Two syntopic new species of the Pristimantis orestes Group (Anura: Strabomantidae) from northwestern Peru

Zootaxa 3249: 47–59 (2012) www.mapress.com / zootaxa/ Copyright © 2012 · Magnolia Press

ISSN 1175-5326 (print edition)

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Two syntopic new species of the Pristimantis orestes Group (Anura: Strabomantidae) from northwestern Peru PABLO J. VENEGAS1,3& WILLIAM E. DUELLMAN² ¹División de Herpetología-Centro de Ornitología y Biodiversidad (CORBIDI), Santa Rita N˚105 Of. 202, Urb. Huertos de San Antonio, Surco, Lima-Perú ²Biodiversity Institute, University of Kansas, 1345 Jayhawk Blvd., Lawrence, Kansas 66045 ³Corresponding author. E-mail: [email protected]

Abstract Two new species of Pristimantis (Anura: Strabomantidae), tentatively assigned to the P. orestes Group, are described from humid puna (3596 m) of Cañaris, Provincia de Ferreñafe, Región Lambayeque on the eastern slopes of the Cordillera Occidental in northwestern Peru. Pristimantis mariaelenae sp. n. is a medium-sized species characterized by the dorsum varying from brown to reddish ochre, groin and posterior surface of the thigh with white blotches, pearly white venter with brown blotches, distinct tympanic annulus and membrane, and tubercles on the eyelid, heel, and outer edge of the tarsus. Pristimantis stipa sp. n. also is a medium-sized species characterized by having narrow discs without marginal grooves, distinct tympanic annulus and membrane present, and Toe III slightly longer than Toe V. Key words: amphibians, Lambayeque, new species, Peru, phenetic assemblage, Pristimantis

Resumen Se describen dos nuevas especies de Pristimantis (Anura: Strabomantidae) procedentes de la puna húmeda (3596 m) de Cañaris, Provincia de Ferreñafe, Región Lambayeque de la vertiente este de la Cordillera Occidental en el noroeste de Perú. Ambas especies son tentativamente asignadas al grupo P. orestes. Pristimantis mariaelenae sp. n. es una especie de tamaño mediano caracterizada por poseer el dorso marrón o rojo ocre, la ingle y la superficie posterior de los muslos con manchas blancas, el vientre blanco perla con manchas marrones, un anillo y membrana timpánica claramente definidos, tubérculos sobre los parpados, talón y borde externo del tarso. Pristimantis stipa sp. n. también es una especie de tamaño mediano caracterizada por tener discos estrechos sin ranuras marginales, un anillo y membrana timpánica claramente definidos, y el Dedo III ligeramente más largo que el Dedo V en los pies. Palabras claves: anfibios, Lambayeque, especie nueva, Perú, agrupación fenética, Pristimantis

Introduction With more than 448 species, the strabomantid frogs of the genus Pristimantis, represent the most speciose genus in the neotropics(Amphibia Web 2012). The major diversity in the genus is in northwestern South America, where the distribution of the genus includes the lowlands to elevations of about 4000 m in the Andes of Colombia, Ecuador, and Peru (Hedges et al. 2008). Strabomantids occupy a variety of habitats, such as dry and humid lowland tropical forest, montane forest, puna, and paramo, but are absent in the arid coastal regions and on the semi-arid Pacific slopes of the Andes (Duellman & Lehr 2009). In Peru, 115 species of Pristimantis are currently known; the majority of species (72%) are restricted to the Andean habitats (Lehr et al. 2010). The Pristimantis orestes Group is a phenetic assemblage (sensu Lynch & Duellman 1997) and currently contains 15 species: P. atrabracus Duellman & Pramuk 1999; P. bambu Arteaga-Navarro & Guayasamin 2011; P. chimu Lehr 2007; P. cordovae Lehr & Duellman 2007; P. corrugatus Duellman, Lehr & Venegas 2006; P. melanogaster Duellman & Pramuk 1999; P. orestes Lynch Accepted by J.M. Padial: 9 Feb. 2012; published: 28 Mar. 2012

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1979; P. pataikos Duellman & Pramuk 1999; P. pinguis Duellman & Pramuk 1999; P. simonbolivari Wiens & Coloma 1992; P. seorsus Lehr 2007; P. simonsii Boulenger 1900; P. stictoboubonus Duellman, Lehr & Venegas 2006; P. ventriguttatus Lehr & Kohler 2007; and P. vidua Lynch 1979. The members of this group are distributed in paramo, puna, and upper humid montane forest at elevations of 2800–4200 m in the Andes of northern Peru and southern Ecuador (Lehr & Duellman 2007). Generally, members of the group are small (females less than 34 mm) with short snouts, robust bodies, relatively narrow heads, proportionately short limbs, Finger I shorter than Finger II, Toe V only slightly longer than Toe III, digital discs narrow and rounded, and the tympanum weakly defined (Hedges et al. 2008). Morphologically, members of this group are like frogs of the Pristimantis myersi Group that inhabit paramo and upper humid montane forest in the Andes of northern Ecuador and southern Colombia (Wiens & Coloma 1992; Hedges et al. 2008). A recent collection from the high grassy region of the Región de Lambayeque revealed two undescribed species of Pristimantis. Herein, we described these new species and assign them to the Pristimantis orestes Group. In so doing, we are following the definition of Pristimantis and the Pristimantis orestes Group and their placement in Strabomantidae as arranged by Hedges et al. (2008) and followed by Duellman & Lehr (2009). We are aware that the morphological characteristics of members of the Pristimantis orestes Group are similar to those of the Pristimantis myersi Group in Colombia and northern Ecuador. Thus, placement of the new species in the Pristimantis orestes Group is based partly on geography.

Material and methods The format for the descriptions follows Lynch & Duellman (1997) and diagnostic characters of Duellman & Lehr (2009). Specimens were preserved in 10% formalin and stored in 70% ethanol, and were sexed externally by the presence or absence of vocal slits and internally by the condition of the gonads. Measurements taken with digital calipers (to nearest 0.1 mm) were as follow: SVL (snout–vent length), TL (tibia length), FL (foot length, distance from proximal margin of inner metatarsal tubercle to tip of Toe IV), HL (head length, obliquely from angle of jaw to tip of snout), HW (head width, at level of angle of jaw), ED (eye diameter), IOD (interorbital distance), EW (upper eyelid width), IND (internarial distance), E-N (eye–nostril distance, straight line distance between anterior corner of orbit and posterior margin of external nares). Fingers are numbered preaxially to postaxially from I–IV. Comparative lengths of Toes III and V were determined when both were adpressed against Toe IV; lengths of Fingers I and II were estimated when adpressed against each other. All specimens were deposited in the herpetological collection of the Centro de Ornitología y Biodiversidad (CORBIDI), Lima, Peru. Specimens examined are listed in the Appendix. The Museo de Historia Natural Universidad Nacional Mayor de San Marcos, Lima, Perú, is abbreviated MUSM.

Descriptions of the new species Pristimantis mariaelenae new species Figs. 1 (A & B), 2(A–F), 3, 4 (A–L) Holotype: CORBIDI 02824, an adult female, from Cañaris, Provincia de Ferreñafe, Región Lambayeque, Perú (6˚7'14.4" S, 79˚18'4.43" W; 3596 m.a.s.l.) obtained on 16 June 2007 by Pablo J. Venegas. Paratypes: CORBIDI 02827–28, two adult females; CORBIDI 02823, 02825–26, 02829, 02831, five adult males; CORBIDI 02830 and 02832, two juveniles, collected with the holotype by Pablo J. Venegas. Diagnosis. A member of the Pristimantis (Pristimantis) orestes Group following the definition of Pristimantis and the Pristimantis orestes Group and their placement in Strabomantidae as arranged by Hedges et al. (2008) and followed by Duellman & Lehr (2009), having the following combination of characters: (1) skin on dorsum shagreen with scattered small tubercles; flanks tuberculate; skin on venter areolate; discoidal fold indistinct, evident as thoracic fold; dorsolateral fold weak or evident; (2) tympanic membrane and tympanic annulus present, distinct; (3) snout rounded in dorsal and lateral views; (4) upper eyelid bearing small, low, round tubercles; upper eyelid width slightly narrower than IOD; cranial crests absent; (5) dentigerous processes of vomers low, indistinct, separated,

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concealed by palatal mucosa; (6) vocal slits and spinous nuptial pads present; (7) Finger I shorter than Finger II; discs on outer fingers barely expanded, rounded to weakly truncate; (8) fingers lacking lateral fringes; (9) ulnar tubercles present, coalesced into a low fold; (10) heel and outer edge of tarsus bearing small tubercles; inner tarsal fold present; (11) inner metatarsal tubercle prominent, elliptical, two times larger than the lower, oval outer metatarsal tubercle; plantar surface areolate; (12) toes lacking lateral fringes; webbing absent; Toe V slightly longer than Toe III; toe discs barely expanded, round, weakly truncate, nearly as large as those on fingers; (13) dorsum entirely reddish ochre, brown or reddish brown with or without markings, such as interorbital bar or longitudinal dark stripes (especially in males); groin, posterior surface of thigh, concealed surface of shank, and dorsal surface of foot colored like dorsum but with white blotches; ventral surfaces pearly white, cream, or orange-ochre with dark brown or black scattered blotches; (14) SVL 16.0–19.4 mm ( x = 17.9; n = 5) in adults males, 23.7–27.8 mm in adult females ( x = 25.5; n = 3).

FIGURE 1. (A, B) Hand and foot of Pristimantis mariaelenae. (C, D) Hand and foot of Pristimantis stipa. Scale bars = 3 mm. Photos by L. Alza.

From the fifteen previously known species of the Pristimantis orestes Group (i.e., P. atrabracus, P. chimu, P. cordovae, P. corrugatus, P. melanogaster, P. orestes, P. pataikos, P. pinguis, P. seorsus, P. simonbolivari, P. simonsii, P. stictoboubonus, P. vetriguttatus, P. vidua; sensu Hedges et al. 2008 and P. bambu assigned to the group by Arteaga-Navarro & Guayasamin 2011), P. mariaelenae is easily distinguished from P. atrabracus, P. melanogaster, P. pataikos, P. pinguis, and P. stictoboubonus by having tubercles on the upper eyelid, heel, and outer edge of the tarsus (absent in the aforementioned species). Furthermore, P. atrabracus differs by having the ventral surfaces of hind limbs black, whereas in P. mariaelenae these are pearly white, cream, or orange-ochre; Pristimantis melanogaster differs by having the skin on dorsum nearly pustular (shagreen with scattered small tubercles in the new species), venter coarsely areolate (areolate), vocal slits and nuptial pads absent (present), and the groin (in life) with contrasting bright yellow spots (with white blotches); P. pataikos differs by having the skin of the dorsum smooth (shagreen with scattered small tubercles), venter coarsely areolate (areolate), and tympanum indistinct (distinct); P. stictoboubonus differs by males lacking vocal slits and nuptial excrescences (present), and fingers and toes bearing broad lateral fringes (lacking lateral fringes); P. pinguis differs by having the venter coarsely areolate (areolate), snout acutely rounded in dorsal view (rounded), dentigerous processes of vomers oblique, prominent, narrowly separated (low, indistinct, separated, concealed by palatal mucosa), and venter (in preservative) tan with brown flecks or reticulation (cream with scattered, brown blotches or dots). Pristimantis mariaelenae can be easily distinguished from P. simonsii and P. stipa sp.nov. by having discs barely expanded well defined by circumferential grooves, dorsum shagreen with scattered small tubercles, and venter areolate, whereas in the last two species, the discs are narrow lacking circumferential grooves, dorsum areolate, and venter coarsely areolate. Moreover, P. mariaelenae differs from P. simonsii (characters in parenthesis), by having the tympanic annulus and tympanic membrane distinct (absent) and from P. stipa by having the ventral surfaces pearly white, cream, or orange-ochre with

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FIGURE 2. Four living adult females of Pristimantis mariaelenae. (A, B) Dorsolateral and ventral views of holotype, CORBIDI 02824, 27.8 mm SVL. (C, D) Dorsal and lateral views of the paratype, CORBIDI 02828, 23.7 mm SVL. (E, F) Dorsal and lateral views of the paratype, CORBIDI 02827, 25.2 mm SVL. Photos by PJV.

dark brown or black scattered blotches (dark brown with white spots or mottling). Pristimantis mariaelenae can be distinguished from P. corrugatus and P. ventriguttatus by having discs barely expanded and fingers lacking lateral fringes (discs broadly expanded and fingers bearing lateral fringes in the two last species). Additionally, P. mariaelenae differs from P. corrugatus by having the skin on dorsum shagreen with scattered small tubercles (shagreen with irregular longitudinal ridges), and low tubercles on the upper eyelid (prominent). Pristimantis ventriguttatus differs from the new species by having the snout acutely rounded in dorsal view (rounded in P. mariaelenae), toes webbed basally (basal web absent), and the venter (in preservative) brown with tan blotches or dots (venter cream with brown blotches). Pristimantis mariaelenae differs from P. chimu and P. seorsus by having discs barely expanded and lacking cranial crests (discs narrow and cranial crests present). Pristimantis mariaelenae differs from P. cordovae by having the snout rounded in dorsal view lacking rostral tubercle or papillae (snout slightly pointed bearing small rostral papillae), and discs round (emarginate). In addition, P. mariaelenae can be distinguished from P. cordovae (in life) by having groin, posterior surface of thighs, and concealed surface of shanks brown or reddish

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brown with white blotches (brown with orange spots), and venter pearly white, cream, or orange-ochre with dark brown or black scattered blotches (ventral surface tan with small gray spots). Pristimantis bambu differs from P. mariaelenae by having the snout acuminate in dorsal view and lacking heel or tarsal tubercles (snout rounded and heel and tarsus bearing tubercles in P. mariaelenae). Pristimantis mariaeleneae differs P. orestes, P. simonbolivari, and P. vidua by having dorsolateral folds weak or evident (absent in the three last species). Furthermore, P. orestes and P. simonbolivari can be distinguished from P. mariaelenae by having (in life) the groin and concealed surfaces of the shanks black with white spots, whereas in the new species these surfaces are reddish ochre, brown or reddish brown with white blotches. Pristimantis vidua differs from P. mariaelenae (characters in parenthesis) by having indistinct tubercles on the eyelids (distinct), narrows discs on fingers and toes (barely expanded), and heel tubercles absent (present).

FIGURE 3. Preserved Pristimantis mariaelenae (holotype) in dorsal (left), and ventral (rigth) views. Photos by PJV.

Apart of Pristimantis orestes and P. simonbolivari (compared above) only three other species of Pristimantis in the Andes of Peru and Ecuador have contrasting white blotches in the groin (i.e. P. caeruleonotus, P. phalaroinguinis, and P. leucorrhinus). These three species differ from P. mariaelenae by having broadly expanded discs on fingers and toes. Lynchius flavomaculatus, known from southern Ecuador and extreme northern Peru, also has pale spots (cream or pale yellow in life) in the groin and on the hidden surfaces of the thighs, but it differs from P. mariaelenae by having smooth skin on the dorsum and venter, Finger I longer than Finger II, and cranial crests. Description of the holotype. Adult female with robust body; head narrow, not as wide as body, wider than long; head width 36.6% of SVL; head length 32% of SVL; snout short, lacking terminal tubercle, rounded in dorsal view, rounded in lateral view; eye–nostril distance 77.4% of eye diameter; nostrils rounded, directed dorsolaterally; canthus rostralis curved in dorsal view, straight in profile; loreal region concave; lips rounded; upper eyelid bearing low diffuse tubercles; upper eyelid width 75.8% of IOD; tympanic annulus present, distinct, with posterodorsal and posteroventral margins obscured by supratympanic fold; tympanic membrane present, distinct; tympanum diameter 58% of eye diameter, tympanum–eye distance 120% of tympanum diameter; one enlarged postrictal tubercle. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous processes of vomers low, indistinct, broadly separated, concealed by palatal mucosa, situated posteromedial to choanae; each vomer bearing two distinct teeth; tongue twice as long as wide, notched behind, free posteriorly for two thirds of its length. Skin on dorsum shagreen with small tubercles, most dense posteriorly; dorsolateral fold weak, discontinuous posteriorly; skin on flanks tuberculate; skin on, belly, chest, throat, and ventral surfaces of thighs areolate; discoidal

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fold indistinct, evident only as thoracic fold; cloacal sheath short; skin in cloacal region tuberculate. Ulnar tubercles coalesced into low fold; palmar tubercles slightly elevated, outer palmar tubercle bifid, approximately twice size of ovoid, thenar tubercle; subarticular tubercles well defined, round in ventral view and round in lateral view; supernumerary tubercle at base of fingers present; fingers lacking lateral fringes; Finger I shorter than Finger II; discs on Fingers I and II narrow, barely expanded on Fingers III and IV; discs round weakly truncate; ventral pads on fingers well defined by circumferential grooves on all fingers (Fig. 1A).

FIGURE 4. Five male paratypes of Pristimantis mariaelenae in life. (A, B) Dorsolateral and ventral views of CORBIDI 02826, 19.4 mm SVL. (C–E) Dorsolateral, lateral, and ventral views of CORBIDI 02823, 17.3 mm SVL. (F, G) Dorsolateral and ventral views of CORBIDI 02831, 16 mm SVL. (H–J) Dorsolateral, lateral, and ventral views of CORBIDI 02830, 14.9 mm SVL. (K, L) Dorsolateral and ventral views of CORBIDI 02832, 13 mm SVL. Photos by PJV.

Hind limbs slender, tibia length 37.4% of SVL; foot length 39.5% of SVL; upper surfaces of hind limbs tuberculate; posterior and ventral surfaces of thighs areolate; heel bearing one low tubercle; outer surface of tarsus bearing low tubercles; short inner tarsal fold present; inner metatarsal tubercle prominent, elliptical, rounded, three times size of low, oval, rounded outer metatarsal tubercle; plantar surface areolate; subarticular tubercles well

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defined, round in ventral view and subconical in lateral view; toes lacking lateral fringes; webbing between toes absent; discs nearly as large as those on fingers, most prominent on Toe IV; discs round, barely expanded, weakly truncate; Toes III, IV, and V having ventral pads well defined by circumferential grooves, less distinct on Toes I and II; relative lengths of toes: 1< 2 5< 4 (Fig. 1B); Toe V slightly longer than Toe III (disc on Toe III not reaching distal subarticular tubercle on Toe IV, tip of the disc on Toe V not reaching the distal border of distal subarticular tubercle on Toe IV). Measurements of the holotype (in mm): SVL 27.8; tibia length 10.4; foot length 11; head length 8.9; head width 10.2; eye diameter 3.1; tympanum diameter 1.8; interorbital distance 2.9; upper eyelid width 2.2; internarial distance 2.4; eye–nostril distance 2.4. Color of holotype in life: The dorsum was entirely reddish ochre (Fig. 2A); the lips, canthus rostralis, and the supratympanic fold were orange-ochre. The groin, posterior surfaces of the thighs, concealed surfaces of the shanks, and dorsal surfaces of the feet were red-ochre with irregular white blotches; the throat, chest, belly, and ventral surfaces of the thighs were pearly white with orange-ochre mottling on the throat and scattered dark brown spots on the belly (Fig. 2B). The palmar and plantar surfaces were orange-ochre, and the iris was dark copper with fine brown reticulations. Color of holotype in preservative (Fig. 3): In ethanol, the dorsum is entirely dark brown; the lips, canthus rostralis, and the supratympanic fold are cream. The groin, posterior surfaces of the thighs, concealed surfaces of the shanks, and the dorsal surfaces of the feet are dark brown with irregular cream blotches. The throat, chest, belly, and, palmar and plantar surfaces are cream with scattered, irregular brown blotches on the belly. TABLA 1. Variation of measurements (in mm) and proportions of the type series of Pristimantis mariaelenae and P. stipa. See text for abbreviations. Pristimantis mariaelenae

Pristimantis stipa

Character

Males (n = 5)

Females (n = 3)

Males (n = 4)

Female (n = 1)

SVL

16–19.4 (17.9 ± 1.4)

23.7–27.8 (25.6 ± 2.7)

15.6–24.4 (18.8 ±3.9)

35.1

TL

6.9–8.1 (7.5 ± 0.4)

10.3–10.4 (10.6 ± 0.5)

5.9–9 (7.5 ± 1.4)

12

FL

6.9–8.5 (7.7 ± 0.6)

8.3–11 (10.6 ± 1.5)

6.5–9.3 (8.2±1.6)

13.9

HL

5.5–6.1 (5.7 ± 0.2)

7.1–8.9 (7.8 ± 0.9)

5.6–8 (6.6 ± 0.9)

10.7

HW

5.7–7.1 (6.3 ± 0.6)

8.7–10.2 (9.4 ± 0.7)

5.8–9.4 (7.7 ± 1.5)

13.2

ED

2.3–2.8 (2.5 ± 0.2)

2.7–3.5 (3.1 ± 0.4)

1.8–2.6 (2.2 ± 0.3)

3.9

IOD

2.1–2.6 (2.4± 0.2)

2.6–2.9 (2.7 ± 0.5)

1.8–2.7 (2.5 ± 0.4)

3.1

EW

1.3–1.7 (1.5 ±0.7)

1.5–2.2 (1.9 ± 0.7)

1.4–3 (1.9 ± 0.7)

3

IND

1.4–2.1 (1.6 ± 0.2)

1.7–2.4 (2 ± 0.6)

1.6–2.2 (1.9 ± 0.9)

2.9

E-N

1.7–1.9 (1.8 ±0.8)

2.2–2.4 (2.6 ± 0.1)

1.3–2.2 (1.7 ± 0.6)

2.7

TY

0.5–1.3 (0.9 ± 0.2)

1.1–1.8 (1.6 ± 0.5)

0.4–1 (0.6 ± 0.5)

1.4

TL/SVL

0.3–0.4 (0.4 ± 0.2)

0.7–0.4 (0.4 ± 0.3)

0.6–0.4 (0.9 ± 0.2)

0.4

FL/SVL

0.4–0.4 (0.4 ± 0.2)

0.5–0.4 (0.9 ± 0.4)

0.8–0.6 (0.4 ± 0.3)

0.4

HL/SVL

0.3–0.3 (0.3 ± 0.1)

0.8–0.3 (0.3 ± 0.2)

0.3–0.8 (0.5 ± 0.2)

0.3

HW/SVL

0.3–0.3 (0.3 ± 0.1)

0.6–0.7 (0.7 ± 0.1)

0.7–0.4 (0.9 ± 0.1)

0.8

HW/HL

1.3–1.6 (1.8 ± 0.6)

1.5–1.3 (1.2 ±0.9)

1.3–1.7 (1.1 ± 0.7)

1.3

E-N/ED

0.6–0.8 (0.7 ± 0.7)

0.6–0.8 (0.7 ± 0.1)

0.9–1 (0.8 ±0.2)

0.9

EW/IOD

0.5–0.7 (0.6 ± 0.9)

0.5–0.8 (0.7 ± 0.5)

0.7–1.1 (0.9 ± 0.5)

0.9

TY/ED

0.2–0.4 (0.3 ± 0.1)

0.4–0.8 (0.7 ± 0.1)

0.8–0.4 (0.9 ±0.1)

0.6

Variation. Males are smaller than females, and the sexes differ slightly in some proportions (Table 1). Males have vocal slits and tan nuptial pads on the dorsal and medial surfaces of the thumb. In respect to coloration and skin texture, Pristimantis mariaelenae is highly variable: CORBIDI 02828 (Fig. 2: C & D), an adult female, has a reddish brown dorsum in contrast to nearly black flanks and a well defined dorsolateral fold; CORBIDI 02827 (Fig. 2: E & F), an adult female, has the same coloration as the holotype except for an orange-ochre dot medially in the TWO NEW SYNTOPIC SPECIES OF PRISTIMANTIS

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FIGURE 5. Distribution of Pristimantis mariaelenae and P. stipa in Peru. The red dot indicates the type locality of both species, the only locality where these species are known to date. White circles are major cities for reference.

scapular region, but the ventral surfaces are orange-ochre instead of pearly white. The adult male paratypes CORBIDI 02825–26 & 29 are identical to the holotype, except for a cream middorsal stripe in the CORBIDI 02826 (Fig. 4: A & B) and the presence of dorsal longitudinal folds in the paratype 02825. Four male paratypes exhibit the

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major variation in color and skin texture. Adult specimen CORBIDI 02823 (Fig. 4: C–E) has a dark brown dorsum with a greenish interorbital stripe and dorsolateral stripes, a thin yellow middorsal stripe, a red blotch on the anterior surface of thighs and axilla, a wrinkled dorsum, and a well defined dorsolateral fold; adult specimen CORBIDI 02831 (Fig. 4: F & G) has a brown dorsum with a dark brown middorsal stripe and well defined longitudinal folds; two juvenile males CORBIDI 02830 (Fig. 4: H–J) and 02832 (Fig. 4: K & L) have a distinct tuberculate dorsum and distinct sinusoidal ridges. Furthermore, these juveniles, CORBIDI 02830 and 02832, have a greenish dorsum. Distribution and ecology. Pristimantis mariaelenae is known only from the type locality in a humid puna above tree line, at an elevation of 3596 m.a.s.l., on the eastern slope of the Cordillera Occidental in northwestern Peru. The type locality is a grassy region dominated by bunch grass (Stipa) with associated small bushes (Baccharis sp.), and some scattered small patches of elfin forest (Fig. 5). All specimens were in the bases of bunch grass and also under rocks near small streams by day. Etymology. The specific name is a patronym for Maria Elena Venegas formerly from Lima, Peru, mother of the senior author. The new species is dedicated to Maria Elena for her continuous support of PJV’s love of nature.

Pristimantis stipa new species Figs. 1(C & D), 6(A–D), 7 Holotype: CORBIDI 02833, an adult female, from Cañaris, Provincia de Ferreñafe, Región Lambayeque, Perú (6˚7'14.4" S, 79˚18'4.43" W; 3596 m.a.s.l.) obtained on 16 June 2007 by Pablo J. Venegas. Paratypes: CORBIDI 02834–37, four adult males, collected with the holotype by Pablo J. Venegas. Diagnosis. This member of the Pristimantis (Pristimantis) orestes Group has: (1) skin on dorsum areolate; that on flanks and venter coarsely areolate; discoidal fold only present as a prominent thoracic fold; dorsolateral fold prominent, discontinuous; (2) tympanic annulus present, weakly defined; tympanic membrane distinct; (3) snout bluntly rounded in dorsal view and in lateral view; (4) upper eyelid tubercles absent; upper eyelid nearly as wide as, or equal to IOD; cranial crests absent; (5) dentigerous processes of vomers prominent, oblique, narrowly separated; (6) vocal slits absent; spinous nuptial pads absent; (7) Finger I shorter than Finger II; discs on fingers narrow, rounded, without marginal grooves; (8) lateral fringes on fingers absent; (9) ulnar tubercles present, coalesced into a low fold; (10) heel and outer edge of tarsus lacking tubercles; inner tarsal fold present; (11) inner metatarsal tubercle prominent, elevated, elliptical, twice as large as lower oval outer metatarsal tubercle; plantar surface areolate; (12) lateral fringes on toes and webbing absent; Toe V slightly shorter than Toe III; toe discs narrow, rounded, without marginal grooves; (13) dorsum entirely brown without markings; groin, posterior surface of thigh, and concealed surfaces of shank and foot dark brown with or without white marks; ventral surfaces dark brown with white spots or mottling; (14) SVL in adult males 15.6–24.4 mm ( x = 18.8; n = 4), in one adults female 35.1 mm. Pristimantis stipa differs from all other species of Pristimantis, including the 15 species currently assigned to the P. orestes Group, except P. simonsii, by the following combination of characters: dorsum areolate; discs on fingers and toes narrow without discernable marginal grooves; and ulnar tubercles coalesced into low fold. However, it is easily distinguished from P. simonsii by having dentigerous processes of vomers prominent (absent in P. simonsii); tympanic annulus and membrane distinct (absent); and Toe V slightly shorter than Toe III (Toes III and V about equal in length). In lacking well-defined marginal grooves on the discs on the fingers and toes, P. stipa can be confused with Phrynopus. The only species of Phrynopus known from the Cordillera Occidental is P. thompsoni from farther south in Región La Libertad. This species of Phrynopus differs from Pristimantis stipa by having longitudinal rows of pustules on the dorsum, tympanic annulus and membrane absent, Fingers I and II equal in length, and venter tan with brown spots. Description of the holotype. Adult female with robust body; head narrow, not as wide than body, wider than long; head width 37.6% of SVL; head length 30.4% of SVL; snout short lacking terminal tubercle, bluntly rounded in dorsal and lateral views; eye–nostril distance 69.2% of eye diameter; nostrils rounded, directed dorsolaterally; canthus rostralis straight in dorsal view, slightly curved in profile; loreal region concave; lips rounded; upper eyelid lacking tubercles; upper eyelid width 96.7% of IOD; tympanic annulus present, weakly defined, with the posterodorsal margin obscured by supratympanic fold, tympanum diameter 35.8% of eye diameter, tympanum–eye distance about 1.5 times tympanum diameter; one enlarged postrictal tubercle on left side and two different-sized postrictal tubercles on right side. Choanae small, ovoid, not concealed by palatal shelf of maxilla; dentigerous pro-

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cesses of vomers prominent, oblique, narrowly separated, situated posteromedial to choanae, each bearing three teeth; tongue twice as long as wide, notched behind, free posteriorly for two thirds of its length.

FIGURE 6. Living adults of Pristimantis stipa. (A, B) Dorsolateral and ventral views of the female holotype, CORBIDI 2833, 35.1 mm SVL. (C, D) Dorsolateral and lateral views of a male paratype, CORBIDI 2834, 24.4 mm SVL. Photos by PJV.

Skin on dorsum areolate; dorsolateral fold prominent, fleshy, and discontinuous; skin on flanks, ventral surfaces of thighs, belly, chest, and throat coarsely areolate; discoidal fold only present as prominent thoracic fold; cloacal sheath short; cloacal region tuberculate. Ulnar tubercles coalesced into low fold; palmar tubercles fleshy, elevated, outer palmar tubercle bifid, approximately twice size of ovoid, thenar tubercle; distal subarticular tubercles weakly defined, except proximally, round in ventral and lateral views; supernumerary tubercle sat bases of Fingers II and III distinct; fingers lacking lateral fringes; Finger I shorter than Finger II; discs on fingers narrow with rounded tips; pads on fingers lacking circumferential grooves (Fig. 1C). Hind limbs slender, tibia length 34.1% of SVL; foot length 39.6% of SVL; upper surfaces of hind limbs areolate; posterior and ventral surfaces of thighs areolate; heel lacking tubercles; outer surface of tarsus lacking tubercles; short inner tarsal fold present; inner metatarsal tubercle prominent, elliptical, twice size of low, oval outer metatarsal tubercle; plantar surface areolate; subarticular tubercles well defined, round in ventral and lateral views; toes lacking lateral fringes; webbing absent; discs on toes narrow with rounded tips, like those on fingers; toes lacking circumferential grooves; relative lengths of toes: 1< 25< 4 (Fig. 1D); Toe III slightly longer than Toe V (disc on Toe V reaching to third subarticular tubercle on Toe IV, tip of the disc on Toe III extending to distal edge of distal subarticular tubercle on Toe IV). Measurements of the holotype (in mm): SVL 35.1; tibia length 12.0; foot length 13.9; head length 10.7; head width 13.2; eye diameter 3.9; tympanum diameter 1.4; interorbital distance 3.1; upper eyelid width 3.0; internarial distance 2.9; eye–nostril distance 2.7. Color of holotype in life: The dorsum was brown (Fig. 6A); the groin, posterior and anterior surface of thighs, concealed surface of shanks and dorsal surface of feet were blackish brown with irregular dirty white blotches, and the venter dark brown with white spots (Fig. 6B). The iris was grayish white with brown mottling. Color of holotype in preservative (Fig. 7): In ethanol, the coloration is as described above, except that the blackish brown is brown, and the dirty white blotches are white; iris greenish is gray with a fine brown mottling.

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FIGURE 7. Preserved Pristimantis stipa (holotype) in dorsal (left), and ventral (rigth) views. Photos by PJV.

Variation. The morphology and coloration of the paratypes are identical to the holotype (Fig. 6: C & D). Males are smaller than females, lack vocal slits, and have white nuptial pads on the dorsal and medial surfaces of thumb. The mottling on the throats of males is brownish cream (Fig. 6D). See Table 1 for measurements and proportions of the type series. Distribution and natural history. Pristimantis stipa is known only from the type locality in a humid puna above tree line, at an elevation of 3596 m.a.s.l., on the eastern slope of the Cordillera Occidental in northwestern Peru (Fig. 5). There it occurs syntopically with P. mariaelenae. Etymology. The specific name is a noun in apposition. It is the generic name of the feather grass (Pooideae) that is common at the type locality and in most areas above tree line and below snow line in Peru. Remarks. The type locality, Cañaris, has been poorly explored herpetologically. It is located in the northeastern corner of the Región Lambayeque at the right margin of the Río Huancabamba (2500–3800 m.a.s.l.). The general landscape is a humid puna with scattered patches of elfin forest and cloud forest partially cleared for cattle ranching and cropland. The destruction of the natural habitats in Cañaris by the local people has been continuous since the 1970s; however, the general area is currently under new human pressure because of mineral exploration. Other surveys by the senior author in Cañaris and the nearby locality of Incahuasi recorded more species of strabomantid frogs, including Lynchius sp., P. chimu, P. petrobardus, P. phoxocephalus, and three species of Prisitmantis. Inasmuch as the unidentified species of strabomantids might be new species, probably all of the recorded species, except for P. phoxocephalus, are restricted to the northern portion of the Cordillera Occidental in Peru. It is well known that most members of the Pristimantis orestes Group have limited distributional ranges (Lehr 2007). Although some large patches of cloud forest still remain, the humid puna is strongly overgrazed by cattle and is frequently burned (Fig. 8); a high probability of future mining activity is a significant new potential threat.

Discussion The Pristimantis orestes group was originally defined by Lynch & Duellman (1997) with three terrestrial species (e.g. P. orestes, P. simonbolivari, and P. vidua) from the Andes of southern Ecuador (Lynch 1979; Wiens & Coloma TWO NEW SYNTOPIC SPECIES OF PRISTIMANTIS

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1992; Lynch & Duellman 1997). The three original species of the Pristimantis orestes Group are structurally similar (Wiens & Coloma 1992) and resemble species of the Pristimantis myersi Group in size, robustness, general proportions, relative lengths of Finger I and II, and sizes of the digital discs (Lynch & Duellman 1997; Hedges et al. 2008). Within the last decade, several new species that have been discovered in the Andes in northern Peru have been assigned to the Pristimantis orestes Group (Duellman & Pramuk 1999; Duellman et al. 2006; Lehr & Duellman 2007; Lehr & Kohler 2007; Lehr 2007; Arteaga-Navarro & Guayasamin, 2011). The osteological characters of the Pristimantis orestes Group remain poorly known. Lynch & Duellman (1997) mentioned that the frontoparietals are fused with the prootics in Pristimantis orestes, P. simonbolivari, and P. vidua, in contrast to the plesiomorphic condition of unfused elements in most species of the large and certainly paraphyletic Pristimantis unistrigatus Group (Hedges et al., 2008). However, this character has not been studied in the Peruvian species of the Pristimantis orestes Group, which has been defined entirely by external morphological characters. Increased taxon sampling is needed for molecular analyses of these frogs; until such is the case, the Pristimantis orestes Group must be recognized as a phenetic group of small terrestrial frogs with short limbs and robust bodies, features that also characterize the Pristimantis myersi Group in the northern Andes. According with Duellman & Lehr (2009), the major threat to strabomantid frogs is habitat destruction through deforestation or agriculture (crop and cattle) and environmental contamination. We cannot state precisely the IUCN Red List status of the species described herein. They might be designated as critically endangered because of the continuing decline in the extent and quality of their habitat at the type locality and their probable restricted ranges (less than 80 km²); both new species are under the same threats as the Critically Endangered (IUCN Red List; Icochea et al. 2004) P. simonsii (also a member of the P. orestes Group) that inhabits the humid puna in neighboring Cajamarca.

FIGURE 8. Habitat at the type locality of Pristimantis mariaelenae and P. stipa on May 2007. The bunch grass is Stipa sp. Photo by PJV.

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Acknowledgments We are indebted with Jesús Córdova and Claudia Torres for allowing access to the herpetological collection of the MUSM. We are grateful with A. Fernandez and K. Garcia-Burneo for arranging the plates. Collecting permits were issued by the Instituto Nacional de Recursos Naturales (110-2007-INRENA-IFFS-DCB), Lima, Peru.

References AmphibiaWeb. (2011) Information on amphibian biology and conservation. [web application]. 2011. Berkeley, California.: AmphibiaWeb. Available:Available from http://amphibiaweb.org/. (Accessed: January 2012). Arteaga-Navarro, A.F. & Guayasamin, J.M. (2011) A new frog of the genus Pristimantis (Amphibia: Strabomantidae) from the high Andes of southeastern Ecuador, discovered using morphological and molecular data. Zootaxa, 2876, 17–29. Boulenger, G. A. 1900. Descriptions of new batrachians and reptiles collected by Mr. P. O. Simons in Peru. Annals and Magazine of Natural History, Series 7, 6: 181–186. Duellman, W.E. & Pramuk, J.B. (1999) Frogs of the genus Eleutherodactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers of the Natural History Museum, University of Kansas, 13, 1–78. Duellman, W.E., Lehr, E. & Venegas, P.J. (2006) Two new species of Eleutherodactylus (Anura: Leptodactylidae) from northern Peru. Zootaxa, 1285, 51–64. Duellman, W.E. & Lehr, E. (2009) Terrestrial-breeding frogs (Strabomantidae) in Peru. Munster, Germany: Nature und Tier Verlag, 382 pp. Hedges, B.S., Duellman, W.E. & Heinicke, M.P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1–182. Icochea, J., Lehr, E., & Puntriano, C. A. (2004) Pristimantis simonsii. In: IUCN 2010. IUCN Red List of Threatened Species. Version 2010.4. . Downloaded on 02 May 2011. Lehr, E. (2007) Seven new species of Pristimantis (Anura: Leptodactylidae) from Peru. Bulletin Museum of Comparative Zoology, 159, 145–178. Lehr, E. & Duellman W.E. (2007) Two new species of Eleutherodactylus (Anura: Leptodactylidae) from the Cordillera Occidental in Peru. Copeia, 2007, 140–149. Lehr, E. & Kohler, G. (2007) A new species of the Pristimantis orestes Group (Anura: Leptodactylidae) from the Cordillera Occidental in northern Peru. Zootaxa, 1621, 45–54. Lehr, E., Moravec, J. & Gagliardi-Urrutia, L.A.G. (2010) A new species of Pristimantis (Anura: Strabomantidae) from the Amazonian lowlands of northern Peru. Salamandra, 46, 197–203. Lynch, J.D. (1979) Leptodactylid frogs of the genus Eleutherodactylus from Andes of southern Ecuador. Miscellaneous Publications of the Museum of Natural History, University of Kansas, 66, 1–62. Lynch, J.D. (1980) A taxonomic and distributional synopsis of the Amazonian frogs of the genus Eleutherodactylus. American Museum Novitates 2696, 1–24. Lynch, J.D. & Duellman, W.E. (1997) Frogs of the genus Eleutherodactylus in western Ecuador: systematics, ecology, and biogeography. Special Publications, Natural History Museum University of Kansas, 23, 1–236. Wiens, J.J. & Coloma, L.A. (1992) A new species of Eleutherodactylus myersi (Anura: Leptodactylidae) assemblage from Ecuador. Journal of Herpetology, 26(2), 196–207.

APPENDIX. Specimens examined. Pristimantis chimu: PERU: Cajamarca: Bosque de Protección de Pagaibamba, 3000 m: CORBIDI 02838, CORBIDI 01598. Pristimantis cordovae: PERU: La Libertad: ca. 8 km NE Quiruvilca, 3542 m: MHNSM 21990 (holotype), MUSM 21991, 21998, 21999 (paratopotypes). Pristimantis corrugatus: PERU: San Martín: Ullilen, 3000 m: MUSM 28063 (holotype); Quintecocha, 3130 m: MUSM 28062, 28064–67 (paratypes); Laurel, 2830 m: CORBIDI 00543–46, 00599–601, 00617. Pristimantis melanogaster: PERU: Amazonas: Área de Conservación Privada Huiquilla, 2857 m: CORBIDI 00384-85. Pristimantis pataikos: PERU: Amazonas: Yuramarca, 2864 m: CORBIDI 00496-97. Pristimantis pinguis: PERU: Cajamarca: Hualgayoc, Cerro Las Gordas MUSM 27021; 23 Km (by road) SW Celendín: KU 181283 (holotype); Celendín, El Punrei (82°35'20.6" W; S 59°43'20.9" W, 3772 m): MUSM 27313–15. Pristimantis simonsii: PERU: Cajamarca: Celendín, 3600–4000: MUSM 27306, 27304, 22516, 22540, 26188, 26187, 26185, 26184, 26186, 26183, 26173. Pristimantis stictoboubonus: PERU: San Martín: Quintecocha, 3130 m: MUSM 24446 (holotype), MUSM 24445 (paratype); Ullilen, 3000 m: MUSM 24447 (paratype); Leimebamba–Los Chilchos trail, 2994 m: CORBIDI 00611.

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