http://zoobank.org/urn:lsid:zoobank.org:act:A8E08511-149C-4DA3-8F74-57C04C7CC302

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

**Etymology:** The specific epithet honors Linda Trueb, one of the most influential amphibian systematist of our days. Linda, as the curator of the herpetological collection of the University of Kansas, has led one of the most prolific and solid research group on amphibian biology, mentoring numerous students (including JMG). Her work on the evolution of skeletal diversity, ontogeny, and scientific illustration is outstanding.

**Holotype:** QCAZ 41572, adult male, collected from Miazi Alto (4.25044◦ S, 78.61356◦ W; 1200 m), Cordillera del Cóndor, Provincia de Zamora Chinchipe, Ecuador, by Juan M. Guayasamin and Elicio Tapia on 8 April 2009.

**Paratopotypes:** QCAZ 41562–71, 41573–74, 41587, 41594, 41597, 41599, 41644–47, 41654–56, 41658, 42446, adult males and females collected by Juan M. Guayasamin, Elicio Tapia, Silvia Aldás, and Holger Braun on 8–12 April 2009.

**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 an absence or reduced webbing between Fingers III and IV; additionally, males lack humeral spines (except *N*. *grandisonae*). *Nymphargus lindae* sp. nov. has the aforementioned traits. Molecular analyses of mitochondrial genes (12S, 16S, ND1) unambiguously place the new species in the clade *Nymphargus* (Figure 16).

**Identification:** *Nymphargus lindae* sp. nov. can be distinguished from most other glassfrogs by having, in life, a green dorsum with dark lavender to black ocelli enclosing yellow to orange spots (Figure 165), and lacking webbing between fingers. *Nymphargus lindae* sp. nov. is most similar to other glassfrogs with dorsal ocellated patterns (*N*. *anomalus*, *N*. *cochranae*, *N*. *ignotus*, *N*. *laurae*, *N*. *ocellatus*). *Nymphargus anomalus* differs by having, in life, a pale brown dorsum and scattered black and lavender flecks between ocelli (green dorsum lacking flecks in *N*. *lindae* sp. nov.); *N*. *ignotus* has a brown dorsum and occurs on the Pacific slopes of the Andes, whereas *N*. *lindae* sp. nov. has a green dorsum and is restricted to the Amazonian slopes of the Cordillera del Cóndor; *N*. *ocellatus* has a dorsal color pattern of large dark rings that, in life, have greenish–white centers (smaller ocelli with yellow to orange centers in *N*. *lindae* sp. nov.); *N*. *laurae*, known from a single adult male, is smaller (SVL = 19.7 mm in *N*. *laurae;* male SVL = 23.0–26.3 mm in *N*. *lindae* sp. nov.), has pointed papillae in Toes I–IV (absent in *N*. *lindae* sp. nov.), and no teeth on the vomers (present in *N*. *lindae* sp. nov.*;* see Taxonomic Remarks). *Nymphargus lindae* sp. nov. differs from its sister species, *N*. *cochranae*, by the relative size of ocelli (minute *in N*. *cochranae* and conspicuously larger in *N*. *lindae* sp. nov.) and female body size (*N*. *cochranae*, SVL = 27.8–30.3 mm; *N*. *lindae* sp. nov., SVL 27.2–27.8 mm).

**Figure 165.** *Nymphargus lindae* sp. nov. in life. (**Left**): Adult female, paratype, QCAZ 41597. (**Right**): Male, also part of the type series. Photos taken at the type locality by Juan M. Guayasamin.

**Diagnosis:** (1) Dentigerous process of the vomer with two to four teeth; (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%–26% 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 about anterior 60% of venter (condition P3); 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 (Figure 165); webbing formula III (2<sup>4</sup>/5–3−)—(2<sup>3</sup>/4–3−) IV; (10) feet about two-thirds webbed (Figure 165); webbing formula: I <sup>2</sup>−—(2<sup>1</sup>/4–21/3)II (1<sup>1</sup>/4–11/3)—(21/4–21/3) III (1<sup>1</sup>/4–11/3)—(22/3–3−) IV (2–2<sup>+</sup>)—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 or slightly shorter (Finger I length 94%–100% of Finger II); (14) disc of Finger III width about 46%–50% of eye diameter; (15) in life, dorsum green with dark lavender to black ocelli enclosing yellow to orange spots (Figure 165); bones green; (16) in preservative, dorsum lavender with black ocelli with white centers; (17) iris white with slight pale yellow hue and thin black reticulation; (18) melanophores usually lacking from dorsal surfaces of fingers and toes, except for few on Toe V; (19) males call from the upper surfaces of leaves; call unknown; (20) fighting behavior unknown; (21) egg deposition site unknown; parental care unknown; (22) tadpoles unknown; (23) small to medium body size; in males, SVL 23.0–26.3 mm ( X = 25.1 ± 0.848, *n* = 12); in females, SVL 27.2–27.8 mm (*n* = 2).

**Description of holotype:** Adult male, SVL 25.8 mm. Head slightly wider than long (head length 95% 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 an angle ~50◦; transverse diameter of disc of Finger III 49% eye diameter. Supratympanic fold low, obscuring upper portion of tympanic annulus; tympanum oriented mostly vertically, but with slight posterolateral inclination; tympanic membrane translucent, pigmented as surrounding skin. Dentigerous process of vomer low, situated transversely between choanae, with three teeth; choanae large, longitudinally rectangular; tongue ovoid, with ventral posterior fourth not attached to floor of mouth and posterior margin notched; vocal slits extending posterolaterally from about the lateral margin of tongue (at about half the length of tongue) to angle of jaws. Humeral spine absent. Low ulnar folds evident on external and internal ventrolateral margins of arm; relative lengths of fingers: III > IV > II > I; webbing absent between Fingers I–III, basal between Fingers III and IV, webbing formula III 3−–23/<sup>4</sup> IV; discs expanded, nearly elliptical; disc pads nearly triangular shaped; subarticular tubercles small, round, simple; low supernumerary tubercles; 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 59% SVL; low inner tarsal fold evident; outer tarsal fold absent; foot two-thirds webbed; webbing formula of foot: I 2−—21/<sup>3</sup> II 11/4—21/<sup>3</sup> III 11/3—22/<sup>3</sup> IV 3−—2− V; discs on toes round to elliptical, lacking papillae; disc on Toe IV narrower that disc on Finger III; disc pads triangular; inner metatarsal tubercle large, ovoid; outer metatarsal tubercle not evident; subarticular tubercles small, round; supernumerary tubercles low. 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. Ventral surface of thighs with pair of enlarged tubercles.

**Coloration of the holotype in life** (Figure 165): Green dorsum with dark lavender to black ocelli enclosing yellow to orange spots. Upper lip white. Anteriorly, about 60% of ventral parietal peritoneum white, posterior portion translucent. Bones green. Iris white with slight pale-yellow hue and thin black reticulation.

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

**Measurements of holotype (mm):** *Nymphargus lindae*, adult male, QCAZ 41572: SVL = 25.8, head length = 7.9, head width = 8.3, eye–nostril = 1.9, nostril–snout = 0.8, IOD = 2.4, upper eyelid width = 2.5, eye diameter = 3.5, tympanum diameter = 0.85, tibia = 15.1, foot length = 11.9, radio–ulna length = 5.5, hand length = 8.1, Finger I length = 5.2, Finger II length = 5.4, disc of Finger III width = 1.7.

**Biology and ecology:** *Nymphargus lindae* sp. nov. is the most abundant species in streams at Miazi Alto, Cordillera del Cóndor. Males were observed calling on leaves and branches about 30–400 cm above a stream. An amplectant pair and two additional males were observed fighting on a branch; males that were not in amplexus continuously approached the amplectant male and tried to push him away with kicks. Parental care is unknown.

**Call:** Not described.

**Tadpole:** Not described.

**Distribution** (Figure 166): *Nymphargus lindae* sp. nov. is only known from the type locality, Miazi Alto (4.25044◦ S, 78.61356◦ W; 1200 m), at Cordillera del Cóndor, Provincia de Zamora-Chinchipe, Ecuador. It inhabits the Eastern Foothill Forest ecoregion.

**Figure 166.** Distribution of *Nymphargus lindae* sp. nov. in Ecuador (yellow dot).

**Conservation status:** We sugges<sup>t</sup> placing *Nymphargus lindae* sp. nov. in the *Endangered* category, following IUCN criteria B2a, B2(iii). The main threats for the species in Cordillera del Cóndor are habitat destruction and contamination associated with mining activities.

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

**Taxonomic Remarks:** *Nymphargus lindae* sp. nov. is similar to *N*. *laurae*. Although the two species are geographically distant (*N*. *lindae* sp. nov. is endemic to the isolated Cordillera del Cóndor, whereas *N*. *laurae* is endemic to the lower slopes of Volcán Sumaco, nearby the town of Loreto), differences between these taxa are limited (body size, vomerine teeth, papillae on toes; see Diagnosis). Since *N*. *laurae* was described based on a single specimen, it is impossible to determine its intraspecific variation. Additional samples from Volcán Sumaco that correspond to the description of *N*. *laurae* need to be examined to support or refute the validity of the specific status of the two species.

*Nymphargus mariae* (Duellman and Toft, 1979 [266]; Figures 167–169).

*Centrolenella mariae* Duellman and Toft, 1979 [266]. Holotype: KU 174713.

Type locality: "Serranía de Sira, ± 1550 m, Departamento Huánuco, Perú".

*Cochranella mariae*—Ruiz-Carranza and Lynch, 1991 [6]; Cisneros-Heredia and McDiarmid, 2006 [267].

*Centrolene mariae*—Duellman and Schulte, 1993 [174].

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

*Centrolenella puyoensis* Flores and McDiarmid, 1989 [268]. Holotype: MCZ 91187. Type locality: "1.0 km W Puyo, Provincia de Pastaza, Ecuador, between 1000–1050 m elevation".

Synonymy by Cisneros-Heredia & Guayasamin, 2014 [175].

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

*Centrolene puyoense*—Stuart, Hoffmann, Chanson, Cox, Berridge, Ramani, and Young, 2008 [269]. *Nymphargus puyoensis—*Guayasamin, Castroviejo-Fisher, Trueb, Ayarzagüena, Rada, and Vilà, 2009[1].

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

**Etymology:** The specific epithet is a patronym for María Koepcke, who devoted her life to biological exploration in Peru [266].

**Identification:** *Nymphargus mariae* can be distinguished from most glassfrogs by having, in life, a green dorsum with relatively large yellow–green spots, a white parietal peritoneum covering the anterior half of the venter, and by lacking humeral spines and webbing between fingers (Figure 167). *Rulyrana flavopunctata*, *T*. *midas*, *N*. *cariticommatus*, *N*. *humboldti* sp. nov., and *N*. *siren* have a similar color pattern, but the yellow spots are conspicuously larger in *N*. *mariae*. Two species, *Sachatamia albomaculata* and *"Centrolene" medemi*, also have large yellow spots on the dorsum, but *S*. *albomaculata* has webbing between Fingers III and IV and inhabits the Pacific lowlands (*N*. *mariae* lacks hand webbing and is found on the Amazonian slopes of the Andes and Amazonian lowlands). Also, *"C*.*" medemi* is larger than *N*. *mariae* (*"C*.*" medemi:* SVL 25.5–30.8 mm in adult males, 34.7–44.3 mm in adult females; *N*. *mariae:* SVL 23.4–30.2 mm in males, SVL 25.1–32.8 mm in females), has more webbing between Fingers III and IV, and males have humeral spines.

**Figure 167.** *Nymphargus mariae* in life. Adult female, QCAZ 37923, from a stream tributary of Río Lliquino, Pastaza province, Ecuador. Photos by Martín Bustamante.

**Diagnosis:** (1) Vomerine teeth present, each vomer with three to four teeth; (2) snout truncated in dorsal aspect, round in lateral profile (Figure 168); (3) tympanum oriented dorsolaterally, with slight posterior inclination, its diameter about 32%–35% of eye diameter; tympanic annulus visible, low supratympanic fold evident, tympanic membrane partially pigmented and clearly differentiated

from surrounding skin; (4) dorsum shagreen with spiculated flat warts corresponding to light spots; (5) venter areolate; pair of enlarged subcloacal warts (Figure 6); (6) white lining on the anterior 40% of the ventral parietal peritoneum (condition P2); translucent peritonea covering intestines, stomach, gall bladder, and urinary bladder (condition V1); (7) liver tetralobed, covered by transparent peritoneum (condition H0); (8) humeral spines absent; (9) webbing absent between Fingers I, II, and III, and reduced or absent between outer fingers, III (3−–3)—(2<sup>1</sup>/3–22/3) IV (Figure 168); (10) webbing formula on foot: I (2–2−)—(2<sup>1</sup>/3–21/2) II 11/2—2−22/<sup>3</sup> III (1<sup>1</sup>/3–12/3)—22/<sup>3</sup> IV (2<sup>+</sup>–3−)—(1<sup>2</sup>/3–2−) V; (11) ulnar fold low; inner and outer tarsal folds low and thin; (12) concealed prepollex; nuptial pad Type I; (13) Finger I about same length as Finger II; (14) disc of Finger III narrow, about 34%–38% of eye diameter; (15) in life, dorsum green to dark green with minute and relatively large yellow–green spots; bones green; (16) in preservative, dorsum dark lavender with minute and large cream spots; (17) iris greyish cream with transverse brown bar and fine dark grey reticulations; pale yellow circumpupilar ring; (18) melanophores absent from Fingers I and II, and Toes I, II, and III; few present on Finger III; numerous on Finger IV and Toes IV and V; (19) males call from the upper side of leaves; call consists of a single note with a duration of 0.015–0.018 s; time between calls is 1.733–1.940 s; dominant frequency is at 3234–4299 Hz; (20) fighting behavior unknown; (21) egg deposition site and parental care unknown; (22) tadpoles unknown; (23) medium body size; in males, SVL 22.4–31.7 mm ( X = 26.3 ± 2.866, *n* = 7); in females, SVL 25.1–30.1 mm ( X = 28.3 ± 2.347, *n* = 4).

**Figure 168.** *Nymphargus mariae*, holotype, female, KU 174713. ( **A**) Head in lateral view. (**B**) Hand in ventral view. Illustrations by Juan M. Guayasamin.

**Color in life** (Figure 167): Dorsal surfaces green to dark green, with minute and large yellow spots. Anterior half of venter white, posterior part translucent. Iris greyish cream with transverse brown bar and fine dark grey reticulations; pale yellow circumpupilar ring.

**Color in ethanol:** Dorsum lavender to dark lavender, with minute and large cream spots; iridophores on anterior 40% of the ventral parietal peritoneum; white pericardium; clear hepatic peritoneum; translucent peritoneum covering intestines, stomach, gall bladder, and urinary bladder.

**Biology and ecology:** Nymphargus mariae is active during the night. Species found at the type locality of N. puyoensis included Boana cinerascens, Pristimantis conspicillatus, P. diadematus, P. lacrimosus, P. lathanites, P. martiae, and P. quaquaversus [268]. Parental care is unknown.

**Call:** Males call from the upper surfaces of leaves. The following call description is based on a recording of one male of *Nymphargus mariae* made by Diego Paucar on 22 February 2008 at stream tributary of Río Lliquino (1.72553 S, 78.98058 W; 400 m), Provincia de Pastaza, Ecuador. The call consists of a single non-frequency modulated note. Each call has a duration of 0.015–0.018 s (*n* = 3); time between calls is 1.733–1.940 s. Dominant frequency is at 3234–4299 Hz; a first harmonic is visible at 7178–7928 Hz, and a second harmonic is visible at 10767–11714 Hz.
