Pristimantis koehleri, Padial & Riva, 2009

Padial, José M. & Riva, Ignacio De La, 2009, Integrative taxonomy reveals cryptic Amazonian species of Pristimantis (Anura: Strabomantidae), Zoological Journal of the Linnean Society 155 (1), pp. 97-122 : 107-111

PRISTIMANTIS KOEHLERI SP. NOV. ( Fig. 7A )

Holotype: MNK-A 6626 (field number JMP 033 ), an adult male from km 6 of Angostura–Samaipata road, Departamento Santa Cruz, Bolivia (18°11′S, 63°34′W), collected by J. M. Padial, 03.i.2003.

Paratopotypes: MNCN 42990-1 , MNK-A 6627 (adult males, field numbers JMP 031–3 ) , same data as the holotype .

Paratypes: Bolivia, Departamento Santa Cruz : MNK-A 7170 (adult male, field number JMP 442 ), 7172 and 7174 (adult males, field numbers JMP 449 and 451), MNCN 43054 (adult male, field number JMP 448 ) from Espejillos (17°50′S / 63°25′W), collected by J. M. Padial and E. Ávila , 26.xi.2003; MNCN 42983 , 42985–6 , 43014 (subadult females, field numbers JMP 152 , 173 , 184 and 153, respectively) from La Chonta, Amboró National Park (17°39′36″S, 63°42′6.6″W) collected by J. M. Padial and R. de Sá , 21–22.iv.2003, and MNCN 43040 (adult male, field number JMP 377 ) collected at the same locality by J. M. Padial & E. Ávila , 05.xi.2003; ZFMK 80005–6 (adult males), ZFMK 80007 (adult female) from Macuñucú, Amboró National Park collected by J. Köhler and S. Lötters , 2.xii.1998; ZFMK 79991 and 79993 (adult females) and 79992 (juvenile) from Mataracú, Amboró National Park , collected by J. Köhler and S. Lötters , 14.i.1999. Referred specimens: BOLIVIA: Departamento Santa Cruz : Espejillos, MNK-A 6447; Km 29 on Santa Cruz de la Sierra – Samaipata road, MNK-A 1000; Río Saguayo, Amboró National Park , MNK-A 189, 191, 224, 358, 361, 364–5, 374; Río Surutú , Amboró National Park , MNK-A 1197; Santa Cruz de la Sierra , BM 1904.10.29.83–101 (general locality, the origin of these specimens is likely to be in the Andean slopes close to Santa Cruz de la Sierra) .

Diagnosis: A member of the Pristimantis conspicillatus Group, as defined by Lynch & Duellman (1997), characterized by: (1) skin on dorsum coarsely shagreen, flanks with larger granules; venter finely granular, smooth only in the middle; posterior surfaces of limbs smooth; discoidal fold conspicuous; dorsolateral folds absent; postrictal glands present; (2) tympanic membrane and annulus round, large, their length about half eye length; supratympanic fold short, very prominent; (3) head slightly longer than wide; snout acuminate in dorsal view, subacuminate in lateral view; canthus rostralis straight in dorsal view, sharp in profile; (4) cranial crests absent; upper eyelid covered by small granules; (5) vomerine odontophores large, situated posteromedial to choanae; (6) males with vocal slits and two nuptial pads on thumb; (7) hands with long and slender fingers, first finger longer than second; subarticular tubercles subconical, prominent; supernumerary tubercles round, smaller than subarticular tubercles; terminal discs of inner two fingers round, those of external fingers enlarged, ovate to truncate; circumferential grooves conspicuous, ungual flap not indented; lateral fringes and keels on fingers absent; (8) ulnar tubercles present; (9) tubercles on heel and tarsus absent, tarsal fold prominent; (10) inner metatarsal tubercle ovate, prominent, outer subconical, prominent; single, round supernumerary tubercle on Toe IV; (11) toes long and slender (foot length 50% SVL); lateral fringes or keels weak, basal toe webbing absent; fifth and third toes reaching midpoint of penultimate subarticular tubercle of Toe IV; tips of toes rounded to ovate, enlarged, ungual flap not indented; (12) dorsal coloration grey to brown with variable pattern of dark spots and flecks, ventral coloration white with fine mottling; posterior surface of thighs brown without light spots; (13) mandibular ramus of the trigeminal nerve passing lateral to the m. adductor mandibulae externus (S condition sensu Lynch, 1986).

This species is very similar to Pristimantis fenestratus ( Fig. 7B ), P. dundeei and P. samaipatae in qualitative characters ( Table 3). It differs, however, from these three species as follows. From P. dundeei by lacking warts on dorsal skin, by lacking basal toe webbing and by having less developed finger fringes. From P. fenestratus by lacking basal toe webbing and having smaller size ( SVL) of adult males (23.8–29.4 vs. 26.0–34.7) and females (34.0–39.5 vs. 37.8–57.2). From P. samaipatae , by having granular skin on belly (smooth), weak finger fringes, and smaller size ( SVL) of adult males (23.8–29.4 vs. 30.1–40.0) and females (34.0–39.5 vs. 44.4–51.4). For additional differences among these three species see Table 3 and results of morphological, bioacoustic and molecular analyses above. For differences of P. koehleri and other members of the group see Table 3.

Description of the holotype: Head longer than wide (head length/head width = 1.2); snout acuminate in dorsal view and subacuminate in lateral profile; nostrils slightly protuberant, orientated posterolaterally; canthus rostralis straight in dorsal view, sharp in frontal profile; loreal region flat; lips not flared; upper eyelid without tubercles but covered by small granules; no cranial crests. Supratympanic fold prominent, short; tympanic membrane and annulus distinct; tympanic membrane nearly round, its length about half of eye length; postrictal glands conical, conspicuous. Choanae not concealed by palatal shelf of the maxillary arch when roof of mouth is viewed from below; choanae large, ovate, separated by distance equal to five times diameter of a choana; vomerine odontophores large, prominent, round in shape, situated posteromedial to choanae, separated by a distance equal to the length of a vomerine odontophore, bearing 4–5 vomerine teeth; vocal sac subgular, vocal slits placed posterolaterally. Skin of dorsal surfaces and posterior parts of hind limbs coarsely shagreen; throat smooth, belly granular, only smooth in the middle; occipital folds absent; dorsolateral folds absent; discoidal fold conspicuous.

Arm with a row of low, round ulnar tubercles; palmar tubercle bifid, flat, equal to elongate, prominent, thenar tubercle; a single supernumerary tubercle on the basis of each finger, low, round, smaller than subarticular; subarticular tubercles prominent, subconical; finger tips small and round on fingers I and II, and large, ovate to truncate on fingers III and IV; fingers lacking lateral fringes; relative length of fingers III > I> II ± IV; a double white glandular, nonspinous nuptial pad on dorsal surface of each thumb.

Toes long and slender (foot length 50% of SVL); heel and tarsus lacking tubercles; tarsal fold prominent, almost in contact with inner metatarsal tubercle and larger than it; inner metatarsal tubercle ovate, prominent, larger than outer; outer metatarsal tubercle round, subconical; only a single inconspicuous supernumerary tubercle on Toe IV; subarticular tubercles conical, prominent; toes with weak lateral fringes; basal toe webbing absent; toe tips round, moderately developed; ungual flap not indented, circumferential grooves evident; relative length of toes IV > III > V > II > I; toes III and V reaching midpoint of penultimate subarticular tubercle of Toe IV.

Measurements (in mm) of the holotype: SVL 26.6, HL 10.7, HW 9.30, EL 4.0, EN 3.7, IND 2.4, EE 4.81, TYH 2.0, TYL 2.0, FIII 1.1, FIV 1.1, FA 4.9, TL 15.1, TH 12.5, FL 13.2, TIV 1.2.

Colour: In preservative, dorsal surfaces light greyishbrown with diffuse and inconspicuous light brown dorsal chevrons; a pair of bold black occipital spots; flanks light greyish-brown with some fine dark mottling; canthus rostralis dark brown; dorsal and loreal regions of snout grey; a fine bold brown interocular stripe; inconspicuous labial bars dark brown and white; subocular stripes absent; tympanic membrane brown, annulus cream; tympanic fold bold black; hind-limbs and arms with transversal dark stripes; plantar surfaces brown; ventral surfaces white to cream with inconspicuous fine greyish-brown mottling; posterior and anterior surfaces of hind limbs brown without spots. The colour pattern in life is very similar, but greyish surfaces tend to be light brown to cream. The ventral surfaces are white and the groin is yellowish-white. The iris is metallic orange with a transverse bold black stripe.

Variation: Males and females are similar in all but sexual qualitative external characters. Females are larger than males but are equal in head and limbs proportions ( Table 4). Dorsal pattern is quite constant, although some specimens, e.g. MNCN 42986, 43054 or MNK-A 7170, 7172, present a fine middorsal stripe from tip of snout to vent. Some dark dorsal marks, such as an interocular stripe, W-shaped occipital mark, X-shaped mid-dorsal mark or sacral chevrons, can be present. The tarsal fold can be poorly developed and rounded, as in MNCN 43054. For measurements, see Table 4.

Etymology: The name is a patronym for Jörn Köhler, German herpetologist and friend, whose studies have greatly contributed to the understanding of Bolivian amphibian diversity.

Distribution: This is a Bolivian endemic species known from the inter-Andean dry valleys of the Departamento de Santa Cruz extending to the northwest along the humid forests of the Andean slopes (see Fig. 8 ). De la Riva (1994) cited this species as Pristimantis peruvianus from Amboró National Park (Departamento Santa Cruz), Bulo Bulo and Valle de Sajta (Departamento Cochabamba). Köhler (2000a) cited this species as P. dundeei for Macuñucú and Mataracú, along the southern edge of Amboró National Park.

Natural history: This species is active by night during the rainy season. Males call from low vegetation of the forest. It has been found in both primary and secondary forest types.

Remarks: Köhler (2000a) described the advertisement call of this species (as Pristimantis dundeei ). His data for the calls are similar to those provided by us, although they differ by a longer note length reported by Köhler (2000a). Reichle’s (2002) recording for P. cf. peruvianus corresponds to P. koehleri . Specimens cited as P. peruvianus by De la Riva (1994) also correspond to P. koehleri as well as Bolivian specimens cited as P. cf. peruvianus by Padial & De la Riva (2005a). With the description of P. koehleri , P. dundeei would no longer occur in Bolivia, according to previous data. However, specimens from Noel Kempff Mercado National Park, Departamento Santa Cruz, Bolivia (see Appendix S1) represent the first country record of P. dundeei .

Figures

Figure 7. A, adult male of Pristimantis koehleri from Km 6 of Angostura–Samaipata road, Departamento Santa Cruz, Bolivia (one from the type series MNK-A 6626–7, MNCN 42990–1); B, adult female of Pristimantis fenestratus from Chalalán, Departamento La Paz, Bolivia (one from the series MNCN 43239–44).

Figure 8. Map of part of South America depicting the approximate distribution of Pristimantis danae, P. koehleri, P. fenestratus, P. reichlei, and P. samaipatae.