Gastrotheca elicioi, Carvajal-Endara & Coloma & Morales-Mite & Guayasamin & Székely & Duellman, 2019

Carvajal-Endara, Sofía, Coloma, Luis A., Morales-Mite, Manuel A., Guayasamin, Juan M., Székely, Paul & Duellman, William E., 2019, Phylogenetic systematics, ecology, and conservation of marsupial frogs (Anura: Hemiphractidae) from the Andes of southern Ecuador, with descriptions of four new biphasic species, Zootaxa 4562 (1), pp. 1-102 : 34-48

Gastrotheca elicioi sp. nov.

urn:lsid:zoobank.org:act:3D26B088-B527-4362-94E8-647FF26C8419

Holotype. CJ 1402 ( Fig. 12 ), an adult male (collected as tadpole in the field and captive raised), from Cajanuma , entrance to Parque Nacional Podocarpus in the Loja-Vilcabamba road, 2456 m (04° 05' 25.51" S, 79° 12' 09.97" W), Loja Province, Ecuador, one of a series obtained on 14 June 2011 by Elicio E. Tapia, Sofía Carvajal-Endara, and Henry Grefa.

Paratypes. ( Total 18: 11 males, 3 females, 4 juveniles). Ecuador: Loja : CJ 413–4 (juvenile, male), 1398–401 (four males), 1941 (male) collected with the holotype ; KU 142603–5 (female, subadults), from 5.5 km W Loja , 2330 m (04° 00' 48.99" S, 79° 13' 50.99" W), on 17 July 1971 by William E. Duellman and Linda Trueb ; KU 142607–8 (subadult, female), 148549–51 (males, female), from 5.5 Km W Loja , 2330 m (04° 00' 48.99" S, 79° 13' 50.99" W), on 23 July 1971 by William E. Duellman and Linda Trueb; KU 202688 (male), from 5.2 km W Loja, 2310 m (03° 58' 58.73" S, 79° 16' 02.21" W), on 10 March 1984 by William E. Duellman ; KU 217511–2 (males), from 6.8 km E Loja ca. Loja-Zamora line (03° 59' 19" S, 79° 09' 21.99" W), on 9 January 1990 by David Kizirian, John J. Wiens, and Luis A. Coloma .

Referred specimens. Ecuador: Loja: QCAZ 22370 (male), from Zamora Huayco , Loja, 3018 m (04°0 5' 49.98" S, 79° 10' 02.5" W), on 2 December 2002 by Diego Almeida-Reinoso; QCAZ 46319–20 (males), from Zamora Huayco , Loja, 3018 m (04° 05' 49.98" S, 79° 10' 02.5" W), on 5 December 2009 by Diego Almeida- Reinoso and Samael D. Padilla; CJ 1841–902 (62 juveniles), from Loja, Barrio La Palmera, Parroquia Sucre , on 22–25 April 2013 by Elicio E. Tapia .

Diagnosis. Included in the genus Gastrotheca by having a closed brood pouch on dorsum of female. A moderately large species (up to 67.5 mm SVL in female, 46.4–67.8 mm SVL in males, n = 13) with tibia length 42¯50% SVL, slightly longer than foot; (2) interorbital distance larger than the width of upper eyelid; (3) skin on dorsum finely granular, not co-ossified with skull, lacking transverse ridges; (4) supraciliary processes absent; (5) heel lacking calcar or tubercle; (6) tympanic annulus smooth to slightly granular; (7) Fingers I and II about equal in length, width of discs about twice width of the digits proximal to discs; (8) fingers unwebbed; (9) webbing between external toes extending nearly to antepenultimate subarticular tubercle on Toe IV and proximal to penultimate subarticular tubercle on Toe V; (10) in life, dorsum green, tan, brown, or reddish-brown with or without dark paravertebral marks; (11) head markings consisting of a pale labial stripe on the posterior margin of the lip, a black canthal stripe and a dark triangular interorbital mark in some individuals; (12) a fragmented, narrow, pale dorsolateral stripe present; (13) flanks bronze-brown, tan, or green with or without small pale spots; groin and anterior and posterior surfaces of thighs lightly mottled with tinges of green or blue; (14) venter cream with small brown flecks or spots; (15) brood pouch single, dorsal.

In comparison with similar species Gastrotheca elicioi is most like G. lojana , G. cuencana , G. litonedis , G. turnerorum , and G. pseustes in Ecuador and G. monticola in Peru. Gastrotheca elicioi differs from G. lojana , G. cuencana , and G. litonedis in color pattern as follows (compare these species in Fig. 10 ). Gastrotheca elicioi and G. litonedis have distinct dark canthal stripes, which are absent in G. lojana and inconspicuous or absent in G. cuencana . The groin and anterior and posterior surfaces of the thighs are slightly mottled in G. elicioi , whereas in G. lojana they are heavily mottled; these surface are translucent cream without marks in G. cuencana and usually brown in G. litonedis . If any marks are present on the dorsum in G. elicioi , they usually consist of a triangular interorbital mark, which is connected or not with two narrow and curved paravertebral marks. In contrast, in G. lojana the interorbital mark is a transverse bar usually connected with two broad paravertebral marks; in G. cuencana and G. litonedis an interorbital mark is absent. Gastrotheca lojana and G. cuencana have a conspicuous, elevated row of dorsolateral warts, whereas they are barely raised in G. elicioi and G. litonedis . Additionally, in G. elicioi the dark bars on the limbs, when present, are shorter, thinner and less defined than in G. lojana , whereas in G. cuencana and in some G. litonedis the bars are replaced by irregular blotches. Gastrotheca elicioi has a cream venter with dark flecks or marks, whereas the venter is uniform creamy white in G. cuencana and pale brownish gray in G. litonedis . Gastrotheca cuencana also differs from G. elicioi by having cream dorsal surfaces of the fingers, whereas they are brown, green, or tan in G. elicoi . Gastrotheca turnerorum and G. pseustes differ from G. elicioi (compare these species in Fig. 10 ) by having the skin on dorsum areolate and weakly areolate respectively, whereas in G. elicioi the skin on dorsum is finely granular. Also, they differ by lacking an interorbital mark, and bars on limbs, which can be present G. elicioi . Gastrotheca pseustes is sympatric with G. elicioi , from which it differs notably by having smaller digital discs (larger in G. elicioi ) and lacking a pale dorsolateral and supracloacal stripes, which are fragmented in G. elicioi . Gastrotheca monticola is the sister species of G. elicioi and their minimum genetic divergence is 3.0% (in a DNA dataset of 438 bp, 16S gene). Gastrotheca monticola differs from G. elicioi by having the axilla and groin, concealed surfaces of the thighs, and the dorsal surfaces of the feet green with well-defined black spots on a green or tan background, whereas in G. elicioi the groin and anterior and posterior surfaces of thighs are lightly mottled with tinges of green or blue. Finally, the advertisement call of G. elicioi is unique amongst the calls of Gastrotheca in southern Ecuador, in that the long, pulsed notes are produced after the short ones, whereas the other species tend to produce the long, pulsated notes before the short ones.

Description of the holotype. An adult male ( Fig. 12 ) collected as a tadpole and raised in the laboratory; body moderately robust; SVL 46.4 mm; head wider that long; snout rounded in dorsal view, bluntly rounded in profile; canthus rostralis round in section; loreal region concave, lips rounded; top of head flat; interorbital distance 142% of width of upper eyelid; internarial area elevated; nostrils not protuberant, directed anterolaterally, posterior to level of anterior margin of lower jaw; diameter of eye greater than its distance from nostril; tympanum round, separated from the eye in a distance approximately one and a half the diameter of tympanum; tympanic annulus and membrane smooth; supratympanic fold moderately weak, extending from corner of eye to tympanum and on to insertion of forelimb. Dentigerous vomerine processes narrowly separated medially, each bearing three teeth.

Arm robust; ulnar tubercles absent; hand and fingers moderately large (TFL 31% of SVL), unwebbed, with distinct narrow lateral fringes; discs much wider than digits, slightly truncated, width of disc on Finger III greater that diameter of tympanum; relative lengths of fingers I=II<IV<III; subarticular tubercles prominent, round, none bifid; supernumerary tubercles, small, numerous, rounded; palmar tubercle bifid, prepollical tubercle elliptical. Hind limb robust; tibia length 43% SVL; foot length 43% SVL; calcar and tarsal tubercles absent, inner tarsal fold on distal half of tarsus; outer metatarsal tubercle absent; inner metatarsal tubercle large, elliptical; toes moderately long; relative length of toes I<II<III<V<IV; basal webbing between Toes I and II; webbing formula for other toes II1— 2III 1— 2IV 2— 1V; subarticlular tubercles moderately small, rounded; supernumerary tubercles, numerous, rounded, present on proximal segments of digits. Nuptial pad absent.

Skin on dorsum finely granular; skin on flanks coarsely granular; skin on throat, venter surfaces of thighs, and arms granular; skin on belly areolate; skin on venter surfaces of shanks smooth; numerous small tubercles lateral to cloacal opening. Vocal sac single, median, subgular. Vocal slits present at posterior lingual margins of mandibles. Tongue broad, suboval, notched posteriorly, fully attached to mouth floor.

Coloration in life (See also under Comments). The dorsum is uniform brown becoming yellowish brown on the flanks. The dorsal surfaces of limbs have an olive-green tinge. A bronze labial stripe is present at the margin of the lip and continues to the insertion of the forelimb. A bronze inconspicuous and highly fragmented dorsolateral stripe is present; a black canthal stripe is present. The flanks are bronze-brown; the groin and anterior and posterior surfaces of the thighs are pale green with white and black spots. The venter is brownish gray with black and white marks; the gular region is gray dark. The iris is copper colored with a few black reticulations.

Coloration in preservative. The dorsum is gray without markings. A narrow pale, fragmented dorsolateral stripe is present. The dorsal surfaces of the limbs are tan with irregular gray blotches. A short pale labial stripe extends from the margin of the lip to the level of the insertion of forelimb; a dark canthal stripe is present. The flanks are dark gray anteriorly and pale gray posteriorly; the groin and anterior and posterior surfaces of the thighs are pale gray with black marks. The venter is cream with dark gray spots; the gular region is gray.

Measurements (in mm). SVL: 46.4, TIBL: 19.8, FL: 20.0, HL: 14.1, HW: 16.2, IOD: 4.9, EW: 4.6 IND: 3.3, ED: 5.6, EN: 4.1, TD: 2.2, FFL: 7.9, TFL: 14.6, TFD: 2.6.

Variation. Morphometric variation of one female and 13 males is summarized in Table 2. The female is larger than most males (67.5 mm; 56.0± 8.1 mm). The skin on the dorsum is finely granular. The tympanic annulus usually is smooth but is slightly granular in some individuals. Each dentigerous vomerine process has 6–8 teeth (6.3±1.5, n = 3).

Color variation in preservative. Preserved specimens have a bluish gray, pale gray, or brownish gray dorsum. Interorbital and paravertebral marks usually are present but the paravertebral marks may be absent. The dark gray interorbital mark, when present, is triangular in most specimens, but it is divided into two blotches in some specimens. A dark canthal stripe is present in all specimens. A short, pale labial stripe is inconspicuous in most specimens. The flanks are gray, darkest anteriorly, with or without white and dark flecks. The groin and anterior surfaces of thighs are pale gray with slight mottling in some; the posterior surfaces of the thighs are pale gray with black flecks and marks usually only distally. In some specimens, the dorsal surfaces of arms and limbs present short dark gray bars. The venter is white with evenly, or densely distributed dark marks.

Color variation in life. ( Figs. 10H, 10I , 13–14 ). In living individuals the dorsum is uniform green (CJ 1843), reddish-brown (CJ 1400), or tan (CJ 1842), with or without contrasting dark gray paravertebral marks. When present, these paravertebral marks usually are narrow, curved, and fragmented; these marks are adherent at the scapular level and may be connected with the interorbital mark. A short cream labial stripe is present on the margin of the lip; the stripe continues posteriorly to the insertion of the forelimb. A dark brown canthal stripe is always present, but it is inconspicuous in some individuals with a dark ground color. The tympanum is brown, tan, or olive green. The iris is usually copper almost without black reticulations. A bronze highly fragmented dorsolateral stripe is present in most individuals. The flanks are brown, bronze-brown, or green. The groin has bluish coloration with slight mottling. The posterior surfaces of the thighs usually are pale green with black flecks and some specimens show black fringes. When present, supracloacal and heel stripes are cream and fragmented. The ventral surfaces usually are cream with dark marks, but some specimens lack dark marks. The gular surface varies from brownish gray to white with black spots. Duellman (1974) provided detailed descriptions of coloration in life of four adult males (KU 148549–51 and 142603, under the name Gastrotheca lojana ).

Tadpoles. Tadpoles belong to Type IV tadpole of Orton (1953), and the exotroph, benthic ecomorphological guild. All measurements are expressed in millimeters. The following description is based on a specimen in Stage 36 (CJ 4311), from a series of 50 tadpoles (CJ 4311) obtained from a pond at Puntzará Alto, near Loja city, 2311 m, Province of Loja, Ecuador, by Luis A. Coloma, Manuel A. Morales-Mite, and Elicio E. Tapia on 28 January 2016. Total length 51.4; body length 19.4 (38% of total length). Body ovoid in dorsal and lateral views, slightly depressed; throat slightly concave in lateral profile, sloping from tip of snout to belly; body width at the level of spiracle 11.9, and height at same position 9.9; head width at level of eyes 11.1. Lateral line system present but barely visible, supraorbital and not evident at level of snout, infraorbital line present at level of the eye, touching the inferior portion of the orbit, and making contact with supraorbital line immediately behind the eye. Inferior oral line visible at the eye level, where it contacts angular line, which descends from eye level. Supraorbital line represented for scattered stiches, which does not make contact with the orbit. Posorbital line forming a circle of stitches, just behind the eye. Anterior oral line and loreal lines not visible; dorsal body and middle bodylines not visible.

Nostril medium sized (in proportion to body length), ovoid, protruding, having a fleshy annulus, its opening directed anterolaterally. Snout–nostril distance 3.5; internarial distance 2.9. Eye directed dorsally; eye length 2, eye width 1.8; interorbital distance 5. Spiracle sinistral, located at midbody level, spiracular opening oriented posteriorly; distance from tip of snout to spiracular opening 13.0; end of spiracular tube rounded, attached to body wall, inner wall of spiracular tube not evident; spiracle length 2.9, tube transverse width 2.9. Vent tube dextral, opening oriented posteriorly, tube length 3.4, vent tube transverse width 3. Tail length 32.4; caudal musculature slender, narrowing gradually until tail terminus; caudal muscle height 4.4, width 3.4; caudal fins well developed and proportional, arising abruptly near tail-body junction and forming a notorious hump, which makes and arc in the body plane. Dorsal fin height 4.18, ventral fin height 3.7; maximum height of tail 11.9; tail terminus rounded, caudal musculature not reaching fin terminus.

Oral disc small, ventral, located near tip of snout, not protruding laterally beyond body; transverse width 4.9. It is surrounded by an uniserial row of marginal papillae, interrupted medially on upper lip; lower lip papillae alternate in orientation, giving appearance of two rows. Upper lip with 23 papillae on right side and 20 papillae on left side; lower lip bearing 52 marginal papillae; upper jaw sheath medium-sized, forming a smooth arch and finely serrated, transverse width 2.6 (53% of oral disc width) height 0.4; lower jaw sheath V- shaped, open and finely serrated, width 4.1, height 1.3. Labial tooth row formula 2/3(1); tooth rows lengths: A1: 4.15, A2: 4.0, P1 right row 1.75, P1 left row 1.75, P1 gap 0.1, P2: 3.75, P3: 3.70. ( Fig. 6D ).

Color in life. Based on a specimen (CJ 4312a) in Stage 37 from a series (CJ 4312) obtained at Loja (neigborhood Puntzará Grande), Loja Province, Ecuador, by Luis A. Coloma, Manuel A. Morales-Mite, and Elicio E. Tapia on 28 January 2016 ( Fig. 5D ). In dorsal and ventral views, body dark brown. Snout and flanks dark brown; guts gray; red gills visible through the throat. Venter gray. Caudal musculature reddish brown with brown stippling, distal portion lacking pigments; dorsal and ventral fins gray with minute cream stippling. Vent tube translucent. Iris reddish-gold.

Variation. Variation of 28 meristic characters of tadpoles in Stages 31–40 (CJ 4306, 4310–13) are shown in Table 6. Total length varies between 15.8 (Stage 31) and 73.2 (Stage 39) and tail length proportion varies from 57% to 70% until Stage 38. Number of marginal papillae varies among specimens and Gosner stages; variation in number of ventral papillae at lower lip is high (43–64).

Based on specimens CJ 1950–2 from a series obtained at Loja (neigborhoods La Palmera and Puntzará Grande), Loja Province, Ecuador, by Elicio E. Tapia on 22–25 April 2013 ( Fig. 15 ). We documented changes in coloration during ontogenetic development of one, mostly brown individual (CJ 1950) ( Figs. 15 A–B). At Stage 40, the dorsum and flanks were cream and brown with a diffuse pattern of brown-gray paravertebral marks and a dark gray stripe bordering the canthus and body dorsolaterally, extending to about level of midbody, bordered dorsally by a diffuse cream area. The caudal musculature is pink proximally, with a gray suffusion distally. The caudal fins are pale gray. The iris is pale red. By Stage 46, the markings on the dorsum of body and limbs are diffuse gray and green on a dark brown background; the fingers and toes are yellowish cream dorsally. The flanks have a wide, brown-gray band. There are well-defined creamy white dorsolateral and labial stripes and supracloacal white marks. Color variation in six additional metamorphs (CJ 1951–2, CJ 4313) from Loja and Loja–Abra de Zamora, in Stage 46 is depicted in Figure 16 . They vary from plain dark brown to plain green, and have a complete supracloacal stripe.

Comparisons. Tadpoles of Gastrotheca elicioi may occur in sympatry with those of G. lojana , G. psychrophila , G. pseustes , and G. turnerorum in the Loja-Abra de Zamora region. Gastrotheca elicioi differs from all of them by having a dorsal gray-pigmented fin that abruptly arises from the body, whereas is nearly translucent and arises gradually in the other species (compare in Fig 5 ). The tadpole described as G. psychrophila by Duellman (2015) is G. elicioi . It has a similar dorsal fin and fit well our description of G. elicioi tadpoles. Nonetheless, until tadpoles of G. psychrophila are described and unequivocally assigned to its species, some doubts will remain.

Vocalization. Seven individuals of Gastrotheca elicioi were recorded from three locations in Loja Province (one individual from the old Loja-Catamayo road, three from Loja, Parque Universitario de Educación Ambiental y Recreación (PUEAR) and one from Loja, Quebrada El Salado) and two individuals in the facilities of Centro Jambatu in San Rafael, Quito, Ecuador (Appendix III), inside outdoor enclosures. Descriptive statistics of the acoustic variables are provided in Table 5. The advertisement call of G. elicioi is a complex call, composed of up to 6, short, single-pulsed notes, and followed by up to 7 longer pulsed notes ( Fig. 17 A–G). It can be characterized as a series of short notes, which become gradually longer, pulsated notes by the end of the call. The long notes have a mean duration of 0.188 s (SD = 0.150) and consists on average of 3.47 (SD = 1.695) distinct pulses, partly fused, without silent intervals (amplitude modulation close but less than 100%). The amplitude of the long note decreases gradually towards the end. The short notes have a mean duration of 0.027 s (SD = 0.014) and the inter-note interval is on average of 0.452 s (SD = 0.125). The mean dominant frequency of the call is 1249.4 Hz (SD = 169.862), with a mean 90% bandwidth of 807.1–1416.8 Hz. The fundamental frequency and harmonics are not clearly recognizable.

Comparisons. The advertisement call of Gastrotheca elicioi is unique amongst the calls of the Southern Ecuadorian species of Gastrotheca in that the long, pulsed notes are produced after the short ones, whereas the other species tend to produce the long, pulsed notes before the short ones. Also, the duration of the long notes of G. elicioi is significantly shorter and there are fewer pulses per long note, compared to all other species with complex calls from southern Ecuador ( Table 5).

Distribution and ecology. Gastrotheca elicioi is known only from the vicinity of the city of Loja, Loja Province, in the Loja Basin and slopes of the adjacent cordilleras ( Fig. 9 ). Its elevational range is 2026–3018 m in an area of extent of occurrence of about 102.7 km 2.

This nocturnal, semiarboreal species inhabits mostly disturbed areas and a few forests in the Evergreen Montane Forest from the south of the Cordillera Oriental of the Andes (Ministerio de Ambiente del Ecuador 2012), where the average annual rainfall is 764–1128 mm and the average annual temperature is 12.6–17.4 °C ( Fick & Hijmans 2017).

At the locality 5.5 km W of Loja, many Gastrotheca elicioi were found in a roadside agave fence-row, where they were sitting on the agave leaves (WED field notes, 17 July 1971). At the locality 5.2 km W of Loja, an individual of G. elicioi (KU 202688) was under an overhanging bank at the edge of small pond in a pasture by day (WED field notes, 10 March 1984). At Zamora Huayco , in the vicinity of Loja, two individuals were sitting on branches of shrubs at approximately 4–5 m above the ground; they were about 7 m from a river within moderately dense mountain forest (Diego Almeida-Reinoso field notes, 5 December 2009). Brooding females under captive conditions are depicted in Figures 10 H–I. The holotype was one of a series of 16 tadpoles that were at the edge of a pond. At the type locality, G. elicioi is syntopic with G. pseustes ; G. elicioi is also syntopic with G. lojana . On 5 December 2017 a pregnant female G. elicioi was caught (Quebrada Shucos, Loja, Loja Province) by Diego Armijos-Ojeda and brought to the laboratory. After 20 days, the female deposited 434 tadpoles, in Gosner Stage 34 (CJ 7874). To the best of our knowledge, this is the largest number of tadpoles deposited by a female Gastrotheca . The female (SVL = 71.7 mm) weighed 43.06 g when caught and 24.38 g after she deposited the tadpoles.

Conservation status. We suggest that Gastrotheca elicioi should be considered as Endangered according to criteria B1ab(i,ii,iii,iv) of the IUCN Red List. We suggest this conservation status because of its small known area of occurrence (102.7 km 2) that is extremely fragmented. Its habitats are in heavily human populated areas. Loja and environs have been deforested and modified for human activities ( Fig. 11 G–H). None of its populations is within the National System of Protected Areas (SNAP), but there is a population in the private Madrigal del Podocarpus Natural Reserve.

Etymology. The specific name elicioi is a patronym of Elicio E. Tapia, an Ecuadorian biologist and member of the Tapia family, whom have contributed substantially to the growth of biodiversity collections in Ecuador ( Pollet 2003, Páez-Vacas et al. 2010). We highlight Elicio´s intense field collecting efforts and enormous contribution to the museums that hold Ecuadorian amphibians, including many tissues, call recordings, ecological data, and specimens of Gastrotheca reported here.

Comments. In the past, Gastrotheca elicioi was confused with G. lojana and G. monticola . It belongs to the subgenus Duellmania and is reported as species B in Duellman (2015: Fig 11.1). Specimens in captivity were not as brightly colored as specimens from the wild; however, the color in life of the holotype does not show remarkable differences from wild-collected specimens.

Figures

FIGURE 5. Live tadpoles of: (A) Gastrotheca cuencana (CJ 1945, Stage 37), from Cuenca, Azuay Province, (B) Gastrotheca litonedis (CJ 6813, Stage 37), from Laguna de Busa, San Fernando, Azuay Province, (C) Gastrotheca lojana (CJ 1948, Stage 35), from Oña, Azuay Province, (D) Gastrotheca elicioi (CJ 4312a, Stage 37), from Loja, Loja Province, (E) Gastrotheca turnerorum (CJ 1957, Stage 36), from Laguna Negra de Jimbura, Parque Nacional Yacuri, Loja Province, and (F) Gastrotheca pseustes 1 (CJ 1949, Stage 37), from Chillacocha, El Oro Province. Scales = 3 mm (A–D, F) (E). Photos by LAC (A, C, E) and MAM (B, D).

FIGURE 6. Oral discs of: (A) Gastrotheca cuencana (CJ 1945, Stage 38), from Cuenca (south control), Azuay Province, (B) Gastrotheca litonedis (CJ 5292, Stage 35), from from Laguna de Busa, San Fernando, Azuay Province, (C) Gastrotheca lojana (CJ 4309, Stage 37), from El Tablón, Loja Province, (D) Gastrotheca elicioi (CJ 4311, Stage 36), from Barrio Puntzará Grande, Loja, Loja Province, (E) Gastrotheca turnerorum (CJ 1959, Stage 39), from Laguna Negra de Jimbura, Parque Nacional Yacuri, Loja Province, and (F) Gastrotheca pseustes (CJ 1949, Stage 37), from Chillacocha, El Oro Province. Not to scale. Photos by LAC.

FIGURE 9. Localities of known occurrence of Gastrotheca cuencana (green), G. litonedis (red), G. elicioi (black), G. psychrophila (orange), G. turnerorum (blue), G. lojana (white), and G. yacuri (yellow). See Appendix II for detailed locality data. Some nearby localities are represented by a single symbol. Inset map indicates location of southern Ecuador (box) in Ecuador.

FIGURE 10. Live adults and brooding females of (A) holotype of Gastrotheca litonedis (KU 202690, SVL = 62.4 mm), of (B– C) G. litonedis (D–G) G. pseustes, G is the holotype (KU 203443, SVL = 55.2 mm) from 7.1 km N San Lucas, 2940 m, (H–I) G. elicioi, (J) G. turnerorum, (K) G. cuencana, (L) G. lojana, (M) female G. psycrophila (KU 164233, SVL = 63.5 mm) from 15 km E of Loja, 2850 m, and (N–O) G. yacuri, N is the holotype (CJ 5822, SVL = 57.4 mm). Not to scale. Photos A, G, M by WED; B–F, H–L by LAC, and N–O by PS.

FIGURE 11. (A) Habitat of Gastrotheca elicioi, G. lojana, G. pseustes, G. psychrophila, and G. turnerorum at the Abra de Zamora, Loja Province, (B) habitat at the type locality of G. cuencana in Cuenca, Azuay Province, (C) habitat of G. turnerorum and G. yacuri at Lagunas Negras, Parque Nacional Yacuri, Loja Province, (D) habitat of G. litonedis, G. lojana, and G. pseustes at Laguna de Busa, Azuay Province, (E) habitat of Gastrotheca lojana at El Tablón, Loja Province, (F) habitat of G. pseustes and (G–H) habitats of G. elicioi at Loja and 15 Km N of Loja, Loja Province. Photos A–B, E, G, H by LAC, C by PS, D by MMM, and F by Jan Verkade.

FIGURE 12. Live holotype of Gastrotheca elicioi (CJ 1402, male, SVL = 46.4 cm): A) dorsal, ventral and lateral views, (B) ventral view of left hand, enlarged 1.8X of A, and (C) ventral view of left foot, enlarged 2.1X of A. Photos by LAC.

FIGURE 13. Live adults and subadults of Gastrotheca elicioi showing variation in dorsal and ventral color pattern: (A) CJ 1841, subadult female, SVL = 33.9 mm, (B) CJ 1399, male, SVL = 48.9 mm, (C) CJ 1843, subadult female, SVL = 40.1 mm, (D) CJ 1400, male, SVL = 52.0 mm, (E) CJ 1941, male, SVL = not taken, and (F) CJ 1842, subadult female, SVL = 33.7 mm. Not to scale. Photos by LAC.

FIGURE 14. Live adults of Gastrotheca elicioi showing variation in lateral, groin, thighs and eye color pattern: (A) CJ 1941, male, SVL = not taken, (B) CJ 1399, male, SVL = 48.9 mm, (C) CJ 1841, subadult female, SVL = 33.9 mm, and (D) CJ 1843, subadult female, SVL = 40.1 mm. Not to scale. Photos by LAC.

FIGURE 15. Ontogenetic changes in live Gastrotheca elicioi. Stages of Gosner: (A) Stage 40 (CJ 1950), (B) Stage 46 (CJ 1950). From Loja, Loja Province, Ecuador. Scale = 5 mm. Photos by LAC.

FIGURE 16. Metamorph variation in live Gastrotheca elicioi in Stage 46 of Gosner: (A) CJ 4313 from Loja-Abra de Zamora, and (B) CJ 1950 from Loja, Loja Province, Ecuador. Photos by LAC.

FIGURE 17. Advertisement calls of Gastrotheca elicioi (A–G) and G. lojana (H–N): (A) oscillogram of two consecutive calls with the first call composed of five short and three long notes and the next call composed of five short and four long notes, (B) oscillogram of a single short note, (C) spectrogram of a single short note, (D) power spectrum of a single short note, (E) oscillogram of a single long note, (F) spectrogram of a single long note, (G) power spectrum of a single long note, (H) oscillogram of three consecutive calls, with the first call composed of one long and one short notes and the next two calls composed of one long and two short notes, (I) oscillogram of a single long note, (J) spectrogram of a single long note, (K) power spectrum of a single long note, (L) oscillogram of a single short note, (M) spectrogram of a single short note, (N) power spectrum of a single short note. All spectrograms at Hanning window function, 512 bands resolution. See text for details.