Oreobates madidi, (PADIAL ET AL., 2005), 2008

Padial, José M., Chaparro, Juan C. & Riva, Ignacio De La, 2008, Systematics of Oreobates and the Eleutherodactylus discoidalis species group (Amphibia, Anura), based on two mitochondrial DNA genes and external morphology, Zoological Journal of the Linnean Society 152 (4), pp. 737-773 : 760-763

OREOBATES MADIDI (PADIAL ET AL., 2005)

COMB. NOV.

Eleutherodactylus madidi Padial et al., 2005 ( Fig. 3J)

Eleutherodactylus madidi Padial, González & De la Riva, 2005: 319 . Holotype: MNK-A 7856. Type locality: ‘ Arroyo Huacataya , Serranía Eslabón , Área Natural de Manejo Integrado Madidi, Provincia Franz Tamayo , Departamento La Paz, Bolivia (14°20′12″S, 68°05′57″W), c. 1500 m a.s.l. ’ ( Fig. 4 ).

Diagnosis: A small Oreobates (SVL of adults, 27.9– 33.0 mm) characterized as follows: (1) skin on dorsum heavily warty, warts regular in size, low, small, rounded, some of them enlarged on the posterior part of the head and scapular region; warts on flanks larger and more elongate than those of dorsum; venter smooth; posterior surfaces of limbs uniformly warty; discoidal fold present, barely visible; no dorsolateral folds; postrictal glands well developed; (2) tympanic membrane and annulus distinct, about half of the eye length; supratympanic fold weak, short; (3) head large, longer than wide; snout round to subacuminate in dorsal and lateral views; canthus rostralis rounded; (4) cranial crests absent; upper eyelid tubercles absent; (5) dentigerous process of vomers large, situated posteromedial to choanae; (6) males with vocal slits and a single faint nuptial pad on thumb; (7) first finger longer than second; subarticular tubercles enlarged and subconical to conical in profile; supernumerary tubercles small; terminal discs of fingers rounded, not enlarged; tips of fingers III and IV round; lateral fringes and keels on fingers absent; (8) ulnar tubercles absent; (9) no tubercles on heel and tarsus; (10) inner metatarsal tubercle ovoid, outer more rounded, subequal; (11) toes lacking lateral fringes or keels; webbing absent; fifth toe shorter than third; discs of toes moderately enlarged, rounded; supernumerary tubercles conical, prominent; (12) axillary gland present; (13) dorsal coloration dark brown to black, with diffuse dark marks; in preservative, dorsal regions brownish grey or purplish grey; snout dark grey with darker marks; throat cream with dense and fine brownish grey mottling; venter cream with scarce to dense brownish grey fine mottling.

Oreobates madidi is most similar to O. cruralis . It differs from O. cruralis by having a homogeneously warty dorsal surface, large warts (skin with small, round, uniform warts, smaller warts with sparse enlarged warts), rounded finger tips (slightly enlarged and truncate tips on fingers III and IV), and a different advertisement call ( Padial et al., 2005b, 2008). It can be distinguished from other small Oreobates ( O. discoidalis , O. heterodactylus , and O. ibischi ) by having rounder finger discs (truncated and enlarged on fingers III and IV); it can be distinguished from larger species of Oreobates by lacking keratinized granules on dorsum.

Remarks: Its advertisement call allows for an unequivocal identification of live male frogs ( Padial et al., 2005b; Padial et al., 2008).

Distribution: Oreobates madidi inhabits the humid montane forests of the Andean slopes of northern Bolivia, at elevations of 900–1500 m a.s.l. ( Figs 4 , 5 ). The species is only known from La Cascada and Serranía Eslabón, two localities in the Department La Paz, separated by a distance of 150 km.

OREOBATES QUIXENSIS JIMÉNEZ DE LA ESPADA, 1872

Hylodes verrucosus Jiménez de la Espada, 1872

Hylodes philippi Jiménez de la Espada, 1875

Borborocoetes quixensis – Boulenger (1882)

Leptodactylus tuberculosus Andersson, 1945

Eupsophus quixensis – Peters (1955)

Eleutherodactylus philippi – Gorham (1966)

Eleutherodactylus verrucosus – Gorham (1966)

Ischnocnema quixensis – Lynch & Schwartz (1971)

Oreobates quixensis – Caramaschi & Canedo (2006) ( Figs 3K, 6I–J )

Oreobates quixensis Jiménez de la Espada, 1872: 87 . Types: not given in the original descriptions. Type locality: San José de Moti , Province Napo, Ecuador ( Fig. 4 ).

Diagnosis: A large Oreobates (SVL of adults, 35–63 mm) characterized as follows: (1) skin of dorsum granular, with round keratinized granules and small, sparse, prominent, and enlarged warts; occipital W-shaped fold and/or x-shaped fold on middorsum; 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, wider than 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 prominent warts and granules; (5) dentigerous process of vomers large, oblique, almost in contact, situated posteromedial to choanae; (6) males with vocal slits, no nuptial pads; (7) hands with long and slender fingers, first finger longer than second; subarticular tubercles large, prominent, round to conical; supernumerary tubercles large, prominent, round to conical, smaller than subarticular tubercles; tips of fingers III and IV round, not enlarged, lacking circumferential grooves and ungual flap; lateral fringes and keels on fingers absent; (8) either one or two ulnar tubercles, round, low; (9) no tubercles on heel or tarsus; (10) inner metatarsal tubercle ovate, prominent; outer metatarsal tubercle smaller, round, prominent; supernumerary tubercles large, prominent, conical, smaller than subarticular tubercles; (11) toes long and slender (foot length 50–60% of SVL), lateral fringes weak or absent, webbing absent; tips of toes round; (12) axillary glands present; (13) dorsal coloration pale brown to dark brown with purple tonalities and cream flecks, sometimes with W-shaped occipital and/or mid-dorsal X-shaped dark marks, and a short longitudinal sacral stripe; ventral surfaces cream with fine to dense brown mottling or reticulations.

Oreobates quixensis is the largest species of the genus. It differs from all small species of Oreobates by the combination of dorsum with enlarged warts and keratinized granules, head wider than long, and rounded finger discs. Among the medium-sized and large Oreobates , it is most similar to O. saxatilis , but O. quixensis has larger subconical tubercles on dorsum and a more heavily mottled venter.

Remarks: The original description by Jiménez de la Espada (1872) is very succinct, but the redescription by Jiménez de la Espada (1875) is very accurate. Several authors have subsequently diagnosed this species (e.g. Lynch & Schwartz, 1971; Duellman, 1978; Rodríguez & Duellman, 1994).

Lynch & Schwartz (1971) placed O. quixensis in the genus Ischnocnema , and considered Hylodes philippi Jiménez de la Espada, 1875 and Hylodes verrucosus Jiménez de la Espada, 1875 as junior synonyms of O. quixensis . These two Hylodes were not described and were only illustrated ( Jiménez de la Espada, 1875). Based on the illustrations, Lynch & Schwartz (1971) considered that the variability of O. quixensis included what Jiménez de la Espada described as H. verrucosus and H. philippi . Moreover, as Duellman was not able to examine the types of O. quixensis and O. verrucosus in the MNCN, Lynch & Schwartz (1971: 107) considered all of the types lost. This status prevailed until this paper (see Frost, 2006). The type locality of O. quixensis was considered ‘la provincia de Quijos’ by Lynch & Schwartz (1971), following the original description. The type locality of H. verrucosus was considered unknown. For H. philippi they provided the correct former holotype number (MNCN 202) and the corresponding locality of origin: San José de Motí, Provincia Napo, Ecuador. Although Jiménez de la Espada never mentioned catalogue numbers in his publications, the types are preserved in the amphibian collection of the Museo Nacional de Ciencias Naturales de Madrid [see González (2006) for a detailed catalogue of the Espada’s collection at the MNCN]. Our examination of the types revealed significant differences between the two Hylodes taxa and O. quixensis . For example, both H. verrucosus and H. philippi lack supernumerary plantar tubercles, a character illustrated in detail by Jiménez de la Espada (1875) for O. quixensis . Moreover, none of the latter species belongs to the genus Oreobates , and both belong to the E. dolops species group (sensu Lynch, 1989). The taxonomic status of these two species will be discussed elsewhere.

Jiménez de la Espada (1875) mentioned 11 specimens of O. quixensis on which he based a more accurate description. In the MNCN there are 14 specimens collected and identified by Jiménez de la Espada. One comes from San José de Moti, Municipio San José de Mote (or Nuevo), Provincia Napo; the other 13 specimens come from Municipio Archidona de Quijos, Provincia Napo, Ecuador. The specimen MNCN 1708 (formerly 330), an adult female from San José de Moti, corresponds to the specimen measured by Jiménez de la Espada (1875: 104). On the reverse of the original catalogue card corresponding to this specimen there is the inscription ‘Cotipo’ (= cotype); at the back, there is the inscription ‘genotipo’ (= type of the genus). Moreover, the card states that this specimen was illustrated in Jiménez de la Espada (1875). The original catalogue number 1154 corresponds to 13 specimens (MNCN 1709–21) from Archidona de Quijos, Napo, Ecuador. Nevertheless, Jiménez de la Espada (1875) only mentioned 11 specimens for this locality. González (2006) mentioned that two of these specimens could correspond to H. verrucosus or H. philippi ; however, we find them all assignable to O. quixensis .

We consider the specimen MNCN 1708 (formerly 330), an adult female from San José de Moti (from Municipio San José de Mote (or Nuevo), Provincia Napo, Ecuador, collected in June 1885 by Marcos Jiménez de la Espada, the lectotype of O. quixensis ( Fig. 6I–J ), and the specimens MNCN 1709–21 (formerly 1154) from Archidona, Ecuador, collected by Marcos Jiménez de la Espada on 4 May 1865, the paralectotypes of O. quixensis . The lectotype corresponds with the specimen measured and illustrated by Jiménez de la Espada (1875).

Distribution: Upper Amazon Basin in Colombia, Ecuador, Peru, Bolivia (Department of Pando), and western Brazil, up to 1000 m a.s.l. (Frost, 2006). The advertisement call was described by Duellman (1978) and Heyer & Gascon (1995), both of which also provide natural history information.

OREOBATES SANCTAECRUCIS ( HARVEY & KECK, 1995)

Ischnocnema sanctaecrucis Harvey & Keck, 1995

Oreobates sanctaecrucis – Caramaschi & Canedo (2006) ( Fig. 3L)

Ischnocnema sanctaecrucis Harvey & Keck, 1995: 57 . Holotype: MNKA 1198 , by original designation. Type locality: ‘ El Chapé, Provincia Florida, Departamento de Santa Cruz, Bolivia, 2060 m’ ( Fig. 4 ), coordinates: 18°03′34′S, 63°54′36′W.

Diagnosis: A medium-sized Oreobates (SVL of adults, 35.1–48.1 mm) characterized as follows: (1) skin of dorsum granular, with round keratinized granules and small, sparse, low, flat warts, 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, short; (3) head large, wider than 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 granules; (5) dentigerous process of vomers large, oblique, almost in contact, situated posteromedial to choanae; (6) males with vocal slits, no nuptial pads; (7) hands with long and slender fingers, first finger longer 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, not enlarged, lacking circumferential grooves and ungual flap; lateral fringes and keels on fingers absent or weak; (8) single ulnar tubercle, weak; (9) no tubercles on heel or tarsus; (10) inner metatarsal tubercle ovate, prominent; outer metatarsal tubercle smaller, round, prominent; supernumerary tubercles round, low; (11) toes long and slender (foot length 50% of SVL), lateral fringes weak or absent, webbing absent; tips of toes round, with ungual flap not indented; (12) axillary 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; a short longitudinal sacral stripe; throat and chest cream with brown mottling; belly cream with brown mottling; in life, conspicuous red to scarlet spots in axiles, dorsum, and shanks.

Oreobates sanctaecrucis can be distinguished from the rest of the species of Oreobates by the combination of dorsum with warts and keratinized granules, round finger and toe tips, and red to scarlet spots in axiles, dorsum, and shanks.

Remarks: Its advertisement call allows for an unequivocal identification of active male frogs ( Reichle, 1999; Köhler, 2000).

Distribution: Oreobates sanctaecrucis inhabits the humid montane forests and cloud forests of the Andean slopes of central Bolivia, in Departments Cochabamba and Santa Cruz, at elevations of 1000– 2100 m a.s.l. ( Figs 4 , 5 ).

Figures

Figure 4. Type localities of members of Oreobates: (1) O. quixensis, San José de Moti, Prov. Napo, Ecuador; (2) O. simmonsi, Río Piuntza, 1830 m a.s.l., Cordillera del Cóndor, Prov. Morona-Santiago, Ecuador; (3) O. saxatilis, Ponga de Shilcayo, 470 m a.s.l., Department San Martín, Peru; (4) O. lehri, Apurimac River Valley, 2445 m a.s.l., Department Cusco, Peru; (5) O. granulosus, Santo Domingo, Carabaya, Department Puno, Peru, 1800 m a.s.l.; (6) O. madidi, Arroyo Huacataya, Serranía Eslabón, 1500 m a.s.l., Department La Paz, Bolivia; (7) O. sanderi, Arroyo Bilunto, Chunirumi Valley, 1800 m a.s.l., near Santa Cruz de Valle Ameno, Department La Paz, Bolivia; (8) O. zongoensis, Valle de Zongo, 1250 m a.s.l., Department La Paz, Bolivia; (9) O. choristolemma, Serranía de Bellavista c. 1000 m a.s.l., Department La Paz, Bolivia; (10) O. cruralis, Department La Paz, Bolivia, 4000 m a.s.l. (in error); (11) O. heterodactylus, gruta Facendinha, State Mato-Grosso, Brazil; (12) O. ibischi, km 68.5 on Santa Cruz de la Sierra-Samaipata road c. 750 m a.s.l., Department Santa Cruz, Bolivia; (13) O. sanctaecrucis, El Chapé, Department Santa Cruz, Bolivia, 2060 m a.s.l.; (14) O. discoidalis, Tucumán, Prov. Tucumán, Argentina.

Figure 5. Altitudinal distribution across habitat types of members of the genus Oreobates.

Figure 6. Type specimens of some members of Oreobates. A–B, holotype of O. cruralis (BM 1947.2.15.70); C–D, holotype of O. simmonsi (KU 147068); E–F, paralectotype of O. discoidalis (BM 1947.2.15.63); G–H, holotype of O. granulosus (BM 1947.2.15.72); I–J, lectotype of O. quixensis (MNCN 1708).