Allobates sp.

Ferreira, Anthony S., Ferrão, Miquéias, Cunha-Machado, Antonio S., Magnusson, William E., Hanken, James & Lima, Albertina P., 2024, Phylogenetic position of the Amazonian nurse frog Allobates gasconi (Morales 2002) revealed by topotypical data, Organisms Diversity & Evolution (New York, N. Y.) 24 (1), pp. 99-118 : 105-113

Allobates sp. “Ucuyali” Réjaud et al. (2020)

Holotype. INPA-H 3082 ( Fig. 3 ), sub-adult male from Jainú West, west bank of the middle Juruá River , municipality of Itamarati, state of Amazonas, Brazil, collected by Claude Gascon on 17 October 1991.

Paratypes. Thirty –three individuals, all collected by Claude Gascon in the Juruá River , Brazil. West bank: Jainú West: six juveniles ( INPA-H 3150–51 , 3172 , 3406 , 3412 and 3415), 19–29 October 1991 ; two adult males ( INPA-H 3512–13 ), 03 November 1991 . East bank: Condor: one juvenile ( INPA-H 2649 ), 19 September 1991 . Jainú East: fifteen juveniles ( INPAH 3066 , 3068 , 3075 , 3078 , 3084–85 , 3087 , 3090 , 3094 , 3220 , 3401 , 3483–84 , 3494 and 3496), 17–29 October 1991 and 2 November 1991 ; three adult females ( INPA-H 3055 , 3079 and 3491), 16–17 October 1991 and 2 November 1991 ; three adult males ( INPA-H 3083 , 3093 and 3249), 17–23 October 1991 . Altamira: one adult male ( INPA-H 3541 ), 9 November 1991 . Nova Vida: one juvenile ( INPA-H 4726 ), 21 March 1992 .

Referred specimens. Thirty-one adults, all from the middle Juruá River , municipality of Itamarati, state of Amazonas, Brazil, collected by A.S. Ferreira, A.P. Lima and M. Ferrão. West bank: Jainú West : five females ( INPA-H 45074–76 and MPEG 44620–21 ) and twelve males ( INPA-H 45067–70 , 45071–73 , 45077–79 and MPEG 44619 , 44625 ), 22–24 January 2021. East bank: Jainú East : one female ( INPA – H 45083 and six males ( INPA – H 45080–82 , 45084 and MPEG 44622–23 ), 23 January 2021. Altamira : one female ( INPA – H 45085 ) and six males ( INPA – H 45086–87 , 45088 , 45089 and MPEG 44626–27 ), 25 and 26 January 2021 .

Reappraisal diagnosis. Allobates gasconi is a small nurse frog diagnosed by the following combination of characters: SVL 14.8–17.1 mm in males (n = 30) and 16.0–18.0 mm in females (n = 10); finger III strongly swollen in adult males; expanded disk on all fingers; one subarticular tubercle on finger IV; basal toe webbing between toes II – III and III – IV; lateral fringes present on pre- and postaxial edges of all fingers and toes; dark brown hourglass-shaped mark on dorsum; males with a gray to dark gray throat; advertisement call comprises 2–4 notes emitted repeatedly, with a dominant frequency of 5,081 –6,761 Hz; larvae with labial keratodont row formula 2(2)/2(1).

Allobates gasconi is easily distinguished from A. amissibilis , A. bacurau , A. caldwellae , A. chalcopis , A. conspicuus , A. femoralis , A. fuscellus , A. granti , A. hodli , A. insperatus , A. juami , A. marchesianus , A. nidicola , A. paleovarzensis , A. sieggreenae , A. subfolionidificans , A. sumtuosus , A. tinae , A. trilineatus , A. vanzolinius , and A. velocicantus by having a distinct dark brown hourglass-shaped mark on the dorsum (absent in the other species). Additionally, A. gasconi differs from A. caldwellae , A. conspicuous , A. chalcopis , A. femoralis , A. granti , A. hodli , A. insperatus , A. juami , A. marchesianus , A. nidicola , and A. tinae by having males with finger III strongly swollen (weakly or not swollen in the other species); from A. paleovarzensis and A. vanzolinius by its maximum SVL 17.1 mm in adult males (minimum SVL 18.3 mm in A. paleovarzensis , 22.9 mm in A. vanzolinius ); from A. sieggreenae , A. subfolionidificans and A. velocicantus by having males with a gray to dark gray throat in life (white to translucent in A. sieggreenae , opaque white in A. subfolionidificans , white centrally and yellow marginally in A. velocicantus ); from A. bacurau and A. sumtuosus by having dark brown transverse bars on the dorsal surface of the thighs (absent in the other species); from A. trilineatus by the absence of ventrolateral stripe (present in A. trilineatus ); from A. amissibilis by having a conspicuous pale dorsolateral stripe present (absent in A. amissibilis ); and from A. fuscellus by having lateral fringes on toes (absent in A. fuscellus ).

The dark brown hourglass-shaped mark on the dorsum of A. gasconi is similar to that of A. algorei , A. flaviventris , A. humilis , A. kamilae , A. melanolaemus , A. ornatus , A. pacaas and A. pittieri . However, A. gasconi differs from A. algorei , A. flaviventris , A. humilis , A. melanolaemus and A. pacaas by having finger III greatly swollen in adult males (weakly swollen in all the other species); from A. flaviventris , A. humilis and A. melanolaemus by the maximum SVL 18.0 mm in adults (minimum SVL 19.3 mm in A. flaviventris , 21.8 mm in A. humilis , and 21.1 mm in A. melanolaemus ); from A. kamilae and A. ornatus by having males with a gray to dark gray throat in life (bright yellow in A. kamilae , pale lemon yellow with scattered melanophores in A. ornatus ); from A. algorei by having basal toe webbing (absent); from A. ornatus by having lateral fringes on toes and oblique lateral stripe (absent); from A. pittieri by having expanded disk on finger III (not expanded) and absence of cloacal sheath (present); and from A. pacaas by having one subarticular tubercle on finger IV (two tubercles).

The advertisement call of Allobates gasconi is easily distinguished from those of A. algorei , A. amissibilis , A. bacurau , A. caldwellae , A. femoralis , A. humilis , A. hodli , A. juami , A. kamilae , A. marchesianus , A. melanolaemus , A. nidicola , A. paleovarzensis , A. sieggreenae , A. subfolionidificans , A. sumtuosus , and A. tinae by having 2–4 notes (calls composed of one note in the other species). Moreover, calls of A. gasconi are emitted with a dominant frequency of 5,081 –6,761 Hz, which differs from A. femoralis , A. humilis , A. hodli , A. nidicola , and A. paleovarzensis (3,100 –3,375 Hz in A. femoralis ; 4,200 Hz in A. humilis ; 2,900 –3,600 Hz in A. hodli ; 3759–4689 Hz in A. nidicola ; and 4050–4930 Hz in A. paleovarzensis ); from A. bacurau , A. caldwellae , A. juami , A. marchesianus , A. melanolaemus , A. sieggreenae , A. sumtuosus , and A. tinae by having calls emitted continuously (calls arranged in call series in the other species). The only species with multi-note calls among those compared here are A. flaviventris , A. granti , A. trilineatus , and A. velocicantus . However, A. gasconi differs from A. granti , A. trilineatus and A. velocicantus by having calls with 2–4 notes that are emitted continuously (call consisting of 2 notes, arranged in call series with 6–11 calls in A. trilineatus ; call series with 1–17 calls and average 6.2 ± 2.6 calls per series in A. granti ; and call composed of 66–138 pulsed notes with a note duration of 5–13 ms in A. velocicantus ), and from A. flaviventris by having a dominant frequency of 5,081 –6,761 Hz (3,618 –4,651 Hz in A. flaviventris ). The advertisement calls of A. chalcopis , A. conspicuus , A. fuscellus , A. insperatus , A. ornatus , A. pacaas , A. pittieri , and A. vanzolinius remain unknown.

Tadpoles of Allobates gasconi have only two tooth rows on the posterior labium, which differs from A. caldwellae , A. femoralis , A. granti , A. hodli , A. kamilae , A. paleovarzensis , A. pittieri , A. subfolionidificans , A. sumtuosus , and A. velocicantus (three rows in the other species) and from A. nidicola (teeth absent). Like A. gasconi , tadpoles of A. marchesianus have two tooth rows on the posterior labium, but A. gasconi also has two tooth rows on the anterior labium (only one row in A. marchesianus ). Tadpoles of A. algorei , A. amissibilis , A. bacurau , A. chalcopis , A. conspicuus , A. flaviventris , A. fuscellus , A. humilis , A. insperatus , A. juami , A. melanolaemus , A. ornatus , A. pacaas , A. sieggreenae , A. tinae , A. trilineatus , and A. vanzolinius have not been described.

Adult morphology. The following description of external morphology is based on adult males of the type series (n = 6) and recently collected males (n = 24) from Jainú West, Jainú East, and Altamira. Coloration of males and females is described in the next section (Intraspecific variation).

A small-bodied species, SVL 14.8–17.1 mm in males and 16.0–18.0 mm in females. Head slightly wider than long ( HW /HL = 106 ± 8%), head width 30 ± 2% of SVL and head length 28 ± 2% of SVL. Snout blunt, rounded to nearly truncate in dorsal view and acutely rounded in lateral view; snout length 55 ± 6% of HL; canthus rostralis predominantly concave in dorsal view; internarial distance 49 ± 3% of HW; nostril opening directed laterally, and only visible laterally and ventrally; eye length 90 ± 5% of EN; tympanic membrane inconspicuous, rounded and small, TYM 38 ± 4% of EL; tongue attached anteriorly, longer than wide, slightly rounded posteriorly and with scattered small papillae; median lingual process absent; vocal sac distinct, single and subgular; teeth present on the maxillary arch.

Forearm slightly thicker and smaller than the upper arm ( FAL / UAL = 92 ± 8%); forearm and upper arm length represent 22 ± 2% and 24 ± 2% of SVL, respectively; black arm glands absent. Hand small, HANDIII 24 ± 2% of SVL; paired dorsal digital scutes present; fingers weakly expanded, but preaxial swelling well developed on finger III ( Fig. 4 ); width of the third phalange of finger III 82 ± 13% of WFD; basal finger webbing absent; when finger IV is appressed, its tip reaches the base of distal subarticular tubercle of finger III in most individuals; relative length of fingers III > I> II > IV; carpal pad absent; excrescences Phylogenetic position of the Amazonian nurse frog Allobates gasconi ( Morales 2002) revealed… 107 on thumbs of males absent; palmar tubercle rounded, diameter 14 ± 2% of HANDIII ( Fig. 4 ); thenar tubercle width 56–133% of the DPT; two subarticular tubercles on finger III, a single proximal subarticular tubercle on fingers I, II and IV ( Fig. 4 ); distal subarticular tubercle of finger IV absent; supernumerary tubercle present on the outer edge of the palm; disc expanded on all fingers; disc on finger III wider than the others; poorly defined basal and lateral keels on pre- and postaxial edges of all fingers.

Hind limbs robust; tibia slightly longer than thigh (TL/ THL = 103 ± 6%); tibia and thigh length 47 ± 4% and 46 ± 3% of SVL, respectively; foot length 96 ± 5% of TL and 99 ± 7% of THL. Relative length of toes IV > III > V > II > I ( Fig. 5 ); black band above elbow absent; basal webbing present between toes II – III and III – IV; tip of toe I reaches the subarticular tubercle on toe II; tip of toe III reaches the medial subarticular tubercle on toe IV; tip of toe V reaches the mid-level of the proximal phalange of toe IV ( Fig. 5 ); discs of toes I and V moderately expanded, moderately to greatly expanded on toes II, III and IV ( Fig. 5 ); width of disc on toe IV 69–150% of disc on finger III ( Fig. 5 ). Inner metatarsal tubercle oval and outer metatarsal tubercle rounded; median metatarsal tubercle absent; tarsal keel short, tubercle-like and strongly curved at proximal end, extending from metatarsal tubercle; foot and tarsal fringe present; weak metatarsal fold present, small and swollen midway between the outer metatarsal tubercle and the first subarticular tubercle of toe V. A single subarticular tubercle on toes I and II, two subarticular tubercles on toes III and V, and three subarticular tubercles on toe IV; proximal tubercles small and poorly defined on toes III – V; disc on all toes wider than the penultimate phalanges; poorly developed lateral keels on preaxial and postaxial sides of all toes; basal keels between toes I– II and IV – V.

Skin smooth on anterior portion of the dorsum, slightly tubercular from mid-body to the sacral region; dorsal surface of forelimbs smooth; dorsal surface of hind limbs with small and scattered irregular tubercles; and flanks and ventral surface smooth.

Intraspecific variation. The adult morphology of the original type series and of additional recently collected specimens are similar. They differ only in the degree of swelling on finger III of males, which is less prominent in the type series, collected in 1991, than in the recent sample collected in 2021, likely due to reproductive state or a preservation artifact. Moreover, all specimens have similar dorsal color patterns, which differ mostly in color contrast. External coloration appears to have faded in the type series in comparison with the fresh material, probably due to the greater time in preservative and possibly due to different procedures of fixation and/or preservation.

Canthus rostralis is concave in 84% of specimens and straight in 16%; finger I longer than finger II in 95% of specimens, same size in 5%; finger II longer than finger IV in 70% of specimens, same size in 27.5% and shorter in 2.5%; finger III strongly swollen in adult males; when appressed, the tip of finger IV reaches the base of the distal subarticular tubercle of finger III in 74% of specimens but does not reach there in 26%; thenar tubercle elliptic in 52% of specimens and sub-circular in 48%; tarsal keel short, tubercle-like in all specimens; basal webbing present between toes II and III in 70% of specimens and between toes III and IV in all; basal keels present between toes of 78% of specimens; a small fold midway between the outer metatarsal tubercle and the first subarticular tubercle of toe V in 48% specimens.

Allobates gasconi is sexually dimorphic in several traits. The snout is longer in females (SL; W = 125.5, p = 0.045), as well as the width of disc on toe IV ( WTD; W = 134.5, p = 0.009). On the other hand, females tend to be slightly larger than males ( SVL; W = 123.5, p = 0.064). Morphometric measurements are summarized in Table 2.

There is no sexual dimorphism in dorsal coloration. In life, ground color varies from orange brown to light brown ( Figs. 6 and 7 ). A dark hourglass-shaped mark extends from the interorbital region to the posterior dorsum, ending before the cloacal opening ( Fig. 8 ); it can be conspicuous ( Fig. 8 ; MPEG 44619 and 44620) or inconspicuous ( Fig. 8 ; MPEG 44622; INPA-H 3491). Scattered small brown, non-keratinized tubercles on posterior dorsum; medial line absent; pale cream dorsolateral stripe with melanophores has a regular border laterally but not medially; dark lateral stripe from snout tip to groin covers the upper half of the tympanum and is fainter but wider posteriorly; oblique lateral stripe present as a whitish but incomplete diffuse line or series of small diffuse spots that extend from the inguinal region to mid-body; ventrolateral stripe absent, but irregular white spots present along the inferior edge of the lateral stripe; arm and forearm pale orange to brown, with dark brown longitudinal band along postaxial sides; anterior portion of arm light yellow or orange-white; dark disks with paired iridescent-white scutes. Dorsal surfaces of hind limbs pale orange or yellowish brown, with a transverse dark brown bar on thigh, shank and tarsus in 90%, 97.5% and 98% of specimens, respectively. Paracloacal mark whitish or cream, short, half-moonshaped in 97% of specimens, always surrounded by a dark brown frame. Fingers brown to pale orange; toes light to dark brown with transverse white bars; paired dorsal digital scutes present on digits I, II and III, with coloration varying from white to black in 71% of specimens, and only white in 29%; digit IV is always black. Iris matte black in 45% of specimens and matte black and silver in 55%, with a thin, dark-brown reticulation ( Fig. 7 ).

Ventral coloration is sexually dimorphic. In life, males have gray to dark gray throats and vocal sacs with a profusion of melanophores ( Figs. 6 and 7 ); females have bright- to whitish-yellow throats with few scattered dark melanophores on jaw edges. The chest is translucent to pinkish grey in males, and bright- to whitish-yellow without dark spots in females. In males, the belly is pinkish-grey anteriorly, whitish-gray centrally, light- or translucent-gray with white and brown small blotches laterally, and yellowish or translucent-gray posteriorly. Some males have melanophores over the entire belly, but most have densely concentrated melanophores until the midbody; one recently collected male has a darkly pigmented belly ( Fig. 8 ; MPEG 44622). In females, the belly is uniformly yellowish-cream to whitish-yellow with few scattered melanophores on the outer margins. Both sexes have pale- to translucent-grey thighs, and shanks and tarsus grayish with a few scattered melanophores. Palmar and plantar surfaces light-brown, dark-grey or dark-brown, respectively.

In preservative ( Fig. 8 ), dark dorsal marks are similar to those observed in life, but orange-brown shades have faded to pale-brown or pale-gray and dark-brown coloration has become blackish-gray. Paracloacal marks are paler than in life. White spots above lateral stripe have disappeared in preserved specimens, and ventral melanophores in males are less intense ( Fig. 8 ; MPEG 44619 and 44622). Bright- and whitish-yellow parts, as well as light-gray and translucent-white ones, have become cream or light-cream in preserved specimens.

Bioacoustics. The advertisement calls of 14 male Allobates gasconi were analyzed, with eight presenting calls with 2–3 notes, three with calls containing 2–4 notes, two with calls consisting of 3–4 notes, and one individual with a call consisting of only 2 notes (each call emitted with one exhalation). Notes were repeated at a rate of 116.2 ± 33.5 notes/min (43.4 notes/ min for two-note calls, 67.2 notes/min for three-note calls, and 5.5 notes/min for four-note calls). The duration of two-, three-, and four-note calls was 59 ± 4 ms, 98 ± 27 ms, and 150 ± 11 ms, respectively. The inter-call interval had a duration of 387 ± 154 ms (161–1,048 ms). The first and last notes had similar durations, with a note duration of 15 ± 4 ms (6–28 ms) and an inter-note interval of 30 ± 12 ms (19–42 ms).

Notes were frequency-modulated ( Fig. 9 ). The first note was usually emitted with a lower amplitude than the last note; the average linear frequency modulation was 130 ± 192 Hz (904–560 Hz). The first note had a low frequency of 5,373 ± 264 Hz (4,795 –6,487 Hz), a high frequency of 5,810 ± 267 Hz (5,261 –6,878 Hz) and a dominant frequency of 5,613 ± 270 Hz (5,082 –6,761 Hz). The last note had a low frequency of 5,309 ± 168 Hz (5,007 –5,643 Hz), a high frequency of 5,790 ± 155 Hz (5,461 –6,059 Hz), and a dominant frequency of 5,573 ± 165 Hz (5,297 –6,059 Hz). However, differences in spectral parameters between the first and last notes were not statistically significant, and frequencies are given as overall values in Table 3.

Tadpole morphology. The following description is based on 14 tadpoles at Gosner stage 36 (lots INPA –H 45090–91) oscillograms of three note arrangements: B two-, C three- and D four notes (each call arrangements corresponds to an expiration). Illustration: A. S. Ferreira from Jainú West and Altamira. Table 4 summarizes 27 morphometric traits of 25 additional tadpoles at stages 34–41.

The body is ovoid in dorsal view, rounded anteriorly but truncate posteriorly, and longer than wide ( BL /BW = 157%; Fig. 10 ); it is flattened in lateral view (BH/BL = 64%). Body length is 31% of total length and head width 92% of body width. The snout is bluntly rounded in dorsal view but slightly pointed in lateral view; snout length slightly less than eye diameter ( NSD /ED = 86%). Large eyes are positioned dorsally and directed laterally; eye diameter 96% of END; interorbital distance 151% of IND and 63% of HWLE. The small naris is located dorsally and directed anterolaterally, visible in dorsal and lateral measurements (in mm) of tadpoles of Allobates gasconi from Jainú West (lot INPA –H

45090) and Altamira (lot

INPA –H 45091), middle Juruá

River, Amazonas, Brazil views; internarial distance 42% of HWLE. The spiracle is sinistral, visible in dorsal, lateral and ventral views, easily seen at approximately mid-body and below lateral midline and directed transversely; inner wall free from body; spiracle-tube length 15% of HWLE. The gut is coiled and visible through the skin; its axis is directed to the left side of the body. The medial vent tube is dextral 1.0 mm, attached to ventral fin, right wall displaced dorsally; vent-tube length 200% of spiracle length. Tail length is 69% of total length; tail maximum height 92% of BH; tail muscle robust and deeper than fins, tail muscle maximum height 63% of MTH. The dorsal fin emerges at the thigh-body insertion and reaches its maximum depth at the central portion of the tail; dorsal fin deeper than the ventral fin along the first two thirds of the tail. Tail tip acuminate, lacks flagellum.

Oral disc is located anteroventrally and emarginated laterally ( Fig. 11 ); oral-disc width 36% of body width. Anterior labium has small pyramidal papillae only on lateral margin, 2 on each side. Posterior labium has a single marginal row of pyramidal papillae of variable length; short papillae are located close to the labium angle, all other papillae are elongate ( Fig. 11 ); elongate papillae twice as long as the shorter ones. Submarginal papillae are absent. Upper jaw sheath is arch-shaped; lower jaw sheath “ V ”-shaped, deeper than upper sheath; cutting edge of each jaw sheath is serrated along its entire length. Labial keratodont row formula is ( LKRF) 2(2)/2(1); tooth row A–1 complete; tooth row A–2 interrupted by a medial gap of ~ 0.5 mm, each segment ~ 0.4 mm long; tooth row P–1 interrupted by a small medial gap; P–1 slightly (0.2 mm) longer than P–2; P–3 absent. Vent tube and tooth row A–2 disappear at stage 41.

In life, dorsal and lateral surfaces of body are grayish brown with scattered irregular brown blotches. Ventral surface of body and tail musculature are the same as in preservative. At stage 36, fins are transparent with scattered irregular brown and silver blotches. In preservative, the dorsal skin of head and body is whitish-cream to yellowish-cream and covered with dark melanophores ( Fig. 10 ). There is a higher concentration of melanophores in the center of the dorsum, but without an obvious hourglass-shaped mark. A dark brown dorsolateral stripe extends from the tip of the snout to the inguinal region in some specimens. Dark black blotches are present on the latero-frontal portion of head. The tail muscle is light cream and fins are translucent or pale with brown to dark brown blotches. Ventral surfaces are yellowish cream with brown irregular-shaped blotches surrounding the posterior labium until the beginning of pericardium and abdomen region; dark melanophores randomly distributed on posterior portion of ventral surface of head and tail; posterior region of ventral surface translucent and unpigmented, intestines visible through the skin.

Distribution and natural history notes. Allobates gasconi is a common diurnal, terrestrial frog that inhabits the leaf litter of seasonally flooded (várzea) forests along the middle and upper Juruá River in Brazil, and along the Yuyapichis River (affluent of the Pachita River, Ucayali Basin) in Peru. The specimen INPA-H 4889 from the paratype locality Vai-Quem-Quer (lower Juruá River) has morphological differences from the other individuals of the type series. In addition, we did not find A. gasconi at this locality, but there is another species with similar call and morphology (Ferrão & Ferreira, pers. comm.). Despite intense sampling in southwestern and western Brazilian Amazonia, A. gasconi has not been recorded in unflooded (terra firme) forests along the Juruá River or in unflooded or seasonally flooded forests along the Madeira, Purus and Solimões (upper Amazon) rivers (unpublished data, A.P. Lima, M. Ferrão, A.S. Ferreira). Breeding coincides with the rainy season from November to March. Males call while perched above the ground on shrub roots, the adaxial surface of live leaves and fallen logs, and within rolled leaves in the leaf litter. They are vocally most active at dawn (0530–0830 h) and dusk (1630–1800 h), and throughout the day on rainy days. Breeding males are territorial and exhibit aggressive behavior; in addition to responding to advertisement-call playback, two males were encountered in wrestling bouts. Territory size, however, remains unknown. When a female approaches a calling male, he emits low courtship calls in her direction, which attract her. We found clutches of 20– 29 eggs deposited on the adaxial surface of live leaves of small shrubs and in dead rolled leaves in the leaf litter. Each oviposition site contained only one clutch. Eggs are enclosed by a layer of transparent jelly, which becomes cloudy during embryonic development ( Fig. 12 ). Adults were not observed transporting tadpoles, nor were tadpoles found in temporary ponds, which leads us to hypothesize that tadpoles complete larval development in flooded forest (várzea) of the Juruá River. Syntopic diurnal species include Allobates femoralis , Ameerega trivittata (Spix, 1824) , and two probably undescribed species of Allobates .

Figures

Fig. 3 A Dorsal and B ventral views of the sub-adult male holotype (INPA-H 3082) of Allobates gasconi from Jainú West, Juruá River, Amazonas, Brazil. Photographs by A. S. Ferreira

Fig. 4 Ventral views of left hands of preserved male and female Allobates gasconi. A Adult male paratype (INPA-H 3249), B, C topotypic adult males (INPA-H 45073 and MPEG 44622, respectively), and D topotypic adult female (MPEG 44620). Photographs by A. S. Ferreira

Fig. 5 A Ventral surface of the right foot of a preserved topotypic adult male (MPEG 44622) and B right foot of topotypic adult female (MPEG 44620) of Allobates gasconi. Photographs by A. S. Ferreira

Fig. 6 Color in life of adult males and females of Allobates gasconi. A, B Topotypic adult males, INPA-H 45067 and INPA-H 45070, respectively. C, D Topotypic adult females, INPA-H 45074 and 45076, respectively. Photographs by A. P. Lima

Fig.7 Coloration in life of A, B, C topotypic adult males and D females of Allobates gasconi. Note the densely pigmented throat and vocal sac of actively calling males. Photographs by B, C A.P. Lima and A, D W. E. Magnusson

Fig. 8 Dorsal, ventral and lateral color-pattern variation among preserved Allobates gas- coni. A–C Adult female, MPEG 44620, from Jainú West. D–F adult male, MPEG 44619, from Jainú West. G–I Adult male, MPEG 44622, from Jainú East. J, K Adult female paratype, INPA-H 3491, from Jainú East. Photographs by A. S. Ferreira

Fig. 9 Advertisement call of Allobates gasconi from its type locality. A Oscillogram showing the continuous emission of 10 calls with distinct note arrangements (2–4 notes per call). Spectrograms and

Fig. 10 A Dorsal, B lateral, and C ventral views of a preserved tadpole of Allobates gasconi, Gosner stage 36, from Jainú West (lot INPA– H 45090). Photographs by A. S. Ferreira

Fig. 11 Detailed views of the oral disc of tadpoles of Allobates gasconi, Gosner stage 36, from Jainú West. A Mouth dyed with blue dye and B mouth without dye. Photographs by A. S. Ferreira

Fig. 12 Allobates gasconi eggs and embryos from Jainú West and Altamira. A Freshly laid clutch on a green leaf contains 42 eggs, each enclosed within transparent jelly and with a pig- mented animal pole. B Embryos of the clutch in A approximately 12 days after deposition. C Clutch laid on humid leaf con- taining 28 eggs approximately 3 days after deposition. D Clutch containing 31 embryos approxi- mately 6 days after deposition. Photographs by A. S. Ferreira