http://zoobank.org/urn:lsid:zoobank.org:act:AD71F7CC-0718-449E-88DC-0A64302D9855.

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

**Etymology:** The specific epithet *humboldti* honors Alexander von Humboldt for his unparalleled contributions to biogeography, integrative perspective of the sciences and arts, humanism, and also for his devotion towards mountains.

**Holotype:** ZSFQ 0388, adult male, collected from Volcán Sumaco (0.61387◦ S, 77.590◦ W; 1738 m), Napo Province, Ecuador, by Jose Vieira on 11 September 2018.

**Paratopotypes:** QCAZ 9402, adult male collected by Juan M. Guayasamin on 19 February 1996; QCAZ 41071, 41073–74, 41077–78, 41081–82, 41150–51, 41314–15, adult males and females, collected by Elicio Tapia and Raúl E. Ruíz on 20–30 March 2009.

**Paratypes:** QCAZ 45712–16, 45730, 47511–14, adult males and females, collected by Elicio Tapia from Río Yana Challuwa Yaku (1.26764◦ S, 78.04797◦ W; 1800–2400 m), Reserva Comunitaria Ankaku, Pastaza Province.

**Generic placement:** The new species is placed in the clade *Nymphargus* (sensu Guayasamin et al. [1]) based on morphological and molecular data. All species in *Nymphargus* share an absence of webbing between Fingers I–III and absence or reduced webbing between Fingers III and IV; additionally, males lack humeral spines (except *N*. *grandisonae*). *Nymphargus humboldti* sp. nov. also has these traits. Molecular analyses of mitochondrial genes (12S, 16S, ND1) unambiguously place the new species in the clade *Nymphargus* (Figure 16).

**Identification:** Among glassfrogs that inhabit the Amazonian slopes of the Andes and the Amazonian lowlands, *Nymphargus humboldti* sp. nov. (Figure 158) is distinguished by having a green dorsum with small yellow spots and moderate body size (male SVL 23.3–25.2 mm; female SVL 24.3–25.9 mm), and lacking hand webbing and humeral spines. *Nymphargus humboldti* sp. nov. is most similar to *N*. *siren*, which is smaller (SVL in males, 19.8–22.6 mm; in females, 22.5–23.3 mm). Other species with a similar dorsal color pattern include *Espadarana audax*, *Nymphargus cariticommatus*, *N*. *siren*, *Rulyrana flavopunctata*, *R*. *mcdiarmidi*, and *Teratohyla midas*. Differences among these species include the presence of hand webbing between Fingers III and IV in *E*. *audax*, *R*. *flavopunctata*, *R*. *mcdiarmidi*, and *T*. *midas* (absent in *N*. *humboldti* sp. nov.), white digestive tract in *T*. *midas* (opaque in *N*. *humboldti* sp. nov.), and humeral spine in males in *E*. *audax* (absent in *N*. *humboldti* sp. nov.). *Nymphargus cariticommatus* differs by having white renal peritoneum and white esophagus (transparent renal pericardium and

opaque esophagus in *N*. *humboldti* sp. nov.) and less foot webbing. Two additional species endemic to Colombia, *N*. *luminosus* and *N*. *spilotus*, can be confused with *N*. *humboldti* sp. nov. Both species are conspicuously larger than *N*. *humboldti* sp. nov. (*N*. *luminosus*, male SVL = 27.8–30.0 mm, female SVL = 32.7 mm; *N*. *spilotus*, male SVL = 25.3–26.4 mm, female SVL = 27.6–28.5 mm); also, *N*. *luminosus* in only found on the Pacific flank of the Andes in Colombia, whereas *N*. *spilotus* is only know from its type locality (Rancho Quemado, Corregimiento Florencia, Departamento de Caldas) on the eastern slope of Cordillera Central.

**Figure 158.** *Nymphargus humboldti* sp. nov. in life. Male from Río Yana Challuwa, Pastaza province, Ecuador, QCAZ 47514, paratype. Photos by Luis A. Coloma.

**Diagnosis:** (1) Vomers with edentate dentigerous process; (2) snout truncated to bluntly rounded in lateral profile; truncated in dorsal view; (3) tympanum oriented almost vertically, with slight lateral and posterior inclinations, its diameter about 20%–27% of eye diameter; upper fourth of tympanic annulus obscured by supratympanic fold; tympanic membrane pigmented as surrounding skin; (4) dorsal skin shagreen, with minute spicules in males; (5) venter areolate; pair of enlarged subcloacal warts; (6) white parietal peritoneum covering anterior 50%–60% of venter (condition P2); white pericardium; translucent peritonea covering intestines, stomach, kidneys, gall and urinary bladders (condition V1); (7) liver tetralobed, covered by transparent peritoneum (condition H0); (8) humeral spines absent; (9) webbing absent between inner fingers, absent or basal between Fingers III and IV; webbing formula III (3−–3)—(2<sup>2</sup>/3–3−) IV; (10) feet about two-thirds webbed; webbing formula: I (2−–2)—(2<sup>+</sup>–21/4) II (1<sup>+</sup>–11/2)—(2<sup>+</sup>–21/2) III (1<sup>+</sup>–11/4)—(21/4–3<sup>+</sup>) IV (2<sup>1</sup>/2–3−)—(1<sup>2</sup>/3–2−) V; (11) ulnar and tarsal folds present, low; (12) concealed prepollex; in males, nuptial pad Type I; (13) Finger I about same length as Finger II (Finger I length 94%–102% of Finger II); (14) disc width of Finger III about 50%–55% of eye diameter; (15) in life, dorsum green with yellow spots (Figure 158); bones green; (16) in preservative, dorsum lavender with small white spots; (18) melanophores usually lacking from dorsal surfaces of fingers and toes, except for few on Toe V; (19) males call from upper surfaces of leaves; call unknown; (20) fighting behavior unknown; (21) egg deposition site unknown; parental care unknown; (22) tadpoles unknown; (23) small body size; in males, SVL 23.3–25.2 mm (X = 24.3 ± 0.684, *n* = 13); in females, SVL 25.5–27.4 mm (*n* = 3).

**Description of holotype:** Adult male, SVL 24.9 mm. Head slightly wider than long (head length 98% of head width); snout truncated in dorsal and lateral profiles; canthus rostralis indistinct, slightly concave; loreal region slightly concave; lips slightly flared; nostril protuberant, closer to tip of snout than to eye, directed frontolaterally; internarial area barely depressed. Eyes large, directed anterolaterally at about a 50◦ angle; transverse diameter of disc of Finger III 53% eye diameter. Supratympanic fold low, obscuring upper portion of tympanic annulus; tympanum oriented mostly vertically, but with slight posterolateral inclination; tympanic membrane transparent, its upper half pigmented as surrounding skin. Dentigerous processes of vomer low, situated transversely between choanae, lacking teeth; choanae large, longitudinally rectangular; tongue ovoid, with ventral posterior fourth not attached (free) to floor of mouth, posterior margin notched; vocal slits extending posterolaterally from a point about midway along lateral margin of tongue) to angle of jaws. Humeral spine absent. Low ulnar fold evident on external ventrolateral margin of arm; inner ulnar fold absent; relative lengths of fingers: III > IV > II > I; webbing absent between Fingers I, II, and III, basal between Fingers III and IV, webbing formula III 3−–22/<sup>3</sup> IV; discs expanded, nearly elliptical; disc pads nearly triangular in shape; subarticular tubercles small, round, simple; supernumerary tubercles absent; palmar tubercle elliptical, simple; nuptial pad large (Type I), ovoid, granular, extending from ventrolateral base to dorsal surface of Finger I, covering proximal half of Finger I. Length of tibia 58% SVL; low inner tarsal fold evident; outer tarsal fold absent; two-thirds webbed; webbing formula of foot: I 2—2<sup>+</sup> II 1<sup>+</sup>—21/<sup>4</sup> III 1<sup>+</sup>—3<sup>+</sup> IV 3−—2− V; discs on toes round to elliptical; disc on Toe IV narrower than disc on Finger III; disc pads triangular; inner metatarsal tubercle large, ovoid; outer metatarsal tubercle round, barely evident; subarticular tubercles small, round; supernumerary tubercles absent. Skin on dorsal surfaces of head, body, and lateral surface of head and flanks shagreen with numerous minute spinules; throat smooth; belly and lower flanks areolate; cloacal opening directed posteriorly at upper level of thighs; cloacal ornamentation absent. Two enlarged tubercles below the level of vent.

**Coloration of the holotype in life** (Figure 158): Green dorsum with small yellow spots; upper lip white. Anterior 60% of venter white, posterior portion transparent. Bones green. Iris greyish cream with slight yellow hue and fine black reticulations; yellow circumpupilary ring.

**Coloration of the holotype in ethanol:** Dorsal surfaces of head, body, and limbs lavender with small white spots; white upper lip. Anterior 60% of ventral parietal peritoneum white. Heart covered by white pericardium; translucent peritonea on gall and urinary bladders; iridophores absent from digestive tract, liver, and kidneys.

**Measurements of holotype (mm):** *Nymphargus humboldti* sp. nov., adult male, ZSFQ 0388: SVL = 24.9, head length = 8.1, head width = 8.3, eye–nostril = 1.9, nostril–snout = 1.1, IOD = 2.6, upper eyelid width = 2.3, eye diameter = 3.5, tympanum diameter = 0.8, tibia = 14.5, foot length = 11.5, radio–ulna length = 5.5, hand = 8.4, Finger I length = 4.9, Finger II length = 5.2, disc of Finger III width = 0.9

**Biology and ecology:** At Río Yana Challuwa Yaku, *Nymphargus humboldti* sp. nov. was found in sympatry with *N*. *anomalus*. During the night, males were calling from the upper surfaces of leaves near a fast-flowing stream. Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 159): *Nymphargus humboldti* sp. nov. is known from few localities on the Amazonian slopes of the Ecuadorian Andes at elevations between 1770 and 2400 m.

**Figure 159.** Distribution of *Nymphargus humboldti* sp. nov. in Ecuador (yellow dots).

**Conservation status:** Following IUCN criteria, we sugges<sup>t</sup> placing *Nymphargus humboldti* sp. nov. in the *Data Deficient* category.

**Evolutionary relationships** (Figure 136): *Nymphargus humboldti* sp. nov. is sister to *N*. *siren*.

*Nymphargus lasgralarias* Hutter and Guayasamin, 2012 [88] (Figures 154 and 160–162).

*Nymphargus lasgralarias* Hutter and Guayasamin, 2012 [88]. Holotype: MZUTI 096.

Type locality: "Five Frog Creek" (0◦01.870- S, 78◦42.358- W; 2150 m) at Reserva Las Gralarias, Pichincha province, Ecuador".

**Common names:** English: Las Gralarias Glassfrog. Spanish: Rana de Cristal de Las Gralarias. **Etymology:** The epithet *lasgralarias* refers to the type locality of the species, Reserva Las Gralarias. The species was dedicated to the reserve and the team of people, led by Dr. Jane Lyons, for their efforts to conserve Ecuadorian cloud forests [88].

**Identification:** Among *Nymphargus* species found on the Pacific versant of the Andes of Ecuador, *N*. *lasgralarias* can only be confused with *N*. *buenaventura* and *N*. *gri*ffi*thsi*. Dorsal texture and color pattern readily separate *N*. *buenaventura*, which, in life, has a light-green dorsum with warts corresponding to pale yellow spots, whereas the dorsum of *N*. *lasgralarias* is homogenously green and lacks warts. Additionally, *N*. *buenaventura* is smaller (male SVL in *N*. *lasgralarias* = 24.6–26.5 mm; male SVL in *N*. *buenaventura* = 20.9–22.4 mm). *Nymphargus lasgralarias* is most similar to *N*. *gri*ffi*thsi*. The two species differ in the following traits: (i) Dorsal color pattern: Homogenously green in *N*. *lasgralarias;* green with very small black spots in *N*. *gri*ffi*thsi* (Figure 154); (ii) body size: Male SVL in *N*. *lasgralarias* = 24.6–26.5 mm, whereas male SVL in *<sup>N</sup>*.*gri*ffi*thsi* = 22.5–24.2 mm; and call: Short, pulsed note lasting 0.016–0.044 s in *N*. *lasgralarias;* long tonal note lasting 0.103–0.148 s in *N*. *gri*ffi*thsi*. Two Colombian species from the Pacific slopes of the Andes, *N*. *cristinae* [255] and *N*. *prasinus* [120], can also be confused with *N*. *lasgralarias*. *Nymphargus lasgralarias* is distinguished from *N*. *cristinae* by being smaller (male SVL in *N*. *lasgralarias* = 24.6–26.5 mm; male SVL in *N*. *cristinae* = 26.0–31.1 mm), having a snout that is truncated in dorsal view and protruding in lateral view (subacuminate in dorsal view, truncated in lateral view in *N*. *cristinae*; see Ruiz-Carranza and Lynch [255]), and lacking palmar supernumerary tubercles (supernumerary small, abundant in *N*. *cristinae*). *Nymphargus prasinus* shares the same color pattern as *N*. *lasgralarias*, but is recognized by having a round snout in dorsal view (truncated *N*. *lasgralarias*), five to seven teeth on each process of the vomer (teeth absent in *N*. *gralarias*), and being considerably larger (male SVL 33.0–34.5 mm; *n* = 3) [88].

**Figure 160.** (**Top Row**): Iris of *Nymphargus lasgralarias*. (**Bottom Row**): Iris of *N*. *gri*ffi*thsi*. Modified from Hutter and Guayasamin [88].

**Characterization.** (1) Vomerine teeth absent; (2) snout truncated in dorsal profile, protruding in lateral profile; (3) tympanum small; supratympanic fold present; tympanic membrane translucent, pigmented only on its upper half; (4) skin texture finely shagreen, with microspiculations; (5) ventral skin areolate, with pair of large subcloacal warts; cloaca surrounded by low warts; (6) upper half of ventral parietal peritoneum covered by iridophores (condition P2), all other peritonea translucent, except for thin layer of iridophores covering heart and renal capsules (condition V1); (7) liver lobed, lacking iridophores (condition H0); (8) humeral spines absent; (9) webbing absent between fingers; (10) foot about half webbed; webbing formula: I (2–2−)—(2<sup>+</sup>–21/2) II (2–2−)—(3−–3) III (2−–2)—(3−–3) IV (3–3+)—2 V; (11) ulnar and tarsal folds low, barely evident; (12) nuptial pad Type I; concealed prepollex; (13) first finger slightly shorter than second; (14) eye diameter larger than width of disc on Finger III; (15) in life: Green dorsum, lacking spots; (16) in preservative: Dorsum pale lavender; (17) iris golden yellow, with numerous small black spots; weakly reticulate; (18) hands and feet yellowish green; melanophores absent from fingers and toes or, when present, restricted to dorsal surfaces of Finger IV and Toes IV and V; (19) males call from upper side of leaves along streams; vocalizations emitted in series of one to five calls; each call pulsed, with a duration of 0.016–0.044 s, and non-modulated dominant frequency at 3618–3963 Hz; (20) fighting behavior unknown; (21) short-term maternal care present; parental care by males absent; (22) egg clutches deposited on upper surface of leaves at terminal margin, transitioning to hanging as eggs develop; (23) tadpoles unknown; (24) SVL in adult males 24.6–26.5 mm (mean = 25.3 ± 0.7368; *n* = 7); in females 26.3–27.2 mm (*n* = 2).

**Color in life** (Figure 154, Figure 160, Figure 161): Dorsum light green, lacking dark spots; flanks yellowish white; bones green; fingers and toes yellow with a faint green tint. Venter white anteriorly and translucent posteriorly. Iris background golden with numerous dark spots and very light reticulation [88].

**Color in ethanol:** Dorsal surfaces of head and body are cream; fingers and toes cream. Upper half of ventral parietal peritoneum covered by iridophores (white), all other peritonea translucent, except for thin layer of iridophores covering heart and renal capsules [88].

**Biology and ecology:** The following information is taken from Hutter and Guayasamin [88]. *Nymphargus lasgralarias* inhabits small, permanent streams (ca. 3 m width) within primary montane forest with minimal disturbance. The species is active during the night and emits advertisement calls from the tops of small-sized ferns, small leaves, and long palm leaves, 1–9 m above the stream. Fighting behavior is unknown, but the description by Duellman and Savitzky [5] could apply to *N*. *lasgralarias* as well. Small clutches (12–36 eggs per mass; *n* = 23) are deposited on the upper surface of a leaf near its terminal margin, and transition into hanging masses as the eggs develop (Figure 160). Tadpoles are unknown. At Reserva Las Gralarias, the species has been found at Ballux Creek, Five-frog Creek, Heloderma Creek, Hercules Creek, Chalguayacu Creek, Kathy's Creek, and Lucy's Creek. *Nymphargus lasgralarias* occurs sympatrically with six other centrolenid species: *Centrolene ballux*, *C*. *heloderma*, *C*. *lynchi*, *C*. *peristicta*, *Nymphargus grandisonae*, and *N*. *gri*ffi*thsi*. Reproductive activity was recorded between April 5 and July 1 2011, but increased dramatically at the end of April and peaked in the middle of May [88]. Females provide short-term parental care; male parental care is absent [25].

**Figure 161.** *Nymphargus lasgralarias* from Reserva Las Gralarias, Pichincha province, Ecuador. (**Top**): Adult male. (**Bottom**): Spider (Ctenidae) preying on egg clutch. Photos by Jaime Culebras.

**Call** (Figure 156): Males emit calls during the night, from the tops of small-sized ferns, small leaves, and long palm leaves, 1–6 m above the stream. Calls are pulsed and emitted in series (one to four calls per call series; mean = 2.7 ± 0.7 calls). Five-call series are emitted sporadically. Each series has duration of 0.033–2.541 s (mean = 1.529 ± 0.597 s) and an interval of 8.6–78.6 s (mean 33.8 ± 18.4 s) between series, with an interval of 0.088–1.513 s (mean = 0.873 ± 0.205 s) between calls within a series. The call repetition rate is 2.0–9.9 (5.5 ± 2.7) calls per minute (*n* = 6 individuals), each call sounding like a "tick" or "click"; call duration is 0.0160–0.0440 s (mean = 0.0257 ± 0.0058; *n* = 119) and dominant frequency is at 3445–3962 Hz (mean = 3691 ± 131.9 Hz) [88].

**Tadpole:** Not described.

**Distribution** (Figure 157): *N lasgralarias* is known from localities on the western slope of the Ecuadorian Andes at elevations between 1850–2300 m. Specifically, it has been found within the limits of the Pichincha and Cotopaxi provinces ([87,88], this work). Individuals that resemble *N*. *lasgralarias*, but are genetically different, are found in northern Ecuador (see Specimens examined).

**Conservation status:** *N. lasgralarias* has not been evaluated by the IUCN. Main threats are habitat destruction and fragmentation due to agriculture and cattle, introduced species (trout), infectious diseases (*Batrachochytrium dendrobatidis*), and climate change. At Reserva Las Gralarias, the species is infected by the chytrid fungus *B*. *dendrobatidis*, but no recent declines have been observed [92]. Following IUCN criteria (B2a, biii, biv), we sugges<sup>t</sup> considering the species as *Endangered*.

**Figure 162.** Distribution of *Nymphargus lasgralarias* in Ecuador (yellow dots).

**Evolutionary relationships** (Figure 136): *N*. *lasgralarias* and *N*. *gri*ffi*thsi* are sister taxa.

**Specimens examined:** *Nymphargus lasgralarias:* Ecuador: *Provincia de Cotopaxi:* Bosque Integral Otonga (0.55 S, 79.46667 W; 2000 m), QCAZ 13115; Reserva Otonga (0.676 S, 76.397 W; 1950 m), QCAZ 11689–90. *Provincia Imbabura: Provincia de Pichincha:* Five Frog Creek (0◦01.870- S, 78◦42.358- W; 2150 m) at Reserva Las Gralarias, MZUTI 096 (holotype), MZUTI 091–095, 097; Kathy's Creek (0◦01.398- S, 78◦43.772- W; 2000 m), Reserva Las Gralarias, MZUTI 091–095; Hercules Giant Tree Frog Creek (0◦01.529- S, 78◦42.243- W; 2175 m), Reserva Las Gralarias, MZUTI 097; Nanegal Grande (0.1167 N, 78.6667 W; 2300 m), QCAZ 46012; 9 km SE Tandayapa (0.01667 S, 78.6833 W; 2160 m), KU 164577–87.

*Nymphargus* <sup>a</sup>ff. *lasgralarias:* Ecuador: *Provincia de Imbabura:* 23.2 km W of Apuela, Cordillera de Intag (0.2666 N, 78.6 W; 2190 m), KU 178122–36; San Antonio de Cuellaje, Finca de Estuardo Ayala (0.4775 N, 78.56263 W), QCAZ 42164; Santa Rosa, Reserva Alto Chocó (0.36939 N, 78.44942 W; 2104 m), QCAZ 31768.

*Nymphargus laurae* Cisneros-Heredia and McDiarmid, 2007 [17] (Figures 163 and 164).

*Nymphargus laurae* Cisneros-Heredia and McDiarmid, 2007 [17]. Holotype: USNM 288453. Type locality: "Loreto, Upper Rio Napo (=Loreto region, near the town of Loreto) (ca. 77◦20- S, 00◦40- W, ca. 500 m elevation), lower slopes of the Sumaco Volcano, on the Cordillera Oriental, eastern slopes of the Andes, Provincia de Orellana, República del Ecuador".

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

**Etymology:** The specific epithet is a patronym for Laura Heredia, DFCH's grandmother, for her support of Diego's interest in animals and science [17].

**Identification:** *Nymphargus laurae* can be distinguished from most centrolenids by the presence of dorsal ocelli (a trait shared only with *N*. *anomalus*, *N*. *lindae* sp. nov., *N*. *cochranae*, and *N*. *ignotus*), two papillae on each toe except Toe V (Figure 163), and relatively small size (SVL = 19.9 mm in male holotype). *Nymphargus anomalus* differs by having smaller and more abundant ocelli, and dark spots and punctuations amidst the ocelli. *Nymphargus lindae* sp. nov. differs from *N*. *laurae* mainly by being larger (in *N*. *lindae* sp. nov., male SVL = 23.0–26.3 mm; female SVL = 27.2–27.8 mm), having vomerine teeth (absent in *N*. *laurae*), and lacking papillae on toe discs. *Nymphargus ignotus* differs by having, in life, a tan dorsal coloration, with smaller and more abundant ocelli, and by being larger (22.3–25.4 mm SVL in males of *N*. *ignota*); further, *N*. *ignotus* is known only from western Colombia around 1900 m, whereas *N*. *laurae* is known from the slopes of Volcán Sumaco, on the Amazonian slopes of the Andes. *Nymphargus laurae* is very similar to *N*. *cochranae;* both species share a similar color pattern and are likely to be sympatric; *N*. *cochranae* however differs by having much smaller ocelli, no ocelli on forearms and shanks, and larger body size (male *N*. *cochranae* SVL = 23.8–26.7 mm), and lacking papillae on the toes. See Taxonomic Remarks.

**Figure 163.** *Nymphargus laurae* in preservative, USNM 288453, from Ecuador, lower slopes of Volcán Sumaco. (**A**) Dorsal view of holotype. (**B**) Ventral view of foot; note presence of two papillae on each toe disc, except Toe IV. Photos by James Poindexter.

**Diagnosis:** The following traits characterize *N*. *laurae:* (1) Vomerine teeth absent; (2) snout truncated in dorsal and profile views; nostrils slightly elevated producing a slight depression in the internarial area; loreal region concave; (3) tympanic annulus evident, oriented dorsolaterally with dorsal inclination; weak supratympanic fold from behind eye to insertion of arm; (4) dorsal skin slightly shagreen with elevated warts corresponding to ocelli, and scattered spicules; (5) ventral skin granular; pair of large, round, flat subcloacal warts; other cloacal ornamentation absent; (6) parietal peritoneum white, covering anterior two-thirds of abdomen (condition P2); white pericardium; all other peritonea clear (condition V1); (7) liver lobed, hepatic peritoneum clear (condition H0); (8) humeral spine absent; (9) webbing basal between Fingers I, II and III, outer fingers III 22/3—21/<sup>2</sup> IV; (10) webbing on feet I 2−—2<sup>+</sup> II 11/2—2<sup>+</sup> III 1<sup>+</sup>—21/<sup>2</sup> IV 21/2—11/<sup>2</sup> V; (11) no dermal folds or tubercles on hands, forearms, feet, or tarsi; (12) unpigmented nuptial pad Type I; concealed prepollex; (13) second finger longer than first; (14) eye diameter greater than width of disc on Finger III; (15) color in life, green with yellow spots surrounded by black ocelli; (16) color in preservative, dorsal surfaces tan cream with dark, reddish–lavender ocelli; (17) iris coloration in life unknown; (18) melanophores absent on fingers and toes; (19) calling site and call unknown; (20) fighting behavior unknown; (21) egg deposition site and parental care unknown; (22) tadpole unknown; (23) minute body size; SVL in male holotype 19.9 mm; females unknown.

**Color in life:** Green dorsum with black ocelli with yellow center [17].

**Color in ethanol** (Figure 163): Dorsal surfaces cream-colored with minute reddish–lavender melanophores appearing as punctuations and forming a reddish –lavender shadow on sides of body. Dark reddish–lavender ocelli on head and body, the center of each encircles a cream-colored wart. Venter cream. Parietal peritoneum with iridophores; white pericardium; all other peritonea clear [17].

**Biology and ecology:** Almost no information is available for *Nymphargus laurae*. The type locality is within the Foothill Evergreen forests. Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 164): *Nymphargus laurae* only is known from its type locality, near the town of Loreto, lower slopes of the Sumaco Volcano (ca. 77◦20- S, 00◦40- W; ca. 500 m), Ecuador [17], within the Amazonian Tropical Rainforest ecoregion.

**Figure 164.** Distribution of *Nymphargus laurae* in Ecuador (yellow dot).

**Conservation status:** *Nymphargus laurae* is classified as *Critically Endangered* by the IUCN [265]; however, given that its type locality has a degree of uncertainty (collected in 1955 from the general area of "Loreto, Upper Rio Napo") and that the species is known from a single specimen, we consider that the category of *Data Deficient* is more appropriate.

**Evolutionary relationships:** It is likely that *Nymphargus laurae* is a close relative of *N*. *cochranae* and *N*. *lindae* sp. nov., given their similar morphologies and color patterns.

**Taxonomic Remarks:** The description of *Nymphargus laurae* was based on a single specimen and, therefore, intraspecific variation is unknown. Given that this species shares a number of traits (general morphology, webbing, color pattern) with *N*. *cochranae*, a relatively abundant species on the lower slopes of Volcán Sumaco (nearby the type locality of *N*. *laurae*), we are presented with two possibilities: (i) The holotype of *N*. *laurae* represents an anomalous individual of *N*. *cochranae*, or (ii) *N*. *laurae* is a valid species, that might occur in sympatry with *N*. *cochranae*. Also, *N*. *laurae* is very similar to *N*. *lindae* sp. nov.*;* although, the two species are geographically distant (*N*. *lindae* sp. nov. is only known from Cordillera del Cóndor, whereas *N*. *laurae* is known from the lower slopes of the Volcán Sumaco); the two taxa differ in body size, vomerine teeth, and papillae on toes (see Diagnosis). As mentioned above, at the moment, it is impossible to determine the extent of intraspecific variation in *N*. *laurae*.

**Specimens examined:** *Nymphargus laurae:* Ecuador: *Provincia de Orellana:* Loreto region, near the town of Loreto, lower slopes of Volcán Sumaco (ca. 77◦20- S, 00◦40- W, ca. 500 m), on the Cordillera Oriental, USNM 288453.

*Nymphargus lindae* new species Guayasamin (Figures 165 and 166).
