Pristimantis gagliardoi Bustamante & Mendelson

Bustamante, Martín R. & Mendelson, Joseph R., 2008, A new frog species (Strabomantidae: Pristimantis) from the High Andes of Southeastern Ecuador, Zootaxa 1820, pp. 49-59 : 50-57

Pristimantis gagliardoi Bustamante & Mendelson , new species

Figs. 1–5

Holotype. QCAZ 27103, an adult female ( Figs. 1–4 ) obtained by MRB at La Libertad, Reserva Mazar, (S 02° 32' 47'' S, 078° 41' 54.1'' W, 2895m). Provincia Cañar, Ecuador, on October 25, 2005.

Paratopotypes. QCAZ 27104, an adult female collected by MRB on 22 October 2005; QCAZ 27568, 27578, 27580–1, adult males, and QCAZ 27577, 27579, two adult females, collected by the authors and Michelle R.Cummer on 26–27 February 2004; QCAZ 29561, an adult male collected by MRB on 16 March 2005; QCAZ 32623, an adult female collected on 20 November 2006 by MRB.

Diagnosis. A member of P. unistrigatus (sensu Lynch and Duellman 1997, as modified by Duellman and Pramuk 1999, Hedges et al. 2008,) diagnosed by having (1) skin texture of dorsum shagreen with scattered tubercles, those on venter areolate; (2) tympanic membrane and annulus evident, 29.7–40% of eye length on females and 25.2–32.3% on males, with supratympanic ridge obscuring upper and posterodorsal edges; (3) snout rounded in profile and dorsal views; (4) upper eyelid bearing one large, and several small scattered, conical tubercles; cranial crests absent; (5) dentigerous process of the vomer triangular and narrowly separated, each bearing 4–6 teeth; (6) vocal slits and vocal sac absent in males, nuptial pads present ( Fig. 5 ); (7) first finger shorter than the second, fingers with expanded discs; (8) fingers with lateral fringes; (9) ulnar tubercles small; (10) heel with one enlarged conical tubercle; tarsus with 3–4 tubercles along the outer edge; (11) two metatarsal tubercles prominent, inner oval, about three times the size of the outer subconical tubercle; numerous supernumerary plantar tubercles; (12) toes with lateral fringes; (13) in ethanol, dorsum tan, gray or dark gray, with darker markings (sometimes chevrons or bars) outlined by white or black lines; venter cream to pale cream with darker spots; palmar and plantar surfaces gray, with Fingers I–II, and Toes I–III being the palest; (14) adults medium sized, in males 19.06–24.33 SVL (= 22.17 + 2.01, n = 6), females 26.83–33.64 SVL (= 30.63 + 3.10, n = 5).

We assign this species to the Pristimantis unistrigatus group based on the following characters: narrow head (head width 38.5–42.6% of SVL in females and 38.3–41.9% in males); absence of cranial crests; upper eyelid as wide as, or wider than, IOD (75.1–96.9% in females, 96.0–112.0% in males); skin on venter aerolate; nuptial pads present; tympanic membrane evident; discs on fingers and toes broad; Finger I shorter than Finger II and toes being unwebbed.

Comparisons. Pristimantis gagliardoi may easily be identified by the presence of large, conical tubercles on the eyelids and heels. This species differs from other species in the group that have some degree of development of such tubercles as follows and by the presence/absece of other characters (condition of P. gagliardoi in parentheses) ( Table 1 ): Pristimantis cryptomelas ( Lynch 1979) is the species most similar to P. gagliardoi , but differs by having a black coloration on the anterior and posterior sides of thighs, the posterior lower flanks, and the concealed portion of the shank (pale reddish), by having smooth skin (shagreen with scattered distinct tubercles), by lacking a conical tubercle over the eye (present), and by having larger pads on the digits of the hands and feet. Pristimantis inusitatus ( Lynch & Duellman, 1980) differs by having a green dorsum and white venter (dorsum pale brown with variable dark brown markings, venter cream with variable dark brown reticulations or markings), conical tubercle on upper eyelid (present, but larger in P. gagliardoi ), snout protruding in profile and subcauminate in dorsal view (rounded in both views). Pristimantis eriphus ( Lynch & Duellman, 1980) has a straight canthus rostralis (weakly angular), adult females have red iris (bronze) and flanks with white areas and slightly oblique white bars on posterior flanks and hidden surfaces of hind limbs (bars on flanks). Pristimantis prolatus ( Lynch & Duellman, 1980) has no ulnar tubercles (few conical), vocal slits and vocal sac present in males (absent), Iris is gray brown (bronze), snout in profile nearly truncate and accuimnate in dorsal view (rounded in both views). Pristimantis nephophilus ( Duellman and Pramuk, 1999) has a few, small tubercles on the posterior eyelid (small tubercles scattered over surface of eyelid, and a single, large, central, conical tubercle), dark brown coloration with cream spots on the posterior thighs (pale reddish), brown coloration on the venter (cream with variable markings), all tubercles on the feet and hands weakly developed (strongly developed), smoother skin, lacking pronounced tubercles (shagreen with scattered distinct tubercles), and discs on the hind digits being slightly smaller than those of fingers (equal in size).

Description of the holotype. Head slightly wider than long, head length 85% of head width ( Fig. 1 ); snout relatively short (snout to eye distance 15.1% of SVL), rounded in dorsal view and in profile ( Figs. 1–2 ); eye to nostril 92.7% length of eye; nostrils not protuberant, directed anterolaterally; canthus rostralis weakly angular; loreal region concave; lips not flared; interorbital area flat, broader than upper eyelid (upper eyelid width 85.9% of interorbital distance); upper eyelid with distinct conical fleshy tubercle situated centrally on outer margin of eyelid; cranial crests absent; supratympanic fold weak, but partially obscuring upper margin of tympanic annulus and tympanum; side of head nearly vertical; tympanic annulus distinct; tympanic membrane weakly pustular, not thickened; length of tympanic annulus 40% length of eye; postrictal tubercles large, subconical, situated ventral and posteroventral to tympanic annulus; skin on head finely granular; choanae very small, oval, not concealed by palatal shelf of maxillary; vomerine odontophores oblique, posteromedial to choanae, oval in outline, about 3– 4 x size of choana, separated medially by distance less than width of odontophore, each bearing 4 teeth; tongue wider than long, its posterior border bilobbed, posterior half not adherent to floor of mouth.

Canthus rostralis Rounded Straight (weakl Sharp Weakly angular Moderately Weakly angular

concave) (barely concave) sharp

Snout in profile Rounded Round Protruding Rounded Nearly truncate Rounded Dorsum of head, body, and limbs with scattered subconical tubercles becoming larger and more abundant on lateral surfaces; dermal ridge from posteromedial edge of eyelid to scapular region; dorsolateral folds absent; flanks granular; cloacal sheath absent; enlarged tubercles in cloacal region present, moderately developed; skin on throat granular; skin on venter areolate; discoidal fold not evident; forearm slender; radio-ulna length 25% of SVL; ulnar tubercles few, round to subconical; thenar tubercle ovoid, about 3x size of bifid subpalmar tubercle; supernumerary tubercles few, large subconical; subarticular prominent, subconical; hand length longer than radioulna length 33.1% of SVL; Fingers II–IV bearing narrow lateral fringes; relative lengths of fingers I <II <IV <III; disc on thumb slightly expanded; discs on Fingers II–IV extensively expanded, truncate, less than twice width of digits; Finger I having ventral pad defined by circumferential groove, absent on other fingers ( Fig. 3 ).

Hind limbs relatively slender; tibia length 52.9% of SVL; foot length 95.3% of tibia length; heel bearing small, subconical tubercle; outer edge of tarsus bearing three large subconical tubercles; inner metatarsal tubercle flat, ovoid, about 3x subconical outer metatarsal tubercle; supernumerary plantar tubercles prominent, round or subconical; subarticular tubercles round, not conical; toes bearing narrow lateral fringes, most developed on Toes IV–V; relative length of toes I <II <III <V <IV; webbing absent; discs on toes equal in size to discs on fingers; tip of Toe V extending to distal edge of distal subarticular tubercle on Toe IV; tip of Toe III not extending to that tubercle ( Fig. 3 ).

Coloration of holotype. In life: Dorsum tan with brown bars outlined by thin black lines, suprascapular region with a dark brown W mark corresponding to suprascapular dermal ridges; dark brown interorbital bar; brown cloacal region. Dorsal surfaces of limbs tan with brown and dark brown bars. Dorsum of hands and feet with black blotches, more abundant towards distal portions of fingers and toes. Flanks tan with black blotches. Axial region and groin pink with brown blotches. Hidden parts of venter and forelimbs pink with brown blotches. Ventral surfaces of tarsus and outer toes dark brown. Yellow belly with dark brown spots, yellowish throat with brown blotches. Two dark brown blotches in ventral scapular region. Venter of hands dark brown, inner fingers (I and II) pale brown; black bars originating below eye and extending towards the lip. Iris bronze with black reticulations ( Fig. 4 ).

In preservative: Dorsum gray with darker interorbital bar and a W over scapular region; dorsal surfaces of proximal forelimbs dull cream, distal forelimbs dull gray with darker markings; dorsal surfaces of Fingers I and II pale cream, Fingers III and IV pale gray with dark gray markings and paler discs; dorsal surfaces of thighs pinkish gray with darker blotches; shanks gray with darker bars, tarsus gray with darker markings, Toes IV and V same color, Toes I–III cream. Dorsum of tympanum with black markings; gray loreal, canthal, and upper labial regions with black markings. Venter tan with dark gray spots, pale cream throat with gray markings; venter of proximal forelimbs pinkish becoming darker on ulna and outer fingers, inner fingers cream; ventral hindlimbs pinkish with brown markings, on tarsus and Toes IV and V becoming dark grey, and Toes I– III cream ( Fig. 1 ).

Measurements of holotype. SVL = 32.49; tibia length = 17.19; foot length = 16.38; head length = 10.84; head width = 12.75; interorbital distance = 4.03; upper eyelid width = 3.46; internarial distance = 2.79; eye to nostril distance = 3.57; snout to eye distance = 5.06; eye diameter = 3.85; tympanum diameter = 1.54; eye to tympanum distance = 1.40; radioulna length = 8.12; hand length = 10.78; and Finger I length = 5.13.

Variation. Variation in measurements and proportions among additional specimens are presented in Tables 2 and 3 . Variation of coloration in life: QCAZ 27570: Dorsum with cream and pale brown bars outlined by thin black lines; flanks cream with bars originating on the dorsum; dorsum of limbs showing the same pattern as the body; groin and hidden surfaces of thighs colorless; venter cream; iris bronze with a brown medial stripe; upper lip with vertical brown and cream bars; QCAZ 27104: Dorsum tan with brown blotches outlined by thin black lines; flanks brown with diagonal bars becoming greenish toward the dorsum; dorsum of forelimbs with brown and greenish bars; tarsus tan with brown bars; dorsum of arms tan with brown blotches; dorsum of hands and feet with black blotches present on outer fingers; groin, armpit, and hidden parts of thighs pale pink with brown blotching; throat tan with brown blotching; venter of limbs tan with dark brown blotching with outer fingers and toes darker; black bars originating below eye and extending towards the lip. Iris bronze with black reticulations; QCAZ 27434: Same pattern as QCAZ 27104 but having diagonal green bars extending from the dorsum onto the hidden surfaces of thighs; groin and armpit are orange.

Etymology. The specific name is a noun in the genitive case and is a patronym for Ron Gagliardo, dedicated collaborator in both research and conservation programs involving Neotropical amphibians. Gagliardo has been important in promoting amphibian research, environmental education, but especially for supporting the establishment of captive breeding programs for the threatened Ecuadorian frogs.

Distribution and ecology. Pristimantis gagliardoi is known only from La Libertad, Reserva Mazar (S 0 2 32 ' 47'' S, 0 78 41' 54.1'' W, 2895m), Provincia Cañar, Ecuador. This site, a small farm, is located on the western versant of the Ecuadorian Andes in Bosque de neblina montano (Montane cloud forest) and Bosque siempreverde montano alto (Evergreen montane forest) according to Valencia et al. (1999) ( Fig. 6 ). The species has been found on leaves and branches at heights varying from 0.40–1.50 m above ground (= 0.775 m), mainly in secondary forests (12 of 16 individuals), and on vegetation over very small stream (2 of 16) and in primary forest (1 of 16). During our visits (six field trips since February 2004) to La Libertad, P. gagliardoi showed seasonal activity. During the wettest months we found it in primary and secondary forest as well as along small streams. In drier months (October and November) we found it along streams. An adult female (QCAZ 32623) obtained on 20 November, 2006, contained mature eggs in the oviducts. Pristimantis gagliardoi occurs sympatrically with Gastrotheca pseustes , Hyloxalus vertebralis , P. orestes , P. pycnodermis , P. riveti and an undescribed Pristimantis species.

Figures

FIGURE 1. Dorsal (left) and ventral (right) views of Pristimantis gagliardoi in preservative; holotype, adult female (QCAZ 27103), SVL = 32.49 mm.

FIGURE 2. Lateral view of the head holotype of Pristimantis gagliardoi, holotype, adult female (QCAZ 27103).

FIGURE 3. (A) Right hand of Pristimantis gagliardoi; (B) right foot of Pristimantis gagliardoi. Holotype, adult female (QCAZ 27103).

FIGURE 4. Dorsolateral (A) and ventral (B) views of Pristimantis gagliardoi in live; holotype, adult female (QCAZ 27103), SVL = 32.49 mm.

FIGURE 5. Nuptial pad on Finger I of a Pristimantis gagliardoi male (QCAZ 27580).

FIGURE 6. Map of Ecuador showing the type locality of Pristimantis gagliardoi.

Tables

TABLE 1. Character states in similar Pristimantis species associated to the unistrigatus group.

Character	P. cryptomelas P. eriphus	P. inusitatus	P. nephophilus P. prolatus	P. gagliardoi
SVL males	28.2–30.2 18.1–25.2	-	- 13.7–18.4	19.1–24.3
SVL females	38.6 25.8–29.0	24.0–24.4	24.6–34.0 20.8–24.1	26.8–33.6

TABLE 2. Measurements (in mm) of adults of Pristimantis gagliardoi. Range, mean SD are given. SVL = snout – vent length.

Character	females n = 5	males n = 5
SVL	26.83–33.64 (30.63 ± 3.10)	19.06–24.33 (22.17 ± 2.01)
Tibia length	15.23–18.17 (16.68 ± 1.29)	10.98–14.97 (12.70 ± 1.50)
Foot length	13.21–17.05 (15.23 ± 1.86)	9.22–13.04 (11.16 ± 1.46)
Head length	9.82–11.43 (10.70 ± 0.60)	6.76–9.05 (7.87 ± 0.85)
Head width	10.71–13.34 (12.22 ± 1.00)	7.73–10.19 (9.02 ± 0.94)
Interorbital distance	3.20–4.03 (3.51 ± 0.33)	2.18–2.85 (2.60 ± 0.28)
Upper eyelid width	2.46–3.46 (3.25 ± 0.39)	2.32–2.83 (2.65 ± 0.22)
Radioulna length	2.58–2.91 (2.81 ± 0.14)	1.83–2.64 (2.23 ± 0.29)
Eye to nostril distance	2.43–3.58 (3.23 ± 0.34)	2.01–2.83 (2.33 ± 0.31)
Snout to eye distance	4.52–5.75 (5.10±0.51)	3.31–4.55 (3.85 ± 0.45)
Eye diameter	3.45–4.12 (3.78 ± 0.27)	2.40–3.00 (2.83 ± 0.24)
Tympanum diameter	1.04–1.54 (1.25 ± 0.19)	0.70–0.80 (0.76 ± 0.03)
Eye to tympanum distance	1.32–1.75 (1.51± 0.17)	1.10–1.35 (1.18 ± 0.10)
Radio-ulna length	6.92–8.50 (7.83 ± 0.65)	5.26–6.94 (5.85 ± 0.66)
Hand length	8.43–10.78 (9.38 ± 1.00)	5.92–7.48 (6.56 ± 0.68)
Finger I length	3.95–5.13 (4.73 ± 0.54)	2.91–3.80 (3.27 ± 0.34)

TABLE 3. Ranges of morphological proportions (in percentages) of adults of Pristimantis gagliardoi. SVL = snout – vent length.

Character	females (n = 5)	males (n =5)
Tibia length/ SVL	51.7–57.5	53.8–61.5
Foot length/SVL	47.5–50.4	47.7–51.4
Foot length/tibia length	85.6–95.3	84.0–92.5
Head width/SVL	38.5–42.6	38.3–41.9
Head length /SVL	33.4–37.8	34.6–37.2
Head length/ head width	85.0–91.7	84.0–92.1
Eye–to–nostril distance/eye diameter	79.5–92.7	72.0–97.9
Upper eyelid width/IOD	75.1–96.9	96.0–112.0
Radioulna length/SVL	24.4–26.6	25.1–28.5
Hand length/radioulna length	103.2–132.7	101.5–125.9
Finger I Length/hand length	46.9–58.2	41.8–66.0