Oreobates granulosus, (BOULENGER, 1903), 2008
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publication ID |
https://doi.org/ 10.1111/j.1096-3642.2007.00372.x |
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persistent identifier |
https://treatment.plazi.org/id/03EF8790-FFC1-FFA4-75B3-739EFD6FF989 |
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treatment provided by |
Felipe |
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scientific name |
Oreobates granulosus |
| status |
comb. nov. |
OREOBATES GRANULOSUS ( BOULENGER, 1903) View in CoL
COMB. NOV.
Hylodes granulosus Boulenger, 1903
Eleutherodactylus granulosus View in CoL – Stejneger (1904) ( Figs 3E–F, 6G–H View Figure 6 )
Hylodes granulosus Boulenger, 1903: 553 . Holotype: BM 1947.2 .15.72 (formerly 1902.11.28.15). Type locality: ‘ Santo Domingo GoogleMaps , Carabaya, S.E. Peru, 6000 feet’ ( Fig. 4 View Figure 4 ), coordinates: 13°49′59.6′′S, 69°38′31.8′′W.
Lynch (1989) synynomized E. granulosus with E. cruralis View in CoL . He was probably influenced by his broad concept that E. cruralis View in CoL included specimens from very different sizes, altitudes, and habitats. Indeed, the series he studied included several species: O. cruralis View in CoL , O. granulosus , and O. sanderi View in CoL . After studying the holotype of both O. cruralis View in CoL and O. granulosus , and additional specimens from around the type locality of O. granulosus , we concluded that O. granulosus is a valid species (see diagnosis and redescription of the holotype below). Moreover, recently collected specimens of O. granulosus by JC at the type locality (Santo Domingo, Cordillera de Carabaya, Provincia Sandia, Department Puno, Peru) confirm this position. This represents a remarkable discovery, because no specimen of this species had been found since Mr Ockenden’s expedition at the end of the 19th century.
Diagnosis: A medium-sized Oreobates (SVL of adults, 22.6–36.9 mm) characterized as follows: (1) skin of dorsum granular, with round keratinized granules and small, sparse, low, flat warts, only some of them slightly enlarged; warts on flanks slightly larger than those of dorsum; occipital W-shaped fold and/or x-shaped fold on mid-dorsum; venter smooth; posterior surfaces of limbs smooth, groin areolate; discoidal fold present; no dorsolateral folds; large postrictal glands; (2) tympanic membrane and annulus distinct, both with length about half of the eye length; supratympanic fold weak and short; (3) head large, as wide as long; snout short, round in dorsal and lateral views; canthus rostralis slightly convex or sinuous in dorsal view, round in profile; (4) cranial crests absent; upper eyelid covered by small warts; (5) dentigerous process of vomers large, oblique, almost in contact, situated posteromedial to choanae; (6) males with vocal slits and nuptial pads; (7) hands with long and slender fingers, first finger equal or slightly shorter than second; subarticular tubercles large, prominent, round to conical; supernumerary tubercles large or small, round to conical, smaller than subarticular tubercles; tips of fingers III and IV round, slightly enlarged, lacking circumferential grooves and ungual flap; lateral fringes and keels on fingers absent; (8) ulnar tubercles absent; (9) no tubercles on heel or tarsus; (10) inner metatarsal tubercle ovate, prominent; outer metatarsal tubercle smaller, round, prominent; supernumerary tubercles inconspicuous, small, or absent; (11) toes long and slender (foot length 50–60% of SVL), lateral fringes weak or absent, webbing absent; toe V reaching the distal margin of the second subarticular tubercle of toe IV, and toe III reaching the proximal margin; tips of toes moderately enlarged, rounded, with ungual flap not indented; (12) axillary, sacral, and inguinal glands present; (13) dorsal coloration pale brown to dark brown with cream flecks, sometimes with W-shaped occipital and/or mid-dorsal X-shaped dark marks, or a pair of cream dorsolateral stripes and a short longitudinal sacral stripe; throat and chest cream with fine brown mottling; belly cream with brown mottling or reticulations on anterior margin.
Oreobates granulosus can be distinguished from other Oreobates (characters of other species in parentheses) as follows: from O. choristolemma by smaller size of females [SVL of the only known adult females, 46.4 mm (holotype) and 41.0 mm (ZFMK 72569)] ( Table 3), and lacking indented ungual flap on discs. From O. discoidalis by having warty and granular dorsal skin (finely shagreened with few enlarged warts in some specimens); tips of fingers III and IV rounded (enlarged and ovate); dentigerous process of vomers posteromedial to choanae (between choanae); canthus rostralis convex or sinuous in dorsal view, and round in lateral profile (straight and sharp). From O. cruralis by having slightly rounded, not enlarged, finger tips (enlarged and truncate tips on fingers III and IV); dorsal skin with low, round, nonpungent keratinized granules and warts (dorsal skin coarsely shagreened, composed of round low warts, without keratinized granules on dorsum); larger size, mean SVL of adult females, 36.9 mm (mean SVL of adult females, 29.3 mm) and head as long as wide (head longer than wide) ( Table 3); inconspicuous supernumerary tubercles, low, few, round (numerous, conical, and prominent supernumerary tubercles on feet). From O. heterodactylus by having dorsal skin with low, round, nonpungent keratinized granules and warts (smooth); rounded, slightly developed tips of fingers III and IV (enlarged and ovate). From O. ibischi by having tympanum length half or less than half of the eye length (more than half of the eye length); granular and warty dorsal skin (smooth to finely shagreened with some enlarged warts); rounded, slightly developed tips of fingers III and IV (enlarged and ovate discs two times wider than the digit); head as long as wide (wider than long). From O. lehri by having dorsal skin with low, round, nonpungent keratinized granules and warts (homogeneously warty); by smaller size, mean SVL of adult males, 26.6 mm (mean SVL of adult males, 32.1 mm) ( Table 3); shorter feet, FL/SVL = 50– 60% (60–70%). From O. madidi by having dorsal skin with low, round, nonpungent keratinized granules and warts (homogeneously warty); head as wide as long (longer than wide); shorter feet, FL/SVL = 50–60% (50%). From O. quixensis by smaller size, mean SVL of males and females, 26.6 and 36.9 mm, respectively [mean SVL of adult males, 39.0 mm, N = 9; mean SVL of adult females, 50.4, N = 14) ( Table 3); dorsal skin with low, round, nonpungent keratinized granules and warts (dorsal skin coarsely tuberculate, with enlarged and prominent warts and granules, some of them keratinized); inconspicuous supernumerary tubercles on feet, low, scarce, round (numerous, conical and prominent supernumerary tubercles on feet). From O. sanctaecrucis by the smaller size of adult females, 36.9 mm (mean SVL of adult females, 46.0, N = 3) ( Table 3); dorsal coloration without scarlet flecks; supernumerary tubercles inconspicuous, scarce (numerous supernumerary tubercles, prominent); longer feet, FL/SVL 50–60% (50%). From O. sanderi by having supernumerary tubercles inconspicuous, scarce (numerous supernumerary tubercles, prominent); longer feet, FL/SVL = 50–60% (FL/SVL = 50%); head as long as wide (wider than long). From O. saxatilis by its smaller size, mean SVL of females 36.9 (adult females, 49.0 and 43.7 mm, holotype and paratype, respectively) ( Table 3); dorsal skin with low, round, nonpungent keratinized granules and warts (dorsal skin coarsely tuberculate, with enlarged and prominent warts and granules, some of them keratinized); head as long as wide (wider than long). From O. simmonsi by having dorsal skin with low, round, nonpungent keratinized granules and warts (densely granular, round, pungent, keratinized granules); ulnar tubercles absent (abundant, round, small, pungent); head as long as wide (longer than wide). From O. zongoensis by having dorsal skin with low, round, nonpungent keratinized granules and warts (densely granular, round, pungent keratinized granules); slightly enlarged and truncate discs on fingers III and IV (rounded finger tips); head as long as wide (wider than long).
Description of the holotype: An adult male with vocal slits and single faint white, glandular, nonspinous nuptial pad on each thumb; head as long as wide; snout round in dorsal view and lateral profile; nostrils small, orientated dorsolaterally; canthus rostralis slightly convex in dorsal view, round in frontal profile; loreal region slightly concave, sloping gradually to the lips; lips not flared; upper eyelid without tubercles but covered by small granules; no cranial crests. Supratympanic fold indistinct; tympanic membrane and annulus distinct; tympanic membrane nearly round, its length about half of the eye length; two or three postrictal glands. Choanae not concealed by palatal shelf of the maxillary arch when roof of mouth is viewed from below; choanae large, round, anterolateral, separated by distance equal to five times the diameter of a choana; dentigerous process of the vomers larger than choanae, oblique, large, prominent, almost in contact, situated posteromedial to choanae, bearing vomerine teeth; vocal slits short, placed posterolaterally. Skin of dorsal surfaces and posterior parts of hind limbs granular, with round, low, keratinized granules and sparse, low, irregular warts; ventral surfaces smooth; W-shaped occipital fold; no dorsolateral folds; a thin mid-dorsal fold; discoidal fold evident, almost reaching the groin.
Ulnar tubercles absent; palmar tubercle round, large, flat; thenar tubercle ovate, large, prominent, its size around two-thirds of the size of the palmar tubercle; supernumerary tubercles low, round, small; subarticular tubercles round, larger than supernumerary tubercles; finger tips round, slightly enlarged, but almost the same width as corresponding digits; fingers lacking lateral fringes and keels; relative length of fingers: I <IV <II <III.
Toes long and slender (foot length 54% of SVL); heel and tarsus lacking tubercles or folds; inner metatarsal tubercle oval to round, prominent, slightly larger than outer metatarsal tubercle; metatarsal tubercle round, conical; supernumerary tubercles small, elongate, inconspicuous; subarticular tubercles prominent, conical, elongate; toes apparently without lateral fringes and keels; toe tips rounded, slightly enlarged; ungual flap not indented; relative length of toes: I <II <V <III <IV.
Colour: In preservative, the dorsal coloration is pale brown to dark brown with cream flecks, a W-shaped occipital dark mark, a pair of faint cream dorsolateral stripes from posterior margin of eyes to occipital region, and a white spot on the middle of the occipital region; head brown with cream flecks, two dark subocular bars; throat and chest cream with fine brown mottling; belly cream with brown mottling or reticulations on anterior margin; flanks cream with fine brown mottling, coalescing in an oblique, broad band posterior to arm insertion; extremities pale brown with diffuse reddish brown transverse bars.
Measurements of the holotype: SVL, 27.6 mm; HL, 10.67 mm; HW, 10.5 mm; EL, 4.6 mm; EN, 3.2 mm; IND, 3.1 mm; EE, 5.2 mm; TYH, 2.1 mm; TYL, 2.1 mm; FA, 6.5 mm; TL, 14.5 mm; TH, 14.4 mm; FL, 14.9 mm.
Variation: There is scarce variation in the qualitative characters other than colour pattern. Juveniles are almost identical to adults. There is marked sexual dimorphism in size (females are larger than males), and males have longer feet and larger EL in relation to HW and EN (see Table 3). There are two basic colour morphs. One is the colour pattern of the holotype, consisting of a pale to dark brown dorsum, with pale and dark irregular flecks, and a W-shaped dark brown or cream mark on the occipital region, and a x-shaped dark or pale mark on mid-dorsum, a pale sacral stripe, and dark transverse bars on extremities ( Fig. 6G View Figure 6 ). The alternative colour pattern (present on five of 26 specimens) consists of a dark brown dorsum delimited by a pair of pale dorsolateral bands, a sacral stripe and two wide dark brown stripes on flanks, one anterior to groin and one posterior to arm insertion, and transverse bars on extremities that are less evident than in the first colour pattern. Both colour patterns are shared by O. cruralis , O. lehri , and O. sanderi . The intensity of brown mottling on the throat and chest also varies. Glandular pads are present on axillary, groin, and sacral regions, but are sometimes completely absent or absent from only one side. The adult breeding males USNM 299011–12 present glandular pads on axillary, groin, sacral regions, and thumb. It is likely that the development of these glandular pads occurs only during the breeding season. In life, the dorsal texture and folds are also similar to preserved specimens. For example, the specimen MHNC 5328 shows incomplete dorsolateral folds composed of warts, an occipital W-shaped fold a x-mid-dorsal fold, and three enlarged round warts on the dorsal surface of the snout.
Description of colour patterns in life are based on the specimens MHNC 5328 and MHNC 5335 ( Fig. 3E, F). The adult female MHNC 5328 showed the following coloration. Head greenish brown, two subocular and one labial irregular dark brown bars, canthus and loreal region with the same coloration; tympanic fold dark brown; lower lip barred with broad dark brown bars separated by light yellow stripes; postrictal glands orange; tympanic membrane purplish brown, annulus greenish brown; an interocular dark brown bar. Dorsum reddish brown with greenish brown tonalities; some scattered red and orange warts on sacral and occipital regions; dark brown W-shaped occipital mark interiorly outlined by orange tones; mid-dorsum with an irregular X-shaped dark brown mark; sacral region with a transverse dark brown mark that continues towards the groin, forming two oblique bands in the posterior margin of the flanks; flanks light orange to pinkish cream, with some dark brown flecks; a broad irregular, oblique band on the anterior and posterior margins of each flank; groin orange. Arms greenish brown with irregular, transverse bands; arm insertion orange; limbs greenish brown with well-defined transverse dark brown bands and orange granules; plantar surfaces dark brown with grey tubercles. Throat purple, with irregular yellow spots; chest purple; belly yellowish; ventral and posterior surfaces of limbs orange. Iris coloration composed of metallic green, yellow, and orange tones, and black reticulations. Pupil horizontal, black, with a vertical black stripe ventrally. The adult male MHNC 5335 showed the following coloration. Head brown with two subocular and one labial irregular dark brown bars, canthus and loreal region with the same coloration; tympanic fold black; lower lip barred with broad dark brown bars separated by light yellow stripes; postrictal glands orange; tympanic membrane purplish brown, annulus brown; an interocular dark brown bar outlined anteriorly with two cream spots. Dorsum dark reddish brown with some scattered orange warts on sacral and occipital regions; dark brown W-shaped occipital mark interiorly outlined by orange; a pair of bold black spots in the dorsolateral region; sacral region with a thin transverse black stripe that continues toward the groin, forming two oblique bands on the posterior margin of the flanks; flanks brown with some light flecks; groin orange. Arms and limbs brown with diffuse transverse bars and orange warts. Throat brown, with white spots; belly yellow with white spots on dark brown groin anterolaterally; ventral and posterior surfaces of limbs fleshy orange. Iris bronze with black reticulations. Pupil horizontal, black, with a vertical black stripe ventrally.
Remarks: The original description by Boulenger (1903) is short but accurate. Although he overlooked the difference in relation to the degree of development of supernumerary tubercles between O. granulosus and O. cruralis , he did note the pivotal difference in skin texture. He mentioned ‘skin slightly rugose’ ( Boulenger, 1902: 580) for O. cruralis , and ‘granulate with small warts’ for O. granulosus ( Boulenger, 1903: 553) . This species, which was known from the holotype only, is now known from 25 additional specimens (see Appendix). BM 1905.5.31.21 corresponds to seven specimens (two small juveniles, two young females, a subadult female, and two adult males) collected at Limbani, Carabaya, Department Puno, Peru, by Ockenden. Boulenger identified them as O. granulosus , but he did not include them as types. BM 1905.5.31.14.20 includes four specimens (two juveniles, a subadult male, and a subadult female) from Agualani, Carabaya, Department Puno, Peru. BM 1907.5.7.17–18 correspond to an adult female and a subadult female, respectively. Both specimens were collected by Ockenden very close to the type locality. A series of specimens (USNM 299006–299012) collected by John Cadle at Ollachea 1800 m a.s.l., Department Puno, Peru, are also O. granulosus . From the series of specimens AMNH 6060–6073 identified as O. cruralis we were able to study the specimens 6060–6064. These include four specimens of O. granulosus and one of O. sanderi from Department Puno, collected by H. Keays. The specimens MHNC 5328 and MHNC 5335 ( Fig. 3E, F) were recently collected by JCC at Santo Domingo, Distrito de Limbani, Provincia de Sandia, Departamento de Puno, Peru (13°49′59.6′′S, 69°38′31.8′′W), at 1400–1500 m a.s.l. The specimen MHNC 5328 (field number 3396) is an adult female collected on 11 November 2006 along the Santo Domingo stream, 1400 m a.s.l., on a rock 1.5 m above ground. The specimen MHNC 5335 (field number 3403) is an adult male collected on 11 November 2006 on the main path to Santo Domingo, 1500 m a. s. l.; it was found calling close to a small waterfall.
Distribution: This species inhabits humid montane forests and cloud forests of the eastern versant of Cordillera Carabaya, Puno, Peru ( Figs 4 View Figure 4 , 5 View Figure 5 ). It is known from four localities between 1400 and 2000 m a.s.l. The most distant localities are Limbani and Ollachea, separated by approximately 200 km.
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