Ameerega pepperi Brown & Twomey, 2009, sp. nov.

Ameerega pepperi sp. nov. Figures 1, 14–16 Phyllobates bassleri: Silverstone 1976 p. 45 (AMNH 42327, 42867, 43402 collected in 1926 and 1928 by H. Bassler at “Pachiza”). Holotype. MUSM 26940 (Fig. 16), an adult male collected on 14 July 2006 by M. Pepper, Provincia Tocache, Departamento San Martín, Peru, 2 km NE of San Francisco, 980 m elevation, 8° 18' 30.3" S, 76° 40' 37.6" W. Found on the ground near a small waterfall. Paratypes. All from San Martín, Peru. One adult female (MUSM 26942), collected 21 July 2006 by M. Pepper and E. Twomey, and two adult females (MUSM 26975, 26976), collected 25 June 2007 by J. Brown, E. Twomey, and K. Fieselman, 390 m elevation, 7.0 km SW of Huicungo at 7° 21' 58.6" S, 76° 48' 44.1" W: Two adult females (MUSM 26979, 26980), collected on 25 June by J. Brown, E. Twomey, and K. Fieselman, 384 elevation, 6.6 km N of Campanilla at 7° 25' 38.6" S, 76° 39' 53.3" W: One adult female (MUSM 26968), collected on 15 July 2006 by M. Pepper, 570 m elevation, SW of Chumanza (alternate spelling: Shumanza) at 7° 34' 7.8" S, 76° 41' 34.4" W: One adult male (MUSM 26941), collected at same locality of holotype, 980 m elevation, 2 km NE of San Francisco at 8° 18' 30.3" S, 76° 40' 37.6" W. Etymology. The specific epithet is a patronym for the discoverer of this species, Mark Pepper, an enthusiastic Canadian explorer and conservationist whose discoveries have greatly contributed to the authors’ understanding of poison frog biogeography and taxonomy. Definition and diagnosis. Assigned to the genus Ameerega on the basis of the following: first finger longer than second, webbing absent between the toes, dorsal skin granular (Myers 1982, Grant et al. 2006). This is a medium-large species of Ameerega with an adult SVL of approximately 28–34 mm. Dorsum has a black ground color, finely stippled brick-red to bright red in southern populations (e.g. San Francisco) to coarsely stippled orange or yellow northern populations (e.g. Campanilla), stippling concentrated near head and is less dense near sacral region. Bright yellow dorsolateral stripes extend from the eyelid to the groin; a yellow labial stripe is also present starting near nares and extending posterior to the axillary region. Faded yellow-green to white spot present above groin in some populations, shank spots and axillary spots absent. Limbs greenish-bronze or black; venter blue with strong black marbling. Teeth present. Appressed first finger longer than second; finger discs moderately expanded, hands and feet lacking webbing between digits. Vocalization consists of a loud retarded trill of whistle-like notes given at a rate of 0.9–1.3 notes per second; dominant frequency 2800–3160 Hz (Fig. 4, Table 2). Ameerega pepperi can be distinguished from most other species of Ameerega by its red, orange, or yellow dorsum. It is similar in appearance to the following species: A. bassleri, A. bilinguis, A. cainarachi, A. macero, A. parvula, and A. yoshina, all of which have (or can have) a solid red, orange, or yellow dorsum. Ameerega parvula and A. bilinguis both lack distinct dorsolateral stripes (vs. dorsolateral stripes present and conspicuous in A. pepperi). Ameerega cainarachi is smaller, females up to 31.3 mm SVL (vs. 34 mm SVL in A. pepperi), and has an advertisement call consisting of a regular chain of notes which are not frequency modulated, repeated at a rate of 9 notes per second (vs. ‘bursts’ of frequency-modulated notes, repeated 0.9–1.3 notes per second in A. pepperi). Ameerega macero is smaller than A. pepperi, females up to 29.5 mm SVL, and has a call consisting of a series of harsh ‘peeps’, repeated regularly at a rate of 10 notes per second. Ameerega bassleri has an advertisement call consisting of a series of regularly-spaced whistle-like notes given continuously for several minutes at a rate of 1.6–2.1 notes per second (vs. notes given at a rate of 0.9–1.3 notes per second in A. pepperi). Ameerega yoshina has an advertisement call given in bursts of 4–31 notes, given at a rate of 3.7–4.1 notes per second, each burst less than 8 seconds in total duration (vs. notes given at a rate of 0.9–1.3 notes per second in A. pepperi). Measurements (in mm) of holotype. The male holotype (Fig. 16) has SVL 28.6; FL 13.4; TL 16.1; KK 28.2; FoL 13.3; HaL 7.5; HL 7.7; HW 7.8; BW 8.3; UEW 5.5; IOD 3.9; IND 4.0; TD 3.0; ED 4.2; DET 1.6; L1F 6.9; L2F 6.1; W3D 0.8; W3F 0.6. Measurements of paratypes are given in Table 5. Description of holotype. Size medium, SVL 28.6 mm (Table 5). Dorsal skin is coarsely granular and is pigmented black with brick red mottling densest near the snout. Thin yellow dorsolateral stripes extend from groin, extending dorsally around snout; posterior to eyes the stripes blend into the red dorsum. Dorsolateral stripe is sightly wider at groin. Flanks are black extending from groin to nares. Forelimbs lack granulation and are colored tan and flecked in olive. Yellow coloration extends up upper arm surface, anteriorly around lip. Legs are coarsely granulated; contain yellow femoral spots, dorsal surface colored olive green. The venter is yellow-gold with olive marbling; marbling extends primarily on throat and margins of legs. Iris black. Widest part of head between jaw articulations and tympanum, head narrower than body. Greatest head width 27.0 % of SVL. Eyes very protuberant. Tongue medium sized, oval. Premaxillary and maxillary teeth present. Vocal slits present. Dorsal skin of head and body is finely granular; skin smooth or nearly smooth on limbs and smooth on sides of head and body and ventral surfaces. Snout sloping laterally; bluntly rounded dorsally; truncate ventrally. Nares situated and directed posterolaterally to the tip of snout; nares visible from front and below but not from above. Canthus rostralis sloped, slightly rounded; loreal region nearly vertical and slightly concave. Upper eyelid 1.4 times wider than interorbital distance. Eye large and prominent with a maximum diameter of 14.6 % of SVL, pupil rounded and horizontally elliptical. Tympanum circular, without tympanic annulus its diameter less than 73.0 % of ED, supratympanic fold absent. Hands relatively small, length being 26.2 % of SVL. Relative length of appressed fingers I ≈ II ≈ IV 300 m), we have never observed A. pepperi greater than ~ 30 m from a stream. This species can occur in moderately disturbed areas provided the streamside habitats are relative

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