reserve (1.05 S, 77.6 W; 395 m), QCAZ 9038–44; Tena, near airport (0.985 S, 77.825 W; 550 m), QCAZ 8819–21. *Provincia de Orellana:* stream nearEstación Científica Yasuní (0.667 S, 76.4 W; 230–240 m), QCAZ 22876, 23895; Estación Científica Yasuní, near the bird observation tower (0.67491◦ S, 76.39834◦ W), QCAZ 19316; Río Napo, Añangu (0.52492◦ S, 76.38445◦ W; 255 m), QCAZ 43939–44; Tiputini Biodiversity Station (0.6167◦ S, 76.167◦ W; 190–270 m), DFCH-D102; km 37–38 on the Pompeya–Iro road (ca. 0.6 S, 76.45 W), QCAZ 17311–12, 17314–16. *Provincia de Pastaza:* Pomona (1.6833 S, 77.883 W; 930 m), QCAZ 33225–26; Estación Hola Vida near Pomona (1.6284◦ S, 77.9095◦ W; 837 m), QCAZ 33225–26; Parroquia Teniente Hugo Ortíz (1.37 S, 77.955 W), QCAZ 33252–53; Río Oglán, Curaray (01◦19- S, 77◦35- W; 600 m), USNM 288437; tributary of Río Lliquino (1.18183◦S, 77.47879◦ W; 400 m), QCAZ 37933–34; tributary of Río Lliquino, Villano B Camp (1.45295◦ S, 77.443◦ W; 350 m), QCAZ 37916–19; Río Pucayacu (1.374148◦S, 77.90955◦ W; 940 m), QCAZ 33252–53. *Provincia de Sucumbíos:* Río Pañayacu (0.40667 S, 76.113 W; 210 m), QCAZ 20329; Monte Tour (0.0315 S, 76.321 W; 290 m); near Lago Agrio (0.0847 S, 76.8828 W; 370 m), KU 125334; Pozo Peña Blanca, E of Dureno and near Pacayacu (0.0001 S, 76.648 W; 260 m), QCAZ 6385; Puerto Bolívar (0.0886 S, 76.142 W; 240 m), QCAZ 28134, 28137; Puerto Libre (0.061 N, 76.75 W; 280 m), KU 123220–23; Estación Científica PUCE at the Reserva de Producción Faunística Cuyabeno (0.083 S, 76.166 W; 220 m), QCAZ 6015; Río Conejo, 2 km N of Santa Cecilia (0.097778 N, 76.99 W), KU 153256; Santa Cecilia (0.05 N, 76.9667 W; 340 m), KU 105283, 107026, 123219, 146625, 150622–23, 152487, 158518.

**Photographic records:** *Teratohyla midas:* Ecuador: *Morona Santiago province:* Kimm, 5 km NW airline distance from Santiago (3.0149 S, 78.0356 W), QCAZ 73554 [294]. Road Peñas-Shaimi, 2.8 Km E of Río Yaupi by road (2.9663 S, 77.84682 W), QCAZ 73546–47 [294]. *Orellana province:* Reserva Río Bigal (0.532913◦ S, 77.423228◦ W; 981 m), photo by Morley Read. *Pastaza province:* Lorocachi (1.625259 S, 75.99035 W), QCAZ 56024 [294].

## *Teratohyla pulverata* (Peters, 1873 [298]; Figures 215–217).

*Hyla pulverata* Peters, 1873 [298]. Holotype: ZMB 7842, according to Duellman, 1977 [118]. Type locality: "Chiriqui", Panama; at the time of the description "Chiriqui" included both

Atlantic and Pacific versants of extreme western Panama, according to Myers, 1982 [299]. *Centrolenepulveratum*—Dunn,1931[236].

*Centrolenellapulveratum*—Taylor,1949[145].

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

*Cochranella petersi—*Goin, 1961 [97]. Type locality: "Rio Durango, N. W. Ecuador". Placed in synonymy by Guayasamin, Cisneros-Heredia, and Castroviejo-Fisher, 2008 [300].

*Centrolenella petersi*—Goin, 1964 [187].

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

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

*Hyalinobatrachium petersi—*Ruiz-Carranza and Lynch, 1998 [27].

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

**Common names:** English: Dusty Glassfrog [24]. Spanish: Rana de Cristal polvosa [24]. **Etymology:** The specific name comes from the Latin *pulvereus*, meaning dusty, and refers to the minutewhitespotsonthedorsumofthespecies.

**Identification:** Among glassfrogs that inhabit the Chocoan forest of Ecuador, *Teratohyla pulverata* is unique by having a green dorsum with small white spots, completely transparent ventral parietal peritoneum (Figure 215), white hepatic and gastrointestinal peritonea, a sloping snout in lateral profile, and by lacking humeral spines. Only species in the genus *Hyalinobatrachium* could be confused with *T*. *pulverata;* however, *T*. *pulverata* has green bones (white in *Hyalinobatrachium*), deposits eggs on the upper sides of leaves (undersides of leaves in *Hyalinobatrachium*), and has conspicuous teeth on the vomers (teeth absent in *Hyalinobatrachium*).

**Figure 215.** *Teratohyla pulverata* in life. Individual (QCAZ 32224) from Silanche, Pichincha province. Photos by Luis A. Coloma.

**Diagnosis:** *Teratohyla pulverata* exhibits the following traits: (1) Dentigerous process of the vomer with two to four teeth; (2) snout rounded in dorsal view, sloping in lateral profile (Figure 216); (3) tympanum visible, relatively small, its diameter 20.2%–23.3% of eye diameter; tympanic annulus visible except for upper border covered by supratympanic fold; tympanic membrane differentiated and translucent, pigmented as surrounding skin; (4) dorsal surfaces shagreen; males with small spicules (only visible under magnification); (5) ventral surfaces granular, lacking pair of enlarged subcloacal warts; (6) ventral parietal peritoneum completely transparent (condition P0); pericardium and gastrointestinal peritoneum white (condition V2); (7) white, bulbous liver (i.e., covered by iridophores; condition H2); (8) humeral spines absent; (9) webbing between Fingers I and II absent or basal; webbing formula: II (1<sup>+</sup>–11/3)—(24/5–3−) III (1<sup>1</sup>/3–12/3)—(1<sup>+</sup>–2−) IV; (10) feet about two-thirds webbed; webbing formula: I (1−–1)—(1<sup>2</sup>/3–2−) II (1−–1)—(1<sup>3</sup>/4–2−) III (1–1+)—(1<sup>2</sup>/3–2+) IV (2−–2<sup>+</sup>)—(1−–1<sup>+</sup>) V; (11) metacarpal, ulnar, metatarsal, and tarsal enameled folds present, with low, enameled tubercles; (12) nuptial pad Type I in adult males; concealed prepollex; (13) Fingers I and II about same length (Finger II length 98%–103% of Finger I length); (14) disc of Finger III small, its width 20.1%–23.5% of eye diameter; (15) in life, dorsum green with white flecks and dots; bones green; (16) in preservative, dorsum cream to light lavender with unpigmented or white flecks and dots; (17) iris greyish white with thin dark grey reticulations and minute yellow flecks; yellow to cream circumpupilary ring borders the pupil; (18) melanophores partially covering dorsal surface of Finger IV, absent from Fingers I–III; (19) males call from upper side of leaves; call emitted as series of three notes (each note = 0.05 s), dominant frequency at 5600–6200 Hz; (20) fighting behavior unknown; (21) egg masses deposited on upper side of leaves; short-term maternal care present; prolonged parental care absent; (22) in tadpoles, tooth row formula 2/3; A-2 tooth row is widely separated in the center; (23) small body size; adult males, SVL 22.0–24.5 mm (*n* = 13); adult females, SVL 25.3–28.3 mm (*n* = 5).

**Figure 216.** *Teratohyla pulverata*, QCAZ 32066. (**A**) Head in lateral view. (**B**) Hand in ventral view. Scale bar = 2 mm. Illustrations by Juan M. Guayasamin.

**Color in life** (Figure 215): Dorsal surfaces of head and body green with small, white dots. Dorsally, limbs green with pale yellow spots, with white centers. Upper lip white; small white dots visible below eye. Ulnar and tarsal folds with low, white tubercles. Cloacal region with numerous small, white warts. Parietal peritoneum transparent (all internal organs visible ventrally). White peritonea covering heart, liver, digestive tract, testes, and gall bladder. Transparent peritonea covering urinary bladder and kidneys. Iris grey white with thin dark grey reticulations and minute yellow flecks; thin yellow line around pupil.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs cream with numerous minute lavender flecks and lacking white iridophores, which are visible in life. Upper lip white; small white spots visible below eye. Low tubercles on ulnar and tarsal folds white, although this coloration is often lost after years of preservation (e.g., KU 116493). Cloacal region with several white warts. Internal organs that are covered by white peritonea in life remain white in preservative (i.e., heart, liver, digestive tract, testes, and gall bladder).

**Biology and ecology:** As in all centrolenid frogs, the reproductive activity of *Teratohyla pulverata* is in vegetation along streams. In Ecuador, *T*. *pulverata* was found sympatrically with *Espadarana prosoblepon* and *H*. *fleischmanni* in Estero Aguacate, and with *Sachatamia ilex*, *S*. *albomaculata*, *E*. *prosoblepon*, and *H*. *aureoguttatum* in Río Bogotá [199]. Delia et al. [25] experimentally demonstrated that short-term brooding provided by females of *T*. *pulverata* greatly impacts the survival of clutches. This form of parental care reduces mortality associated to dehydration, predation, and fungal infection [25].

**Call:** Males of *Teratohyla pulverata* call from the upper surfaces of leaves [24]. The call of is a quick high-pitched "tik", normally repeated several times with roughly a second pause between calls [24]. The dominant frequency is at 5.7–6.2 kHz [149,150].

**Egg masses:** The following information was mostly obtained from Hawley [301]. Unlike other glassfrog egg clutches, the jelly surface is crenulated [17]. The eggs have a yellowish–green coloration and are deposited in a single layer on the upper surface of leaves. The coloration persists for the first five days of development, then changes to pink over the next five days, and lastly to darker shades of red during the remainder of development. The number of eggs per clutch is 23–57 eggs (*n* = 62). Development from oviposition until the last egg hatched was 6–24 days. There was a positive correlation between hatching and nocturnal rainfall. Survivorship per clutch was 85% ± 4% (*n* = 59); 42% percent of clutches hatched without embryonic mortality. Mortality in embryos was caused mainly by flooding (5% or 128 eggs). Desiccation caused mortality in July and August, whereas flooding produced mortality in September. Hatching in embryos in clutches with high mortality is earlier than embryos in clutches with low mortality, perhaps as a response to avoid fungal infection on decaying embryos or predation. Delia et al. [25] reported that clutches contain 59 eggs (±7.5, *n* = 50).

**Tadpole:** The tadpole of *Teratohyla pulverata* was described by Ho ffmann [302]; herein, we present a summary of his description. The ecomorphological guild of the tadpole is exotroph, lotic, and burrower. When hatching (Gosner Stage 24–25) tadpoles have a dark reddish coloration due to the dorsal star-like pigmentation spots and the reddish visceral and muscular system. Young larvae have a nearly circular shape in cross-section, whereas later larval stages change to a typical flat shape. The LTRF is 2/3; the A-2 row had a wide gap; P-3 is slightly shorter than the equal sized P-1 and P-2. The upper jaw sheath forms a smooth arc and has a pointed serration. The lower jaw sheath is smooth and V-shaped. Marginal papillae are present on both sides of the oral disc; the papillae are arranged in a single row but have a wide dorsal gap on the anterior labium. The eye position is dorsal.

**Distribution** (Figure 217): *Teratohyla pulverata* is known from the humid lowlands on the Atlantic versant from north-central Honduras, and on the Pacific slope from southwestern Costa Rica up to 960 m; it is also known from the lowland forest on the western slopes of the Andes of Colombia and Ecuador at elevations below 400 m ([84,101,148,198,199,300,303], this work). In Ecuador, this species is known from localities in the provinces of Esmeraldas and Pichincha at elevations below 400 m (Specimens Examined). In Ecuador, the potential distribution of the species is 25,146 km<sup>2</sup> within the Chocoan Tropical Forest region.

**Conservation status:** Globally, *Teratohyla pulverata* is listed as *Least Concern* by the IUCN [304]. Major threats for species inhabiting the Chocó Ecoregion are contamination associated to mining and deforestation because of agriculture (mainly oil palm) and pasture lands. In Ecuador, because of the mentioned threats, we sugges<sup>t</sup> that the species should be considered as *Endangered*, following IUCN criteria B2a, B2(iii).

**Evolutionary relationships** (Figure 210): Guayasamin et al. [1] mentioned that the placement of *Teratohyla pulverata* in the genus *Teratohyla* is supported by a dataset that includes nuclear and mitochondrial genes, and the combined nuclear genes. They also mentioned that the topology recovered by one particular mitochondrial gene (ND1) strongly supports a clade composed of *Vitreorana castroviejoi* + *V*. *antisthenesi* + *T*. *pulverata*. Most genetic data at hand favor a sister relationship between *T*. *pulverata* and *T*. *amelie* ([2,3,19], this work). Given the lowland distributions of these two species— *T*. *amelie* east of the Andes and *T*. *pulverata* from west of the Andes—the most likely mechanism of speciation is vicariance through the uplift of the Andes.

**Figure 217.** Distribution of *Teratohyla pulverata* in Ecuador (yellow dots).

**Remarks:** From our observations, it is evident that iridophores (white iridophores) are usually maintained in preserved specimens during long periods of time. For example, the specimen KU 85476 was collected on August 1964 and still maintains the white colorations in most of its internal organs (only a portion of the digestive tract has lost the iridophores and now is cream). However, small amounts of iridophores are diffused in ethanol. In *Teratohyla pulverata*, the white dots on the dorsal surfaces and ulnar and tarsal folds are not visible in some preserved specimens. Given the importance of the presence/absence of iridophores in centrolenid taxonomy, it is important that researchers take the time to carefully record and photograph the coloration in life.

**Specimens examined:** *Teratohyla pulverata:* Costa Rica: *Puntarenas:* Rincón de Osa, UCR 17417, USNM 219379–87. Ecuador: *Provincia de Esmeraldas:* Río Durango (1.04186◦ N, 78.62405◦ W; 243 m), BM 1902.5.27.24 (holotype of *H*. *petersi*), BM 1902.5.27.25, QCAZ 32066, QCAZ 45414, DHMECN 2612, 3194–3195; Río Zapayo (0.78333◦ N, 78.9333◦ W), BM 1902.7.29.36–37; Río Bogotá, DHMECN 3194–95; Reserva Ecológica Cotacachi-Cayapas, Charco Vicente, 60 m, QCAZ 11367–68; Estero Aguacate (00◦39- N, 80◦03- W; 10–64 m), Parroquia San Francisco del Cabo, DHMECN 3194–95. *Provincia de Pichincha:* Silanche (0.1333◦ N, 79.1333◦ W; 400 m), QCAZ 32224. Nicaragua: Matagalpa: Finca Tepeyac, 10.5 km N and 9.0 km E of Matagalpa, 960 m, KU 85476. Panama: Coclé: Quebrada Guabalito, Palmarazo, Parque Nacional Omar Torrijos, CH 5122; Darién: Río Jaque, 1.5 km above Río Imamado, 50 m, KU 116493. Honduras: Olancho, USNM 342214–21.

*Teratohyla spinosa* (Taylor 1949 [145]; Figures 218–221).

*Centrolenella spinosa* Taylor, 1949 [145]. Holotype: KU 23809. Type locality: "Los Diamantes, one mile south of Guápiles, (Cantón de Pococí, Provincia de Limón,) Costa Rica". Savage, 1974 [183], commented on the type locality. *Teratohyla spinosa*—Taylor, 1951 [15]. *Cochranella siren*—Ruiz-Carranza and Lynch, 1991 [6]. *Teratohyla spinosa*—Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, Vilà, 2009 [1].

**Common names:** English: Dwarf Glassfrog [24], Minute Spiny Glassfrog. Spanish: Rana de Cristal Enana [24], Rana de Cristal espinosa.

**Etymology:** The specific name comes from the Latin *spina*, meaning thorn, and refers to the spine-like prepollex that is evident next to the thumb (Figure 12).

**Identification:** Among glassfrogs that inhabit the lowlands of western Ecuador (Chocó Ecoregion), *Teratohyla spinosa* is unique in having a uniform green dorsum, a white to yellowish–white iris with contrasting black reticulations (Figure 218), small size (SVL < 21.0 mm), and lacking humeral spines. Also, this species is exceptional by having a spine (prepollex) that is clearly separated from Finger I (Figures 12 and 219).

**Figure 218.** *Teratohyla spinosa* in life. (**Top row**): Adult male from Durango, Esmeraldas province, QCAZ 45410. (**Bottom row**): Amplectant pair from Reserva Otokiki, Esmeraldas province, Ecuador. Photos by Luis A. Coloma.

**Diagnosis:** (1) Vomers with dentigerous process that bear or lack teeth; (2) snout truncated in dorsal aspect; truncated to slightly protruding in lateral profile (Figure 219); (3) tympanum with pronounced lateral inclination, its diameter about 23.5%–32.0% of eye diameter; upper border of tympanic annulus obscured by supratympanic fold; tympanic membrane clearly di fferentiated from surrounding skin; (4) dorsal skin smooth, lacking spicules; (5) ventral skin smooth to slightly areolate; lacking pair of enlarged subcloacal warts; (6) white parietal peritoneum covering anterior 50%–60% of venter (condition P3); 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 absent between Fingers I and II, reduced between Fingers II and III, and extensive between Fingers III and IV (Figure 219); formula: II (1<sup>2</sup>/3–2−)—(3<sup>+</sup>–31/4) III (2−–2)—(1<sup>+</sup>–12/3) IV; (10) distinct prepollex; in males, nuptial pad Type I (see Remarks); (11) Finger I slightly longer than Finger II (Finger II length 90.1%–92.5% of Finger I); (12) ulnar fold low; tarsal folds absents; (13) feet with relatively extensive webbing; formula: I 1—(2−–2) II (1−–1)—(2−–2<sup>+</sup>) III (1<sup>+</sup>–11/2)—(2<sup>+</sup>–21/2) IV (2−–21/2)—(1−–1) V; (14) width of Finger III disc about 33.1%–40.4% of eye diameter; (15) in life, dorsum uniform green; bones green; (16) in preservative, dorsum creamy lavender; (17) in life, iris white to yellowish white, with black contrasting reticulations; (18) melanophores present only on dorsal surfaces of Finger IV (and sometimes Finger III) and Toes IV and V; (19) males call from the upper surfaces of leaves; each call consists of one to three pulsed notes; the dominant frequency is at 6469–6937 Hz; (20) fighting behavior unknown; (21) eggs deposited on the underside of leaves; short-term maternal care present; prolonged parental care absent; (22) oral apparatus of tadpoles with tooth row formula 0/0; (23) minute body size; in males, SVL 18.0–19.7 mm ( X = 19.0 ± 0.525, *n* = 10); in females, SVL 19.7–20.7 mm (*n* = 3).

**Figure 219.** *Teratohyla spinosa*, adult male. ( **A**) Head in lateral view, QCAZ 31321. (**B**) Hand in ventral view, KU 164668. ( **C**) Finger I in dorsal view, QCAZ 10450; note nuptial pad and prepollical spine. Illustrations by Juan M. Guayasamin.

**Color in life** (Figure 218): Yellowish–green dorsum. Anterior half of venter white, posterior part transparent. Green bones. Iris silvery white to yellowish white, with marked black reticulations.

**Color in ethanol:** Dorsal surfaces of head, body, and limbs cream with light lavender hue. Anterior half of venter white, posterior part translucent. Translucent peritonea covering gall and urinary bladders. Iridophores absent from digestive tract, liver, and kidneys.

**Biology and ecology:** In Ecuador, *Teratohyla spinosa* has been found breeding along streams near San Francisco de Bogotá (25 May 2006) and Alto Tambo (April 2010). Individuals were active during the night; several males were calling and amplectant pairs were observed. Females provide short-term parental care, whereas any form of prolonged parental care is absent [25].

**Call** (Figure 220): Males call from the upper surfaces of leaves. Ibáñez et al. [197] described the call as a series of harsh "creep-creeps" followed by a considerable pause, with a dominant frequency at 6.8–7.2 kHz. The description provided below is based on a recording obtained by JMG at Reserva

Otokiki (0.91104◦ N, 78.57369◦ W; 706 m), Esmeraldas province, Ecuador. Calls are emitted every 12–35 s; each call usually has two notes (range = 1–3 notes, X = 1.92 ± 0.515, *n* = 12; Figure 220); calls with one note have a duration of 0.124–0.139 s (*n* = 2); calls with two notes last 0.431–0.594 s (X = 0.545 ± 0.067, *n* = 9); a single three-note call had a duration of 0.793 s. Notes are pulsed, with 5–15 pulses per note (X = 7.87 ± 2.096, *n* = 23). The dominant frequency is at 6469–6937 Hz (X = 6664 ± 129.2, *n* = 12); a first harmonic is visible at 12,843–13,687 Hz (X = 13351 ± 305.6, *n* = 12); a second harmonic is sometimes visible at 19,312–20,812 Hz (X = 20039 ± 9885, *n* = 8).

**Figure 220.** Call of *Teratohyla spinosa*, LBE-C-002, recorded at Reserva Otokiki, 706 m, Esmeraldas province, Ecuador.

**Egg masses and tadpoles:** Egg masses are attached on the underside of the leaves near their margins and 0.5 to 2.5 m above the water and sometimes over the shore [24,146,147]. When laid, the eggs of *Teratohyla spinosa* are greenish white; the embryos are initially yellow and become darker during development, reaching a dark grey, brownish, or reddish–tan color at hatching [24,146,147]. Egg masses contain 15–25 eggs, with each egg and associated capsule measuring 5–7 mm in diameter; clutch mass is small, measuring about 21 × 25 × 7 mm [24,146,147]. The following description of the tadpole is primarily derived from Savage [148], but with some modifications (RWM, pers. obs.). Tiny on hatching, total length about 11 mm at stage 25; body elongated, depressed; mouth ventral; nares and crescent-shaped eyes dorsal; midlateral, posterior, sinistral spiracle; vent tube median; tail long with rounded tip; caudal fins reduced. Oral disc complete, moderate, with single row of marginal papillae (21–45) laterally and ventrally, wide dorsal gap above mouth; LTRF probably 2/3, but when rows are present, rows of teeth are weaker and sometimes irregular; A-1 row often not evident [147]; A-2 usually present, but often visible only as traces of teeth on the lateral extremes of the tooth ridge; jaw sheaths robust and well developed, upper broadly arched and lower one v-shaped, both with sharp serrations along their entire lengths. Body and tail greyish brown dorsally, lighter ventrally; bright yellow yolk mass colors posterior half of the body and covers the liver and intestine; tail musculature mostly brown except pale area anteroventrally; fins are opaquely transparent with scattered dark spots. Starrett [146] and Hoffmann [147] also provided descriptions of the tadpoles. Starrett mentioned that her specimens lacked labial teeth and suggested that possibly this species does not develop labial teeth. Savage [148] considered her specimens to be anomalous and possibly the result of raising the tadpoles

in captivity. Hoffmann agreed, but also noted how variable the tooth rows were in his specimens, which also were lab-raised. A better understanding of the nature of the labial teeth will only become available with the examination of field collected larvae of *T*. *spinosa*.

**Distribution** (Figure 221): *Teratohyla spinosa* is distributed from eastern Honduras, through Nicaragua, Costa Rica, and Panama in Central America [84,134,145,148,305–308]. In South America, it is found in the Chocó Ecoregion of Colombia south to Río Palenque in Ecuador [86,101,134]. In Ecuador, the species has been reported from the provinces of Esmeraldas, Pichincha, and Los Ríos at elevations below 700 m. In Ecuador, the potential distribution of the species is 38,671 km2, within the Chocoan Tropical Forest and the Western Foothill Forest regions.

**Figure 221.** Distribution of *Teratohyla spinosa* in Ecuador (yellow dots).

**Conservation status:** Globally, *Teratohyla spinosa* is listed as *Least Concern* by the IUCN [309]. In Ecuador, the habitat of the species (Chocó) is continuously being fragmented by logging activity and habitat transformation towards oil palm plantations; therefore, we sugges<sup>t</sup> that the species should be considered as *Endangered*, following IUCN criteria B2a, B2(iii).

**Evolutionary relationships** (Figure 210): *Teratohyla spinosa* is sister species to a clade formed by *T*. *midas* + *T*. *adenocheira*. *Teratohyla spinosa* is found in the Chocó, whereas *T*. *midas* and *T*. *adenocheira* are endemic to the Amazon basin. The most like speciation mechanism that separated *T*. *spinosa* from its closest relatives is vicariance through the uplift of the Andes.

**Specimens examined:** *Teratohyla spinosa:* Ecuador: *Provincia de Cotopaxi:* 6 km W of Guasaganda (0.797◦ S, 79.212◦ W), QCAZ 6809. *Provincia de Esmeraldas:* 2 km E of San Francisco de Bogotá, on the San Francisco–Durango road (1.08585◦ N, 78.68575◦ W; 77 m), QCAZ 32124–25; 1.3 km E of San Francisco de Bogotá, on the San Francisco–Durango road (1.0888◦ N, 78.69563◦ W), QCAZ 32138; near Durango (1.07934◦ N, 78.66954◦ W; 74 m), QCAZ 31321; 4 km N of Durango (1.44307◦ N, 77.99667◦ W; 253 m), QCAZ 33319; Bosque Integral Otokiki (0.91241◦ N, 78.58092◦ W; 700 m), QCAZ 48283; 7 km N of Durango in the Durango–San Lorenzo road (1.07934◦ N, 78.66954◦ W; 74 m), QCAZ 31321. *Provincia Los Ríos:* Estación Biológica Río Palenque (0.55◦ S, 79.3667◦ W; 220 m), KU 164663–68. *Provincia Pichincha:* Río Blanco, Río Yambi (00◦01- S, 79◦08- W; ca. 700 m), USNM 288443.

**Localities from the literature:** Ecuador: *Provincia de Esmeraldas:* Estación Biológica Bilsa (0.36667 N, 79.750 W), Reserva Ecológica Mache Chindul [310].

**Photographic record:** Ecuador: *Provincia de Esmeraldas:* Reserva Tesoro Escondido (0.5337 N, 79.1445 W), QCAZ 65398 [294].

*Vitreorana ritae* (Lutz in Lutz and Kloss, 1952 [311]; Figures 222–226).


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

*Centrolenella ametarsia* Flores, 1987 [314]. Holotype: MCZ 96522. Type locality: "the headwaters of Río Caiwima, a tributary of the Río Amaca-Yacu, ca. 70 km NNE Puerto Nariño, Amazonas, Colombia (approximately 3◦20- S, 70◦20- W)". Placed in synonymy by Cisneros-Heredia, 2013 [312].

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


*Vitreorana ritae—*Cisneros-Heredia, 2013 [312].

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

**Etymology:** The name *Vitreorana* has its origin in the Latin words *vitreum*, meaning glass, and *rana*, meaning frog. The name refers to the total or partial ventral transparency of these frogs [1]. The specific name *ritae* is a patronym to Rita Kloss, assistant of Bertha Lutz.

**Identification:** Among glassfrogs, *Vitreorana ritae* is unique by having a green dorsum with small black spots (Figure 222), a small size (≤24 mm), white gastrointestinal peritoneum, and a distinct prepollex (Figure 12). The only glassfrog that could be confused with *V*. *ritae* is *V*. *helenae*, which differs by having a yellow iris (greyish white with a fine dark reticulation in *V*. *ritae*), light greenish–yellow dorsum with dark punctuations (green with dark punctuations in *V*. *ritae*; Figure 222), and a mostly white hepatic peritoneum (hepatic peritoneum mostly transparent, showing the brown liver, except for some iridophores on the upper border in *V*. *ritae* [300].

**Diagnosis:** *Vitreorana ritae* has the following combination of traits: (1) Dentigerous process of vomer with one tooth or lacking teeth; (2) snout rounded in dorsal and lateral views (Figure 223); (3) tympanum visible, moderate in size, its diameter 25.8%–35.4% of eye diameter; tympanic annulus visible except for upper border covered by supratympanic fold; tympanic membrane differentiated and translucent, pigmented as surrounding skin; (4) dorsal surfaces shagreen; spinules absent; (5) ventral surfaces granular, pair of enlarged subcloacal warts; (6) anterior 25%–40% of ventral parietal peritoneum white, posterior portion transparent (condition P2–P3); pericardium and gastrointestinal peritoneum white (condition V2); (7) lobed liver covered by transparent peritoneum, except for anterior border that may be covered by thin layer of iridophores (condition H0); (8) humeral spines absent; (9) webbing between Fingers I–III absent, moderate between Fingers III and IV (Figure 223); webbing formula: III (2−–21/3)—(1<sup>+</sup>–2−) IV; (10) webbing between toes moderate; webbing formula: I 1—(2−–2) II (1–1+)—(2–2<sup>1</sup>/4) III (1<sup>+</sup>–11/2)—2<sup>+</sup> IV (2–2<sup>1</sup>/3)—1 V; (11) low ulnar fold, lacking iridophores; low inner tarsal fold present, lacking iridophores; outer tarsal fold absent; (12) nuptial pad of males Type I; distinct prepollex; (13) Finger I slightly longer than Finger II (Finger II length 84%–92% of Finger I); (14) disc of Finger III moderate, its width 31.0%–42.3% of eye diameter; (15) in life, dorsum green with small dark spots (Figure 222); bones green; (16) in preservative, dorsum lavender with dark spots; (17) iris greyish white with thin dark reticulation; (18) melanophores covering dorsal surfaces of Fingers III and IV; (19) males call from the upper side of leaves; single and double note advertisement call of 0.10–0.15 s duration, dominant frequency at 4640–5160 Hz; (20) fighting behavior unknown; (21) eggs deposited on upper or underside of leaves; short-term maternal care present; prolonged parental care absent; (22) tadpoles at stage 25 with labial tooth row formula 0/1–2; oral disc small and ventral with one row of large marginal papillae laterally and posteriorly; upper jaw sheath wide and robust, lower jaw sheath wide, V-shaped; dorsum reddish brown, venter whitish, tail muscle reddish and tail fins transparent (Figure 224); (23) minute body size; adult male SVL 17.1–20.1 mm (X = 18.8 ± 1.250, *n* = 6); two adult females SVL 19.8–19.9 mm; Lima et al. [315] provided the following data for the species in central Amazonia: SVL in males 17–21 mm, in females 21–24 mm.

**Figure 222.** *Vitreorana ritae* from Yasuní National Park, Ecuador. Photo by Jaime Culebras.

**Figure 223.** *Vitreorana ritae*. (**A**) Head in dorsal view, MCZ 96522. (**B**) Head in lateral view, MCZ 96522. (**C**) Hand in ventral view, with exposed prepollex, ICN 50847. (**D**) Toe I in ventral view, with papilla at tip. Not drawn to scale. (**A**,**B**) Modified from Flores [314]; (**C**,**D**) by Juan M. Guayasamin.

**Color in life** (Figure 222): Green dorsum with small dark blue to black spots. Green bones. Iris greyish white with thin dark reticulation. Venter white anteriorly, turning transparent posteriorly; white digestive tract.

**Color in ethanol:** Dorsum lavender with small dark lavender spots. Color of parietal and visceral peritonea variable; holotype and one Colombian specimen (ICN 50847) with transparent parietal and visceral peritonea. Two specimens (ICN 50846, QCAZ 28138) with white parietal peritoneum on anterior half of venter, and white peritoneum on intestines and stomach. We consider that the white lining has been dissolved in the holotype and ICN 50847, but it is equally possible that *Vitreorana ritae* is polymorphic for the presence of white lining on the parietal and visceral peritonea. It may also be possible that the species is a composite of more than one species.

**Biology and ecology:** During the night, *Vitreorana ritae* has been found on vegetation, about 1–5 m above streams, and on the trunk of a *Ceiba*, about 7 m above the ground [300,314]. At Reserva Florestal Adolpho Ducke in Manaus, Amazonas, Brazil, breeding individuals and clutches of *V*. *ritae* were observed on vegetation above small streams in forests [300]. Adults were found throughout the year, and vocalizations were heard throughout the night and were more common between January and May (from the middle to the end of the rainy season). Short-term maternal care is present, whereas any form of prolonged parental care is absent [25].

**Egg masses and tadpoles** (Figure 224): The information shown below is from Menin et al. [316]. At Reserva Florestal Adolpho Ducke, Amazonas, Brazil, clutches were deposited on the upper (*n* = 4) or undersides (*n* = 2) of leaves hanging over a stream, from 0.5 to 1 m above water surface. Mean clutch size was 16 eggs (*n* = 4; range 12–18). The eggs were green and deposited in a single layer with adhered jelly capsules. Tadpoles were green while in the egg capsules but became reddish brown with transparent tail after hatching. Tadpoles from a single clutch raised in the laboratory hatched at Gosner's [191] stage 24. Tadpoles were found buried in the leaf litter along the edges of slow-flowing streams. The description below is based on eleven tadpoles at stage 25. In life, dorsum reddish brown, venter whitish, tail muscle reddish and tail fins transparent. In preserved specimens, dorsum transparent with dark melanophores, venter whitish with melanophores on posterior part and tail translucent with melanophores on tail musculature. Body elongate and depressed in lateral view, and rectangular and elongate in dorsal view. Body and tail 27%–28% and 72%–73% of total length, respectively. Body wider than deep; maximum body width behind eyes. Snout broad and truncated in dorsal and ventral views and flattened in lateral view. Eyes small, close together, and facing dorsolaterally. Interorbital distance six times larger than maximum eye diameter. Narial openings small, nearer snout than eyes. Spiracle single, short, sinistral, positioned on longitudinal axis of body and directed posteriorly. Vent tube short, positioned along ventral midline and attached directly to ventral fin. Tail musculature robust; caudal fins low, similar in height, deeper than the robust caudal musculature only on the posterior one-third of the tail. Dorsal fin originates at tail–body junction and increases in depth throughout the first third of the tail, and then gradually decreases to a rounded tip. Ventral fin originating on the tail musculature, slightly arched, and maintaining the same height throughout the distal two-thirds of the tail. Tail tip rounded. Oral disc small and ventral (Figure 224), bordered laterally and posteriorly by one row of large marginal papillae with pointed tips. Marginal papillae on posterior border larger than lateral ones. Labial tooth row formula (LTRF) 0/1–2. All labial tooth rows nearly the same length. Upper jaw sheath wide and robust; lower jaw sheath wide, V-shaped; both hardly serrated.

**Figure 224.** Tadpole of *Vitreorana ritae* at Gosner stage 25. (**A**) Lateral view. (**B**) Dorsal view. (**C**) Oral disc. Modified from Menin et al. [316].

**Call** (Figure 225): At central Amazonia of Brazil, males of *Vitreorana ritae* were observed calling on vegetation above streams, about 0.5 to 3 m high on leaves of aquatic or stream side plants (*n* = 20) [316]. Herein we report a call obtained by Morley Read at Boanamo community (LBE-C-039; 240 m) and Shiripuno Lodge (LBE-C-039; 250 m), Orellana province, Ecuador. We analyzed 31 notes contained within 10 calls from two individuals. The typical advertisement call is relatively short (range = 288–1095 ms, mean = 685.2 ms, SD = 280.1 ms) and contains two to five notes per call (mean = 3.3, SD = 1.1). Each note has a duration of 61–122 (mean = 76, SD = 16.3) ms. Notes are strongly pulsed and have 10–19 (mean = 13.7, SD = 2) amplitude peaks throughout the note, whereby some pulses are difficult to delimit because of a lower amplitude. Pulses within a note have a rate of 131–224 (mean = 171, SD = 24) pulses per second. Notes have their peak amplitude in the last 50% of the note (relative peak time: Range = 0.4425–0.8815, mean = 0.719, SD = 0.106), where the amplitude increases from a lower to a higher amplitude at the end of the note. The dominant frequency of a note measured at peak amplitude is 4688–5344 (mean = 5044, SD = 182) Hz and is contained within the fundamental frequency. The fundamental frequency has a lower limit of 4312–4875 (mean = 4605, SD = 164) Hz and a higher limit of 4969–6029 (mean = 5284, SD = 229) Hz. The call described for the Ecuadorian populations matches the general information provided by Zimmerman and Bogart [317] for Brazilian populations.

**Figure 225.** Calls of *Vitreorana ritae*, LBE-C-039, recorded by Morley Read at Boanamo Community, 240 m, Orellana province, Ecuador. (**A**) A single call with four notes. (**B**) A single note.

**Distribution** (Figure 226): *Vitreorana ritae* is found from the Guianas across western Amazonia [17,222,229,300,313–316,318]. In Ecuador, the species has been reported from five localities in the provinces of Orellana and Sucumbíos at elevations below 260 m (Specimens Examined). In Ecuador, the potential distribution of the species is 42,600 km<sup>2</sup> within the Amazonian Tropical Rainforest region.

**Conservation status:** *Vitreorana ritae* is listed as *Data Deficient* by the IUCN [297]. This species is known only from a handful of localities but is likely to occur more widely in the Amazon basin. Further surveys with emphasis on sampling bromeliads may provide a better understanding of the population dynamics of this species [166,319]. Given the broad distribution of the species in the Amazon basin, we sugges<sup>t</sup> that it should be placed in the *Least Concern* category.

**Evolutionary relationships** (Figure 227): *Vitreorana ritae* is sister to *V*. *helenae*.

**Taxonomic remarks:** Cisneros-Heredia [312] placed *C*. *ametarsia and C*. *oyampiensis* under the synonymy of *C*. *ritae*, arguing that several of the traits described by Lutz [311] were misinterpreted and that Lutz's description of apparently "large size" of the discs of *C*. *ritae* was misrepresentative. We adopt the change proposed by Cisneros-Heredia [312] but note that new material of *ritae* from its type locality ("Benjamin Constant, Alto Solimões", Estado do Amazonas, Brazil) will provided a definitive solution to any taxonomic uncertainty.

**Figure 226.** Distribution of *Teratohyla ritae* in Ecuador (yellow dots).

**Figure 227.** Evolutionary relationships of species in the glassfrog genus *Vitreorana*. The tree was inferred using maximum likelihood and Bayesian criteria.

**Specimens examined:** *Teratohyla ritae:* Colombia: *Departamento de Amazonas:* headwaters of Río Caiwima, tributary of the Río Amayaca-Yacu, MCZ A-96522 (holotype of *C*. *ametarsia*, neotype of *V*. *ritae*); Leticia, ICN 50846–47, ICN (JDL 24472). Ecuador: *Provincia de Orellana:* Río Yasuní, 200 km upstream from Río Napo (0.85◦ S, 76.383◦ W; 200 m), KU 175216; Tiputini Biodiversity Station (0.65◦ S, 76.13◦ W; 210 m), DFCH-USFQ D162; Estación Científica Yasuní PUCE, 240 m, QCAZ 16652; Yasuní, km 8 on the Pompeya-Iro road, 260 m, QCAZ 22709. *Provincia de Sucumbíos:* Puerto Bolívar (0.09◦ S, 76.14◦ W; 240 m), 260 m, QCAZ 28138.
