NEW SPECIES OF ELEUTHERODACTYLUS (ANURA: LEPTODACTYLIDAE) FROM THE EASTERN ANDES OF CENTRAL PERU...

NEW SPECIES OF ELEUTHERODACTYLUS (ANURA:LEPTODACTYLIDAE) FROM THE EASTERN ANDES OF CENTRAL

PERU WITH COMMENTS ON CENTRALPERUVIAN ELEUTHERODACTYLUS

EDGAR LEHR1,4,5, MIKAEL LUNDBERG

1, CESAR AGUILAR2, AND RUDOLF VON MAY

3

1Staatliche Naturhistorische Sammlungen Dresden, Museum fur Tierkunde, Konigsbrucker Landstrasse 159,D-01109 Dresden, Germany

2Museo de Historia Natural, Departamento de Herpetologıa, Universidad Nacional Mayor de San Marcos,Av. Arenales 1256, Jesus Marıa, Ap. 14-0434, Lima, Peru

3Department of Biological Sciences, Florida International University, 11200 SW 8th Street, Miami, FL 33199, USA

ABSTRACT: Four new species of Eleutherodactylus are described from montane forests at elevations of1330–3000 m in the Cordillera Oriental in the central Peruvian Departamentos Huanuco, Junın and Pasco,Three of the new species are assigned to the Eleutherodactylus unistrigatus Group. Two of them lacka tympanum, but have distinct colorations and iris patterns: one has the groin and hind limbs partly orangeto red and the iris with a brown vertical and black horizontal streak forming a cross, whereas the other hasthe groin and hind limbs partly yellow and the iris with a brown horizontal streak and a black vertical streakfrom the pupil to the lower margin of the eye forming a ‘‘T’’. The third species, assigned to theEleutherodactylus conspicillatus Group, is similar to E. rhabdolaemus, but differs from that in certainmorphological characters and coloration. The fourth species, assigned to the E. lacrimosus assemblage, hasa yellowish-tan dorsum with dark brownish-purple blotches and streaks and white flanks and venter. Itinhabits terrestrial bromeliads and is compared with E. schultei from northern Peru. Morphological anddistributional data are provided for seven previously described Eleutherodactylus from central Peru. Theelevational distributions of Andean Eleutherodactylus, Phyllonastes, and Phrynopus from central Peru iscompared.

Key words: Andes; Anura; Eleutherodactylus; Leptodactylidae; New species; Peru.

KNOWLEDGE OF ELEUTHERODACTYLINE FROGS

in central Peru (Departamentos Huanuco,Pasco, Junın, and southern part of SanMartın) is limited. The species compositionof Phrynopus seems to be relatively wellknown (Hedges, 1990; Duellman, 2000; Lehr,2002; Lehr et al., 2005a), but only one speciesof Phyllonastes (Lehr et al., 2004a) and 10species of Eleutherodactylus have been de-scribed for central Peru within the last 10years (Dwyer, 1995; Lehr et al., 2004b; Lehr2005; Duellman and Hedges, 2005). Intensivefieldwork since 1998 on the lower and upperforested slopes of the eastern Andes incentral Peru has resulted in the collection ofseveral, mostly unidentified, species ofEleutherodactylus, two of which were recentlydescribed by Lehr et al. (2004b) and Lehr(2005). The taxonomy of Eleutherodactylus is

complex because of the high number ofspecies (more than 700; Frost, 2004), highinterspecific similarity, high intraspecific poly-morphism, and the difficulty of identifyingdiagnostic characters (coloration, skin texture)in poorly preserved specimens. Access toscientific collections that harbor many typespecimens as well as identified species ofEleutherodactylus is extremely helpful asa source of comparative material for speciesdescriptions. This was made possible whenthe senior author visited the Division ofHerpetology at The University of KansasNatural History Museum and BiodiversityResearch Center. The examination of recentlycollected specimens and comparison withmuseum material, led to the recognition offour undescribed species. Herein, we describethe new species, add information aboutmorphology and distribution for seven othercentral Peruvian Eleutherodactylus, and com-pare their elevational distributions with thoseof Phyllonastes and Phrynopus in the Andes ofcentral Peru.

4 PRESENT ADDRESS: Natural History Museum andBiodiversity Research Center, Division of Herpetology,The University of Kansas, Jayhawk Boulevard 1345,Lawrence, KS 66045-7561, USA

5 CORRESPONDENCE: e-mail, [email protected]

Herpetological Monographs, 20, 2006, 105–128

E 2006 by The Herpetologists’ League, Inc.

105

MATERIALS AND METHODS

The format for the diagnoses follows thatof Lynch and Duellman (1997), with in-clusion of a description of a holotype. Fieldnotes on the new species were recorded byE. Lehr in August 2003, and by M.Lundberg in March 2000, March 2002, andJanuary 2004. Additionally, photographswere used for descriptions of coloration inlife. Definitions of the Peruvian ecoregionsfollow Brack (1986). Specimens collectedbetween 2000 and 2002 were preserved in4% formalin; those collected in 2003 and2004 were preserved in 96% ethanol; allwere stored in 70% ethanol. If secondary sexcharacters were present, specimens weresexed externally; otherwise, they were dis-sected to determine the presence or absenceof ovaries. We follow the definition ofa tympanum by Lynch and Duellman(1997). The otic region was dissected inorder to evaluate the condition of thetympanic annulus. Measurements taken withdigital calipers and rounded to the nearest0.1 mm are: SVL (snout–vent length), TL(tibia length), FL (foot length, distance fromproximal margin of inner metatarsal tubercleto tip of Toe IV), HL (head length, from

angle of jaw to tip of snout), HW (headwidth, at level of angle of jaw), ED (eyediameter), IOD (interorbital distance), EW(upper eyelid width), IND (internarial dis-tance), E–N (eye–nostril distance, straightline distance between anterior corner oforbit and posterior margin of external nares),TY (tympanum diameter, horizontal).Lengths of Toes III and V were determinedwhen both were adpressed against Toe IV;lengths of Fingers I and II were determinedwhen adpressed against each other. Alldrawings were made by the senior authorusing a stereomicroscope with drawing tubeattachment (Nikon SMZ 1000). Specimenswere deposited in the herpetological collec-tions of the Museo de Historia NaturalUniversidad Nacional Mayor de San Marcos(MHNSM) in Lima, Peru, and the Museumfur Tierkunde Dresden (MTD), in Dresden,Germany. Codes for museum collectionsfollow those of Leviton et al. (1985) andFrost (2004). For specimens examined, seeAppendix I.

Eleutherodactylus cruciocularis sp. nov.

Holotype.—MHNSM 18685 (Figs. 1A, B),an adult female obtained from Pampa Her-

FIG. 1.—Eleutherodactylus cruciocularis (MHNSM 18685, holotype, SVL 21.8 mm) in lateral (A) and ventral (B)views, photos by E. Lehr; and Eleutherodactylus flavobracatus (MHNSM 19871, SVL 21.5 mm) in lateral (C) andventral (D) views, photos by M. Lundberg.

106 HERPETOLOGICAL MONOGRAPHS [No. 20

mosa (10u 599 33.30 S, 75u 259 58.00 W) at1540 m between 28 and 29 August 2003,Distrito de Huasahuasi, Provincia deTarma, Departamento de Junın, Peru, by E.Lehr.

Paratopotypes.—All collected with the ho-lotype by C. Aguilar, E. Lehr, and R. von May:6 females (MHNSM 18682–84, MTD 45634,45637–38), 15 males (MHNSM 18686–91,18697, MTD 45635–36, 45639–44).

Referred specimens.—Three females (MH-NSM 19988, MTD 46832–33), and fivejuveniles (MHNSM 19985–87, MTD 46834–35), all obtained from San Alberto (10u 34938.50 S, 75u 239 32.20 W) at 1850 m on 25June 2005, Distrito de Oxapampa, Provinciade Oxapampa, Departamento de Pasco, Peru,by M. Lundberg; one female (AMNH 91548)obtained from about 30 km (airline) NETingo Marıa, Cordillera Azul, [5 5 km byroad SW high point (1640 m) on Tingo Marıa-Pucallpa Road] at 1330 m on 3 November1974, Departamento de Huanuco, Peru, by C.W. Myers and J. Daly.

Diagnosis.—A member of the Eleuthe-rodactylus (Eleutherodactylus) unistrigatusGroup having the following combination ofcharacters: (1) Skin on dorsum shagreen withsmall scattered spicules, that on venterareolate; discoidal fold not evident; dorsolat-eral folds absent; (2) tympanic membrane andtympanic annulus absent; (3) snout short,rounded in dorsal and lateral views; (4) uppereyelid with small tubercles; width of uppereyelid narrower than IOD; cranial crestsabsent; (5) vomerine teeth small; (6) maleslacking vocal sac, vocal slits, and nuptial pads;(7) Finger I shorter than Finger II; discs onouter fingers broadly expanded, rounded; (8)fingers without lateral fringes; (9) ulnar andtarsal tubercles present; (10) heel with singletubercle; inner tarsal fold absent; (11) innermetatarsal tubercle moderate, ovoid, 33 aslarge as outer; outer metatarsal tubercle small,rounded; few low, diffuse supernumeraryplantar tubercles present; (12) toes withoutlateral fringes; toe webbing absent; Toe Vmuch longer than Toe III; toe discs slightlysmaller than those on fingers; (13) dorsum andventer dark grayish brown; groin and anteriorsurfaces of thighs orange to red; iris gold, withfine, black reticulations and a dark brown,

broad horizontal streak across pupil anda narrow black vertical streak across pupilforming a cross; (14) SVL in adult females18.7–21.8 mm (n 5 7), in adult males 11.4–15.4 mm (n 5 7).

Eleutherodactylus cruciocularis is readilydistinguished from all other species of thegenus by the combination of its lack ofa tympanic annulus and membrane, posses-sion of an iris with a cruciform mark and thepartly orange to red groin and hind limbs.Five other species of Eleutherodactylus (E.colodactylus, E. flavobracatus, E. imitatrix, E.lirellus, and E. rhabdocnemus) from theeastern Andes and Amazonian lowlands ofcentral Peru lack a tympanum, but none hasan iris with a cruciform mark. Eleutherodac-tylus cruciocularis differs from the similar-sized E. imitatrix (females to 20 mm [Duell-man and Hedges, 2005] vs. up 21.8 mm in E.cruciocularis) by having vomerine teeth (ab-sent in E. imitatrix). Eleutherodactylus cru-ciocularis is smaller than E. rhabdocnemus(females to 21.8 mm vs. 24.5 mm in E.rhabdocnemus [Duellman and Hedges,2005]), and has a dark gray belly with palebrown spots (pale belly with brown flecks andsmall brown spots).

Eleutherodactylus cruciocularis is mostsimilar to E. flavobracatus, but differs fromit in the following features (characters of E.flavobracatus in parentheses): maximum SVLin females 21.8 mm (23.4) and in males15.4 mm (19.6), groin and hind limbs withorange or red blotches (yellow), posteriorsurface of thighs brown with small, darkorange spots (brown with large yellow longi-tudinal stripe extending to shanks), uppereyelid tubercles low (prominent), supratym-panic fold absent (weak), and eye with darkbrown horizontal and narrow, black verticalstreak forming a cross (dark brown horizontalstreak present, black vertical streak from pupilto venterolateral margin of eye). Eleuthero-dactylus cruciocularis shares with E. altama-zonicus a nearly uniform dark venter, andorange to red blotches in groin and anteriorsurfaces of thighs, but differs as follows(characters of E. altamazonicus in parenthe-ses): maximum SVL in females 21.8 mm(33.9 mm; Lynch 1980), tympanum absent(present, although partly concealed in some

2006] HERPETOLOGICAL MONOGRAPHS 107

individuals), and inner tarsal fold absent(present).

Description of the holotype.—Head as wideas body and nearly as wide as long; head width35% of SVL; head length 33% of SVL; snoutshort, rounded in dorsal and in lateral view(Figs. 2A, B); eye diameter larger than eye–nostril distance (eye–nostril distance 83% oflength of eye); nostrils slightly protuberant,directed dorsolaterally; canthus rostralisslightly curved in dorsal view, rounded inprofile; loreal region slightly concave; lipsrounded; upper eyelid bearing small, lowtubercles; upper eyelid width 74% of IOD;supratympanic fold, tympanic annulus andtympanic membrane absent; enlarged post-rictal tubercles absent. Choanae small, ovoid,not concealed by palatal shelf of maxilla;vomerine teeth minute, concealed by palatalmucosa, situated posteromedial to choanae,each vomer bearing four teeth in an obliquerow, narrowly separated; tongue 1.83 as longas wide (length 6.8 mm, width at midlength oftongue 3.7 mm), not notched posteriorly,posterior one-half free.

Skin on dorsum smooth with few, scatteredspicules; dorsolateral folds absent; skin on

flanks smooth with few, scattered spicules;skin on belly and chest areolate, skin on otherventral surfaces smooth; discoidal fold notevident; cloacal sheath short; large tuberclesabsent in cloacal region. Ulnar and tarsaltubercles present, both low; palmar tuberclesslightly elevated, outer palmar tubercle bifid,approximately 2.53 the size of ovoid, innerpalmar tubercle; supernumerary tuberclesbelow Fingers III–IV, round, low, slightlysmaller than subarticular tubercles; subarticu-lar tubercles well defined, round in dorsalview and subconical in lateral view, mostprominent on base of Finger IV; fingerswithout lateral fringes; Finger I shorter thanFinger II; discs on fingers expanded, mostprominent on Fingers III–IV, rounded termi-nally; discs on Finger I slightly expanded; allfingers having ventral pads well defined bycircumferential grooves (Fig. 3A). Hind limbsslender, tibia length 50% of SVL; foot length39% of SVL; upper surfaces of hind limbssmooth; posterior and ventral surfaces ofthighs weakly areolate; heel bearing single,round tubercle; tarsus with row of small, lowtubercles; tarsal fold absent; inner metatarsaltubercle elevated, elliptical, about 33 sub-conical outer metatarsal tubercle; few, lowplantar supernumerary tubercles; subarticulartubercles well defined, round in dorsal viewand subconical in lateral view; toes withoutlateral fringes; toe webbing absent; discs ontoes about equal in size to those on fingers,most prominent on Toes IV–V, toe discs welldefined by circumferential grooves; relativelengths of toes: 1 , 2 , 3 , 5 , 4 (Fig. 3B);Toe V much longer than Toe III (disc on ToeIII not reaching distal subarticular tubercle onToe IV, tip of the disc on Toe V extending todistal border of distal subarticular tubercle onToe IV).

Measurements (in mm) of holotype: SVL21.8; TL 11.0; FL 8.5; HL 7.2; HW 7.6; ED2.9; IOD 2.7; EW 2.0; IND 1.8; E–N 2.4.

Coloration of holotype in preservative.—Dorsum grayish brown with dark browninterorbital bar and blotches in scapula andsacral regions; limbs with dark brown trans-verse bars; three dark brown labial bars;narrow, dark brown stripe extending fromposterior margin of eye diagonally to forearminsertion; venterolateral posterior half of

FIG. 2.—Lateral (A) and dorsal (B) views of head ofEleutherodactylus cruciocularis (MHNSM 18685); lateral(C) and dorsal (D) views of head of E. flavobracatus(MHNSM 19871).


flanks dark brown with pale cream blotches;groin and anterior surface of thighs brownwith pale cream blotches, posterior surface ofthighs brown with small pale cream spots;throat pale gray, chest and belly grayish brownwith pale gray spots; posterior part of bellyand anterior half of thighs with a pale creamblotch, separated medially by a narrow, gray-ish-brown stripe; outer surface of left shankwith three pale cream blotches, right shankwith five pale cream blotches; tips of fingersand toes pale gray; iris pale gray with fine,black reticulations and black horizontal andvertical streaks that form a cross in the irisacross the pupil.

Coloration of holotype in life.—Dorsumpale brown with small red spots on back, darkbrown interorbital bar, dark brown blotches inscapula and sacral regions; limbs with darkbrown transverse bars; three dark brown labialbars; narrow, dark brown supratympanic stripeextending from posterior margin of eye di-agonally to insertion of forearm; posteroventralhalf of flanks black with pale cream blotches;groin with dark orange blotches; anteriorsurfaces of thighs brown with dark orangeblotches, posterior surfaces of thighs brownwith small dark orange spots; ventral surfacesdark gray with pale cream spots; posterior partof belly and anterior half of thighs with a darkorange blotch, separated medially by a narrow,dark gray stripe; outer surface of left shank withthree dark orange blotches, right shank withfive orange blotches; tips of fingers pale yellow,of toes, pale red; iris gold with fine, blackreticulations and dark brown horizontal andblack vertical streaks that form a cross in theiris across the pupil.

Variation.—The paratypes differed in dor-sal and lateral coloration in life. MHNSM18684, 18688, MTD 45638, and 45642 had tanflanks and a dark brown dorsum. In MHNSM18682, 18691, MTD 45635, and 45644, thedorsum was pale brown with a dark brownmiddorsal stripe and two dark brown lateralstripes extending from about the naris to thecloaca; the flanks were dark brown. In MTD45643, there was a single, narrow, pale brownmiddorsal stripe and a pale brown stripe onthe throat. Also, the specimens differ in thenumbers and sizes of the orange blotches.Coloration in life of a female (AMNH 91548)was noted by C. W. Myers as follows: ‘‘Patternof contrasting, brown markings above, withvivid bright orange flash mark in groin. Rearthigh black with gray flecking and few dots ofpale orange; some small pale orange spots onshank.’’ See Tables 1, 2 for measurements ofthe type series; see Table 3 for ranges andproportions.

Etymology.—The specific name is derivedfrom the Latin noun crux, meaning cross, andthe Latin noun oculatus, meaning ‘‘havingeyes.’’ The name refers to the cruciform irispattern of this species.

Distribution and ecology.—Eleutherodacty-lus cruciocularis is known from three localities

FIG. 3.—Ventral views of hand (A) and foot (B) of E.cruciocularis (MHNSM 18685); ventral views of hand (C)and foot (D) of E. flavobracatus (MHNSM 19871).


(Pampa Hermosa, San Alberto, and 30 km[airline] NE Tingo Marıa) at elevations of1330–1850 m in the Selva Alta Ecoregion ofcentral Andean Peru. The type locality(Fig. 4) is near the Chanchamayo Valley and

supports undisturbed primary forest. Allspecimens were found at night on vegetationapproximately 1 m above the ground or on theground. Sympatric anurans include Eleuther-odactylus bipunctatus, E. bromeliaceus, E. cf.

TABLE 1.—Measurements (in mm) of adult females of Eleutherodactylus cruciocularis.

Character MHNSM 18685 MHNSM 18684 MTD 45637 MTD 45638 MHNSM 18683 MHNSM 18682 MTD 45634

SVL 21.8 21.6 20.8 20.7 19.6 18.9 18.7TL 11.0 10.9 10.5 11.1 10.6 9.7 9.6FL 8.5 8.6 7.6 7.7 7.5 7.2 7.5HL 7.2 7.6 7.9 7.2 7.4 6.9 7.6HW 7.6 8.0 7.9 7.6 7.5 7.1 7.4ED 2.9 2.6 2.8 2.7 2.5 2.6 2.6IOD 2.7 2.9 2.9 2.7 2.7 2.7 2.8EW 2.0 1.7 1.4 1.5 1.5 1.9 1.4IND 1.8 1.8 1.7 2.0 1.9 1.6 1.6E–N 2.4 2.2 1.9 1.9 2.0 2.1 2.2

TABLE 2.—Measurements (in mm) of selected males of Eleutherodactylus cruciocularis.

Character MHNSM 18697 MTD 45635 MTD 45644 MTD 45640 MTD 45641 MTD 45639 MHNSM 18690

SVL 15.4 14.7 14.1 13.7 12.9 12.4 11.4TL 8.0 8.0 7.7 7.2 6.8 6.7 6.2FL 6.5 6.2 5.7 5.5 5.3 5.1 4.6HL 6.6 5.6 5.1 5.5 4.6 4.6 4.7HW 5.8 5.4 5.2 4.8 4.7 4.7 4.7ED 2.0 2.4 2.2 1.7 1.9 1.5 1.7IOD 2.0 2.0 2.2 1.7 1.9 1.9 1.5EW 1.2 1.5 1.2 1.2 1.2 1.3 1.2IND 1.4 1.2 1.1 1.4 1.2 1.4 1.3E–N 2.0 1.8 2.2 1.5 1.4 1.4 1.5

TABLE 3.—Ranges (in mm) and proportions of adult Eleutherodactylus cruciocularis and E. flavobracatus; rangesfollowed by means and one standard deviation in parentheses.

Character

E. cruciocularis E. flavobracatus

Females (n 5 7) Males (n 5 7) Females (n 5 2) Males (n 5 2)

SVL 18.7–21.8 (20.3 6 1.2) 11.4–15.4 (13.5 6 1.2) 21.5–23.4 (22.5 6 1.0) 18.2–19.6 (18.9 6 0.7)TL 9.6–11.1 (10.5 6 0.6) 6.2–8.0 (7.2 6 0.6) 11.3–11.4 (11.4 6 0.0) 10.3–10.4 (10.4 6 0.0)FL 7.5–8.6 (7.8 6 0.5) 4.6–6.5 (5.6 6 0.6) 8.8–9.1 (9.0 6 0.2) 8.2–8.4 (8.3 6 0.1)HL 6.9–7.9 (7.4 6 0.3) 4.6–6.6 (5.2 6 0.7) 8.1–8.5 (8.3 6 0.2) 8.0–8.1 (8.1 6 0.0)HW 7.1–8.0 (7.6 6 0.3) 4.7–5.8 (5.0 6 0.4) 7.7–8.4 (8.1 6 0.4) 7.0–7.1 (7.1 6 0.0)ED 2.5–2.9 (2.7 6 0.1) 1.5–2.4 (1.9 6 0.3) 2.5–2.6 (2.6 6 0.1) 2.2–2.3 (2.2 6 0.0)IOD 2.7–2.9 (2.8 6 0.1) 1.5–2.2 (1.9 6 0.2) 2.6–2.7 (2.7 6 0.1) 2.8–2.9 (2.9 6 0.1)EW 1.4–2.0 (1.6 6 0.2) 1.2–1.5 (1.3 6 0.1) 1.7–1.8 (1.8 6 0.0) 1.6–1.8 (1.7 6 0.1)IND 1.6–2.0 (1.8 6 0.1) 1.1–1.4 (1.3 6 0.1) 1.6–1.9 (1.8 6 0.2) 1.4–1.6 (1.5 6 0.1)E–N 1.9–2.4 (2.1 6 0.2) 1.4–2.2 (1.7 6 0.3) 2.4–2.5 (2.5 6 0.0) 1.9–2.2 (2.1 6 0.2)TL/SVL 0.52–0.54 0.50–0.54 0.48–0.53 0.53–0.56FL/SVL 0.40–0.42 0.37–0.40 0.39–0.41 0.42–0.46HL/SVL 0.36–0.43 0.33–0.41 0.36–0.38 0.41–0.45HW/SVL 0.35–0.41 0.35–0.40 0.36 0.36HW/SVL 0.35–0.41 0.35–0.40 0.36 0.36–0.38HW/HL 0.87–1.02 0.97–1.06 0.99–1.08 0.86–0.89E–N/ED 0.74–1.00 0.68–0.85 0.96 0.68–0.76EW/IOD 0.55–0.80 0.48–0.74 0.65–0.67 0.57–0.62


platydactylus, E. sagittulus, Hybsiboas cf.melanopleura, and Gastrotheca atympana.See Figure 5 for comparative altitudinal dis-tributions.

Eleutherodactylus flavobracatus sp. nov.

Holotype.—MHNSM 19871 (Figs. 1C, D),an adult female obtained at Km 34 on road

from Oxapampa to Yaupi (10u 449 44.40 S, 75u309 02.29 W) at 1770 m on 18 October 2003,Distrito de Chontabamba, Provincia de Ox-apampa, Departamento de Pasco, Peru, by M.Lundberg.

Paratopotypes.—Four, all collected by M.Lundberg: 1 female (MTD 45908, on 18October 2003), 2 males (MHNSM 19848,MTD 45716, on 25 May 2004), 1 juvenile(MTD 45717, on 1 June 2003).

Diagnosis.—A member of the Eleuthe-rodactylus (Eleutherodactylus) unistrigatusGroup having: (1) skin on dorsum shagreenwith small scattered tubercles, that on venterareolate; discoidal fold not evident; dorsolat-eral folds absent; (2) tympanic membrane andtympanic annulus absent; (3) snout short,rounded in dorsal and lateral views; (4) uppereyelid with two tubercles, one at midlength ofeyelid near inner margin, the other at itsposterior end; upper eyelid narrower thanIOD; cranial crests absent; (5) vomerine teethsmall; (6) males lacking vocal sac, vocal slits,and nuptial pads; (7) Finger I shorter thanFinger II; discs on outer fingers broadlyexpanded, rounded; (8) fingers without lateralfringes; (9) ulnar and tarsal tubercles present;(10) heel with single tubercle; inner tarsal fold

FIG. 4.—Map with type localities of the new speciesdescribed herein: Cillapata (E. ornatus), Huasahuasi (E.pardalinus), Km 34 on road from Oxapampa to Yaupi (E.flavobracatus), and Pampa Hermosa (E. cruciocularis).

FIG. 5.—Range of altitudinal distribution in Andean Eleutherodactylus, Phyllonastes, and Phrynopus (excludingspecies in which taxonomic changes will appear in the near future) in central Peru. Ranges are indicated by bars,whereas dots indicate single collecting sites; a circle with a cross indicates species that lack a tympanum. For sources ofdistribution and precise distributional data, see Appendix II.


absent; (11) inner metatarsal tubercle moder-ate, ovoid, 23 as large as outer; outer meta-tarsal tubercle small, rounded; few supernu-merary plantar tubercles present, low, diffuse;(12) toes without lateral fringes; toe webbingabsent; Toe V much longer than Toe III; discson toes slightly smaller than those on fingers;(13) dorsum and venter dark brown; groin andanterior and posterior surfaces of thighsyellow; shank yellow ventrally; iris gold withfine, black reticulations and a broad, darkbrown horizontal streak across pupil anda narrow, black vertical streak from pupil tolower margin of eye forming a ‘‘T;’’ (14) SVLin adult females 21.5–23.4 mm (n 5 2), inadult males 18.2–19.6 mm (n 5 2).

Eleutherodactylus flavobracatus is readilydistinguished from all species of the genus bythe combination of lacking a tympanum andpossessing an iris with a dark brown horizontalstreak and a dark brown vertical streak frompupil to lower margin of eye forming a ‘‘T,’’and having the groin and hind limbs partlyyellow. Five other species of Eleutherodac-tylus (E. colodactylus, E. cruciocularis, E.imitatrix, E. rhabdocnemus, and E. sp. nov.)from the eastern Andes and Amazonian low-lands of Peru lack a tympanum, but in nonedoes the eye have a T–shaped mark. Further-more, E. flavobracatus is slightly larger thanE. imitatrix (SVL to 23.4 mm in female E.flavobracatus vs. 20 mm in female E. imitatrix[Duellman and Hedges, 2005]) and hasvomerine teeth (absent in E. imitatrix).Eleutherodactylus flavobracatus differs fromE. rhabdocnemus (which has a similar size,24.5 mm in females [Duellman and Hedges,2005]) by having prominent tubercles on theupper eyelid (absent in E. rhabdocnemus),and by having a brown belly with dark brownspots (belly pale with brown flecks and smallbrown spots). Eleutherodactylus flavobracatusis most similar to E. cruciocularis; forcomparison of the two species see diagnosisof E. cruciocularis.

Description of the holotype.—Head as wideas body, slightly longer than wide; head width36% of SVL; head length 38% of SVL; snoutshort, rounded in dorsal and lateral views(Figs. 2C, D); eye diameter barely larger thaneye–nostril distance (eye–nostril distance 96%of length of eye); nostrils slightly protuberant,

directed dorsolaterally; canthus rostraliscurved in dorsal view, rounded in profile;loreal region slightly concave; lips rounded;upper eyelid bearing two prominent tuber-cles, one located at midlength of eyelid nearouter margin, the other at its posterior end;upper eyelid width 65% of IOD; supratympan-ic fold weak, tympanic annulus and tympanicmembrane absent; enlarged postrictal tuber-cles absent. Choanae small, ovoid, not con-cealed by palatal shelf of maxillary arch;vomerine teeth small, concealed by sur-rounding tissues, situated posteromedially tochoanae, each vomer bearing four teeth in anoblique row and narrowly separated; tongue1.23 as long as wide (length 5.2 mm, widthat midlength of tongue 4.5 mm), not notchedposteriorly, posterior one-third free.

Skin on dorsum smooth with few, scatteredspicules; dorsolateral folds absent; skin onflanks smooth with few, scattered spicules; skinon belly and chest weakly areolate, skin onother ventral surfaces smooth; discoidal foldnot evident; cloacal sheath short; large tuber-cles in cloacal region absent. Ulnar and tarsaltubercles present, both low; palmar tuberclesslightly elevated, outer palmar tubercle bifid,approximately 23 size of ovoid, inner palmartubercle; supernumerary tubercles below Fin-gers II–IV, round, low, slightly smaller thansubarticular tubercles; subarticular tubercleswell defined, round in dorsal view andsubconical in lateral view, most prominent onbase of Finger I; fingers without lateral fringes;Finger I shorter than Finger II; discs on fingersexpanded, most prominent on Fingers III–IV,rounded terminally; disc on Finger I slightlyexpanded; all fingers having ventral pads welldefined by circumferential grooves (Fig. 3C).The right hand had been cut for tissue source.Hind limbs slender, tibia length 53% of SVL;foot length 41% of SVL; upper surfaces of hindlimbs smooth; posterior and ventral surfaces ofthigh areolate; heel bearing single, slightlyelevated, round tubercle; tarsus with row ofsmall, low tubercles; tarsal fold absent; innermetatarsal tubercle elevated, elliptical, about23 subconical outer metatarsal tubercle; few,low plantar supernumerary tubercles present;subarticular tubercles well defined, round indorsal view and subconical in lateral view; toeswithout lateral fringes; toe webbing absent;


discs on toes slightly smaller than those onfingers, most prominent on Toes IV–V, welldefined by circumferential grooves; relativelengths of toes: 1 , 2 , 3 , 5 , 4 (Fig. 3D);Toe V much longer than Toe III (disc on ToeIII not reaching distal subarticular tubercle onToe IV, tip of the disc on Toe V extending todistal border of distal subarticular tubercle onToe IV).

Measurements (in mm) of holotype: SVL21.5; TL 11.4; FL 8.8; HL 8.1; HW 7.7; ED2.6; IOD 2.6; EW 1.7; IND 1.6; E–N 2.5.

Coloration of holotype in preservative.—Dorsum dark brown with tan blotch anteriorto nares; hind limbs with transverse darkbrown bars; ill–defined, dark brown labialbars present; narrow, dark brown supratym-panic stripe extending from posterior marginof eye diagonally to insertion of forearm;flanks tan dorsolaterally, dark brown ventro-laterally; groin with a white blotch; anteriorsurfaces of thighs white, bordered by darkbrown bars, posterior surfaces of thighs witha transverse white streak; ventral surfacesbrown with dark brown spots, except forposterior part of belly, proximal half of thighs,and ventral surfaces of shanks which arewhite; tarsus white dorsally; finger tips andtoe tips pale gray. Iris black.

Coloration of holotype in life.—As de-scribed above with the exception that thewhite areas were yellow, and the iris gold withfine, black reticulations, a dark brown hori-zontal streak across pupil, and a black verticalstreak from pupil to lower margin of eyeforming a ‘‘T.’’

Variation.—Variation exists in the dorsaland lateral coloration, and in the amount of

yellow coloration. In MTD 45908, the dorsumand flanks were dark brown, and there wasa yellow blotch in the groin. The holotype hadthe most yellow coloration in the groin and onthe hind limbs. See Table 4 for measurementsof the type series; see Table 3 for ranges andproportions.

Etymology.—The specific name is derivedfrom the Latin noun flavus, meaning yellow,and the Latin adjective bracatus, meaning‘‘wearing trousers.’’ The name refers to theyellow coloration of groin and parts of thehind limbs of this species.

Distribution and ecology.—Eleutherodacty-lus flavobracatus is known only from the typelocality (Fig. 4), which is in the Selva AltaEcoregion. All specimens were found at nightat 1900–2200 hr on vegetation 1–3 m aboveground, next to a road. The roadside is nearlyvertical and covered with secondary vegeta-tion, which is cut several times a year as partof road maintenance. There are several smallcreeks near the type locality, and a roadsideditch contained running water. At night theleaves were covered with water drops evenwhen it was not raining. No calls were heard.Sympatric anurans include Atelopus sp.,Eleutherodactylus bipunctatus, E. rhabdocne-mus, and Hyalinobatrachium bergeri. SeeFigure 5 for comparative altitudinal distribu-tions.

Eleutherodactylus ornatus sp. nov.

Holotype.—MHNSM 20664 (Figs. 6A, B),an adult female obtained at Cillapata (ca.1.5 km NNE Auquimarca, approximately 10u439 520 S; 75u 429 480 W) at 2900 m on 11March 2002, Distrito de Paucartambo, Pro-

TABLE 4.—Measurements (in mm) of type series of Eleutherodactylus flavobracatus.

Character MTD 45908 MHNSM 19871 MHNSM 19848 MTD 45716 MTD 45717

Sex Female Female Male Male JuvenileSVL 23.4 21.5 19.6 18.2 14.7TL 11.3 11.4 10.4 10.3 8.8FL 9.1 8.8 8.2 8.4 6.8HL 8.5 8.1 8.0 8.1 5.6HW 8.4 7.7 7.1 7.0 5.6ED 2.5 2.6 2.2 2.3 1.9IOD 2.7 2.6 2.8 2.9 2.0EW 1.8 1.7 1.6 1.8 1.3IND 1.9 1.6 1.4 1.6 1.6E–N 2.4 2.5 1.9 2.2 1.7


vincia de Pasco, Departamento de Pasco,Peru, by M. Lundberg.

Paratypes.—Twenty (7 males, 10 females, 3juveniles), all from Provincia de Pasco,Departamento de Pasco, Peru, collected byby M. Lundberg and E. Lehr: Paucartambo:Chinche, 3000 m: MTD 44769 (4 April 2002);

Paucartambo: Cillapata (approximately 1.5 kmnorth-northeast of Auquimarca), approxi-mately 10u 439 520 S; 75u 429 480 W,2900 m: MHNSM 17831 (14 April 2000),20665–71 (March–April 2002), MTD 44766–68 (11 March 2002), MTD 44770–72 (March2002), MTD 45073 (1 June 2002); Paucar-tambo: near road from Uchuhuerta to Auqui-marca (10u 419 05.60 S; 75u 459 28.70 W,2900 m): MTD 45897–98 (10 December2003); Huachon: Uchuhuerta, 2400 m:MHNSM 19883 (22 October 2003); Huachon:km 77 at the road Ninacaca-Huachon-Puag-maray-Oxapampa, 2550 m: MTD 46319 (27January 2004).

Diagnosis.—A member of the Eleuthero-dactylus (Eleutherodactylus) conspicillatusGroup having: (1) skin on dorsum finelyshagreen with small scattered tubercles, thaton venter smooth; discoidal fold present;dorsolateral folds present; (2) tympanic mem-brane and tympanic annulus present, upperand posterolateral margins concealed bysupratympanic fold; (3) snout long, subacumi-nate in dorsal view, round in lateral view; (4)upper eyelid without tubercles; width ofupper eyelid much narrower than IOD;cranial crests absent; (5) vomerine teeth small,oblique; (6) males with vocal slits, vocal sacnot visible externally, nuptial pads absent; (7)Finger I shorter than Finger II; discs on outerfingers broadly expanded, rounded; (8) fingerswithout lateral fringes; (9) ulnar and tarsaltubercles absent; (10) heel without tubercles;inner tarsal fold absent; (11) inner metatarsaltubercle moderate, ovoid, 23 as large asouter; outer metatarsal tubercle small, round-ed; few supernumerary plantar tuberclespresent, low; (12) toes without lateral fringesand webbing; Toe V slightly longer than ToeIII; toe discs smaller than those on fingers;(13) dorsum tan with dark brown blotchesforming two irregular chevrons; extremitieswith irregular dark brown blotches; darkbrown canthal and supratympanic stripes;venter whitish gray with brown blotches onchest and throat; groin and anterior andposterior surfaces of thighs reddish brown;lower one-half of iris dark brown, upper one-half pale gold with fine, black reticulations;(14) SVL in adult females 20.7–27.3 mm (n 511), in adult males 16.7–20.5 mm (n 5 7).

FIG. 6.—Eleutherodactylus ornatus (MTD 45897, fe-male, SVL 21.1 mm), in lateral (A) and ventral (B) views,photos by E. Lehr; Eleutherodactylus rhabdolaemus (KU173237, male, SVL 22.7 mm) in lateral view (C), photo byW. E. Duellman.


Eleutherodactylus ornatus is most similar toE. rhabdolaemus (Fig. 6C), a member of theE. conspicillatus Series according to Lynchand Duellman (1997), which is known fromthe Peruvian Departamentos Huanuco, Aya-cucho, and Cusco, southwards to Departa-mento Santa Cruz in Bolivia at elevations of1000–2700 m (Frost, 2004). Both E. ornatusand E. rhabdolaemus have dorsolateral foldsand dark brown canthal and supratympanicstripes (broader in E. ornatus), but thesespecies can be distinguished by the followingfeatures (characters of E. rhabdolaemus inparentheses): maximum SVL in females27.3 mm (31.7; Duellman, 1978a) and inmales 20.5 mm (24.0; Duellman, 1978a),dorsolateral fold weakly defined (clearly dis-tinct), diameter of eye approximately 33 thediameter of tympanum (approximately 23),diameter of tympanum equal to eye–tympa-num distance, Figs. 7A, B (one-half of tym-panum diameter, Figs. 7C, D), males withvocal sac not visible externally (subgularvocal sac visible), vomerine teeth in a clump(oblique row), scapular tubercles absent(present), venter smooth (coarsely areolate),supratympanic fold prominent (weak), termi-nal discs on fingers not broadly expanded,

1.23 width of digit, Fig. 8A (terminal discs onfingers broadly expanded, 23 width of digit,Fig. 8C), small tubercles on heel absent(present), Toe III slightly shorter than ToeV, Fig. 8B (much shorter, Fig. 8D); Toe IIInot reaching distal subarticular tubercle onToe IV (reaching), Toe V reaching lowermargin of distal subarticular tubercle on ToeIV (overlapping distal subarticular tubercle),irregularly shaped, dark brown blotches onbody forming ‘‘interrupted’’ chevrons (clearlydefined dark chevrons), extremities with darkbrown blotches, rarely presenting diagonalbars (clearly defined diagonal bars present).

Superficially, Eleutherodactylus ornatuscan be confused with E. toftae, a member of

FIG. 7.—Lateral (A) and dorsal (B) views of head ofEleutherodactylus ornatus (MTD 45897), and lateral (C)and dorsal (D) views of head of E. rhabdolaemus(KU 173246).

FIG. 8.—Ventral views of hand (A) and foot (B) of E.ornatus (MTD 45897), and ventral views of hand (C) andfoot (D) of E. rhabdolaemus (KU 173246).


the E. unistrigatus Group, which is restrictedto forests of the Amazonian lowlands to lowerAndean foothills of southern Peru to Cocha-bamba, Bolivia (Frost, 2004). These speciesare similar in size (maximum SVL 27.4 mm infemales of E. toftae; Duellman, 2005) andboth have a long snout, dark brown canthaland supratympanic stripes, and dorsolateralfolds (absent in E. toftae according to theoriginal species description and a recentspecies account by Duellman [1978b, 2005]).However, E. ornatus has fingers and toeswithout lateral fringes (present in E. toftae),tan flanks with brown blotches (dark brown inE. toftae), and lacks a labial stripe (present inE. toftae). Eleutherodactylus ornatus and thecentral Peruvian E. aniptopalmatus havesimilar elevational distributions, smooth ven-tral skin, and dark brown canthal andsupratympanic stripes, but E. ornatus is larger(maximum SVL in females 27.3 mm vs.22.0 mm in E. aniptopalmatus; Duellmanand Hedges, 2005), has a long snout (moder-ately long in E. aniptopalmatus) and dorsolat-eral folds (absent in E. aniptopalmatus).Juvenile E. ornatus can be confused withjuvenile E. bipunctatus, a member of the E.conspicillatus Group. Both species have longsnouts, dark brown canthal and supratympa-nic stripes, dorsolateral folds, and lack lateralfringes, but E. ornatus is much smaller(maximum SVL 41.5 mm in E. bipunctatus),and lacks black scapular warts (present in E.bipunctatus). Furthermore, E. bipunctatus isrestricted to lower elevations (230–2320 m),whereas E. ornatus occurs at higher elevations(2400–3000 m).

Description of the holotype.—Head as wideas body, as wide as long; head width 36.4% ofSVL; head length 36.8% of SVL; snout long,subacuminate in dorsal view, round in lateralview (Figs. 7A, B); eye diameter nearly equalto as eye–nostril distance; nostrils slightlyprotuberant, directed dorsolaterally; canthusrostralis straigth in dorsal view, rounded inprofile; loreal region concave; lips rounded;upper eyelid without tubercles; width ofupper eyelid much narrower than IOD (uppereyelid width 58.6% of IOD); supratympanicfold prominent; tympanic membrane andtympanic annulus present, tympanum length33% of eye length, separated from eye by

distance equal to tympanum length; tympa-num slightly higher than long, its upper andposterolateral margin concealed by supratym-panic fold; two enlarged, conical postrictaltubercles present. Choanae small, ovoid, notconcealed by palatal shelf of maxilla; vomerineteeth small, oval, situated posteromedially tochoanae, each vomer bearing five teeth,vomers widely separated; tongue 1.33 as longas wide (length 6.1 mm, width at midlength oftongue 4.6 mm), not notched posteriorly,posterior half free.

Skin on dorsum finely shagreen with small,scattered tubercles; weakly defined dorsolat-eral fold on each side of body; skin on flanksfinely shagreen with scattered tuberclesslightly larger than those on dorsum; skin onproximal posteroventral surfaces of thighsweakly areolate, other ventral surfacessmooth; discoidal fold present; cloacal sheathshort; large tubercles absent in cloacal region.Ulnar and tarsal tubercles or folds absent;palmar tubercles slightly elevated, outerpalmar tubercle bifid, approximately 2.53size of ovoid, inner palmar tubercle; fewsupernumerary tubercles present, round,low, slightly smaller than subarticular tuber-cles; subarticular tubercles well defined,round in dorsal view and subconical in lateralview, most prominent on bases of fingers;fingers without lateral fringes; Finger I shorterthan Finger II; discs on fingers expanded,most prominent on Fingers III–IV, disc onFinger III nearly truncate terminally, otherdiscs on fingers rounded, disc on Finger Islightly expanded; all fingers with ventral padswell defined by circumferential grooves(Fig. 8A). Hind limbs slender, tibia length58.1% of SVL; foot length 52.6% of SVL;upper surface of hind limbs smooth; posteriorand ventral surfaces of thighs weakly areolate;heel without tubercles; tarsus without tuber-cles and fold; inner metatarsal tubercleelevated, elliptical, about 23 subconical outermetatarsal tubercle; few, low plantar super-numerary tubercles present; subarticular tu-bercles well defined, round in dorsal view,subconical in lateral view; toes without lateralfringes and webbing; discs on toes slightlysmaller than those on fingers, most prominenton Toes IV–V; toe discs well defined bycircumferential grooves; relative lengths of


toes: 1 , 2 , 3 , 5 , 4 (Fig. 8C); Toe Vslightly longer than Toe III (tip of the disc onToe III reaches middle of disc on Toe V; discon Toe III not reaching penultimate subartic-ular tubercle on Toe IV, tip of the disc on ToeV reaching middle of penultimate tubercle onToe IV).

Measurements (in mm) of holotype: SVL25.3; TL 14.7; FL 13.3; HL 9.3; HW 9.2; ED2.4; TY 0.8; IOD 2.9; EW 1.7; IND 3.3; E–N2.3.

Coloration of holotype in preservative.—Dorsum grayish tan with dark brown blotchesarranged in three ill-defined, interrupted(without contact at midline) chevrons, andnarrow, dark brown interorbital bar; darkbrown blotches on lower arm and outersurface of hand; anterior surfaces of armswith a dark brown longitudinal stripe; hindlimbs with dark brown blotches, and thighswith ill-defined, transverse, dark brown bars;dark brown canthal and supratympanicstripes; flanks slightly paler than dorsum, withdark brown blotches forming three irregulardiagonal bars; groin pale cream with small,

pale brown spots; anterior and posteriorsurfaces of thighs grayish tan with small palebrown spots; ventral surfaces cream with smalldark brown spots, most densely on throat andchest; iris black.

Coloration of holotype in life.—Unknown.Variation.—Coloration in life for female

MTD 45897 is based on photos. Dorsum tanand brown with blackish-brown blotchesforming interrupted chevrons; canthal andsupratympanic stripes blackish brown; upperhalf of flanks pale tan with blackish-brownblotches forming three irregular diagonal bars;lower half of flanks white; groin, anterior andposterior surfaces of thigh reddish brown;belly, chest, and throat whitish gray withgrayish-brown blotches more dense on throat;upper half of iris pale gold with blackreticulations, lower part with a broad, darkbrown, horizontal stripe. See Tables 5, 6 formeasurements of the type series; see Table 7for ranges and proportions.

Etymology.—The specific name is derivedfrom the Latin noun ornatus, meaning deco-rated. The name refers to the black markings

TABLE 6.—Measurements (in mm) of adult males of Eleutherodactylus ornatus.

Character MTD 44770 MTD 44769 MHNSM 19883 MHNSM 20668 MTD 44772 MHNSM 20671 MTD 44771

SVL 20.5 20.3 20.1 19.9 19.8 18.4 16.7TL 12.2 12.1 11.1 11.3 11.3 10.9 10.7FL 10.9 9.9 10.4 9.9 10.3 8.9 8.6HL 8.0 8.3 7.5 7.4 7.4 6.7 7.3HW 8.3 8.1 7.9 7.6 7.9 7.0 6.7ED 21.0 2.2 2.4 1.9 1.9 2.0 1.8TY 1.0 0.7 0.9 0.8 0.8 0.8 0.8IOD 2.4 2.1 2.5 2.2 2.1 2.1 2.1EW 1.4 1.2 1.5 1.5 1.2 1.4 1.3IND 2.6 2.4 2.4 2.3 2.6 2.2 2.2E–N 2.1 2.3 2.1 2.3 2.1 2.0 2.0

TABLE 5.—Measurements (in mm) of selected adult females of Eleutherodactylus ornatus.

Character MHNSM 20665 MTD 44768 MTD 45073 MTD 44767 MHNSM 20667 MTD 45897 MTD 45898



on the dorsum; these are reminiscent ofemblems (chevrons) used on military uni-forms.

Distribution and ecology.—This species isknown from several localities at elevations of2400–3000 m in the Provincia de Pasco(Fig. 4). Eleutherodactylus ornatus was foundat night between 1900–2300 hr on the groundor on vegetation up to 1 m above the ground.During the day, specimens were found amongleaf litter and under stones. One pair (femaleMHNSM 20665, male MTD 44771) was inaxillary amplexus on 5 March 2002. Sympatricanurans include Bufo multiverrucosus, E.lundbergi, E. mendax, E. platydactylus, Gas-trotheca stictopleura, Phrynopus juninensis,and Phyllonastes duellmani. See Figure 5 forcomparative altitudinal distributions.

Eleutherodactylus pardalinus sp. nov.

Holotype.—MHNSM 19806 (Fig. 9), anadult female obtained at Huasahuasi (11u 16906.10 S, 75u 399 28.30 W) at 2640 m on 1September 2003, Distrito de Huasahuasi,Provincia de Tarma, Departamento de Junın,Peru, by E. Lehr.

Paratopotypes.—15, all collected with theholotype by C. Aguilar, E. Lehr, and R. vonMay: 7 females (MHNSM 19807–08, MTD45666–69, 45671), 8 males (MHNSM 19809–13, MTD 45670, 45672–73).

Diagnosis.—A member of the Eleuthero-dactylus lacrimosus Assemblage of theEleutherodactylus (Eleutherodactylus) uni-strigatus Group having: (1) skin on dorsumshagreen with small tubercles, that on ventercoarsely areolate; discoidal fold not evident;dorsolateral folds absent; (2) tympanic mem-brane and tympanic annulus distinct, upperand posterolateral margins of tympanumconcealed by supratympanic fold; (3) snoutacuminate in dorsal view, rounded in lateralview; (4) upper eyelid with small tuberclesand usually one prominent tubercle at centerof eyelid; width of upper eyelid narrower thanIOD; cranial crests absent; (5) vomerine teethwidely separated, in oblique clump; (6) maleswith vocal slits and nuptial pads; (7) Finger Ishorter than Finger II; discs on outer fingersbroadly expanded, rounded; (8) fingers withbroad lateral fringes; (9) ulnar and tarsaltubercles present; (10) heel with small, low

TABLE 7.—Ranges (in mm) and proportions of adultEleutherodactylus ornatus; ranges followed by means and

one standard deviation in parentheses.

Characters Females (n 5 11) Males (n 5 7)

SVL 20.7–27.3 (24.4 6 2.3) 16.7–20.5 (19.4 6 1.3)TL 12.3–15.2 (14.1 6 0.9) 10.7–12.2 (11.4 6 0.5)FL 10.4–13.8 (12.6 6 1.0) 8.6–10.9 (9.8 6 0.8)HL 8.4–10.3 (9.3 6 0.7) 6.7–8.3 (7.5 6 0.5)HW 8.3–9.9 (9.3 6 0.6) 6.7–8.3 (7.6 6 0.5)ED 2.2–2.8 (2.5 6 0.2) 1.8–2.4 (2.0 6 0.2)TY 0.8–1.2 (1.0 6 0.2) 0.7–1.0 (0.8 6 0.1)IOD 2.9–3.3 (3.1 6 0.2) 2.1–2.5 (2.2 6 0.2)EW 1.2–1.9 (1.6 6 0.2) 1.2–1.5 (1.4 6 0.1)IND 2.2–3.3 (2.8 6 0.2) 2.2–2.6 (2.4 6 0.2)E–N 2.3–2.9 (2.6 6 0.2) 2.1–2.3 (2.1 6 0.1)TL/SVL 0.53–0.68 0.57–0.64FL/SVL 0.47–0.59 0.48–0.53HL/SVL 0.35–0.49 0.36–0.44HW/SVL 0.33–0.48 0.63–0.70HW/HL 0.88–1.08 0.92–1.07E–N/ED 0.89–1.17 0.88–1.21EW/IOD 0.38–0.64 0.57–0.68TY/ED 0.33–0.48 0.32–0.50

FIG. 9.—Lateral (A), dorsal (B), and ventral (C) views ofEleutherodactylus pardalinus (MHNSM 19806, female,SVL 34.1 mm). Photos by E. Lehr.


tubercles and one prominent tubercle; innertarsal fold absent; (11) inner metatarsaltubercle ovoid, 33 outer; conical outer meta-tarsal tubercle; many supernumerary plantartubercles present, low, diffuse; (12) toes withbroad lateral fringes; toe webbing absent; ToeV much longer than Toe III; toe discs equal insize to those on fingers; (13) dorsum yellowishtan with dark, brownish-purple blotches;flanks white; venter white; groin, anteriorand posterior surfaces of thighs yellowishtan; iris brownish red or tan with horizontal,dark brown band and with fine, blackreticulations; (14) SVL in females 26.1–34.1 mm (n 5 8), in males 21.1–26.7 mm (n5 8).

With respect to habitus and coloration,Eleutherodactylus pardalinus resembles otherbromeliad-inhabiting species in the CordilleraOriental, these are E. bromeliaceus, E. lacri-mosus, E. mendax, E. olivaceus, and E.schultei. Eleutherodactylus pardalinus isslightly smaller than E. schultei, but largerthan the other species: the maximum SVL offemale E. pardalinus is 34.1 mm, whereas thatof female E. bromeliaceus is 28.1 mm (Lynch,1979), E. olivaceus 24.2 mm (Kohler, 2000),E. lacrimosus 32.5 mm (Lynch, 1980), and E.mendax 28.1 mm (Duellman, 1990). Further-more, in E. pardalinus the dorsum is sha-green, whereas the dorsum is smooth in E.bromeliaceus. Eleutherodactylus pardalinushas robust hind limbs, whereas the hind limbsare slender in E. lacrimosus (Lynch andDuellman, 1980). Eleutherodatylus pardali-nus has white flanks, whereas E. mendax hasyellow flanks (Duellman, 1978c). Eleuthero-dactylus pardalinus has the dorsum yellowishtan with brownish-purple blotches, whereasthe dorsum is uniform olive-green in E.olivaceus.

Eleutherodactylus pardalinus is most simi-lar to E. schultei, which is known at elevationsof 2400–2850 m (Duellman, 1990) at twolocalities in northern Peru (Departamento deAmazonas). Both species are about the samesize (males: 21.1–26.7 mm in E. pardalinusand 23.5–26.6 mm in E. schultei; females:26.1–34.1 mm in E. pardalinus and 28.4–34.0 mm in E. schultei; Duellman, 1990) andhave white bellies. However, they can bedistinguished by the following features (char-

acters of E. schultei in parentheses): body andextremities robust (slender); fingers and toeswith broad lateral fringes (narrow); males withnuptial pads (absent); disc on Toe III reachesabove distal subarticular tubercle on Toe IV(not reaching above); 3–5 vomerine teeth (2–4, or absent [Duellman, 1990]); dorsal color-ation yellowish tan (dull green, greenishyellow, creamy tan, pale yellowish tan, orreddish brown [Duellman, 1990]); dorsumwith large purple blotches and streaks (dor-sum uniform, bordered by black stripes, orwith small dark flecks [Duellman, 1990]);flanks white (tan to greenish yellow [Duell-man, 1990]).

Description of the holotype.—Head nar-rower than body, slightly wider than long;head width 38.4% of SVL; head length 36.7%of SVL; snout moderately short, acuminate indorsal view, rounded in lateral view (Fig. 10A,B); eye diameter less than eye–nostril distance(eye–nostril distance 114.7% of length of eye);nostrils slightly protuberant, directed dorso-laterally; canthus rostralis straight in dorsalview, rounded in profile; loreal region slightlyconcave; lips rounded; upper eyelid bearingsmall tubercles and one prominent tubercle at

FIG. 10.—Lateral (A) and dorsal (B) views of head, andventral views of hand (C) and foot (D) of E. pardalinus(MHNSM 19806).


center of eyelid; width of upper eyelid 50% ofIOD; supratympanic fold prominent; tympan-ic annulus and tympanic membrane present,tympanum round, its length 47.1% of eyelength, separated from eye by distance aboutone-half length of tympanum, its uppermargin concealed by supratympanic fold;one enlarged, conical postrictal tubercles oneach side. Choanae ovoid, not concealed bypalatal shelf of maxillary arch; vomerine teethsmall, situated posteromedially to choanae,right vomer bearing four teeth (vomerineteeth on left side completely concealed bybuccal tissues) in an oblique row, widelyseparated; tongue 1.13 as long as wide (length6.3 mm, width at midlength of tongue5.6 mm), slightly notched posteriorly, posteri-or third free. Skin on dorsum shagreen withsmall tubercles; dorsolateral folds absent; skinon upper half of flanks same as dorsum,venterolaterally coarsely areolate; skin onthighs, belly, and chest coarsely areolate, skinon other ventral surfaces smooth; discoidalfold not evident; cloacal sheath short; no largetubercles in cloacal region.

Venterolateral surface of forearm bearingrow of low tubercles; palmar tubercles slightlyelevated, outer palmar tubercle bifid, approx-imately 43 the size of narrow, ovoid, innerpalmar tubercle; numerous low round super-numerary tubercles, slightly to much smallerthan subarticular tubercles; subarticular tu-bercles well defined, round in dorsal view andsubconical in lateral view; fingers with broadlateral fringes, outer fringe of Finger IVextends to outer edge of palm with its shapeslightly undulated; Finger I shorter thanFinger II; discs on fingers broadly expanded,most prominent on Fingers III–IV, roundedterminally; all fingers with ventral padswell defined by circumferential grooves(Fig. 10C). Hind limbs robust, tibia length48.1% of SVL; foot length 47.5% of SVL;upper surfaces of hind limbs shagreen withsmall tubercles; posterior and ventral surfacesof thighs coarsely areolate; heel bearing small,low, round tubercles, two on each heel moreprominent; inner ventral surface of tarsus withrow of low tubercles partially coalesced intofold; inner metatarsal tubercle elevated, elon-gate, about 2.33 subconical outer metatarsaltubercle; numerous, low plantar supernumer-

ary tubercles; subarticular tubercles well de-fined, round in dorsal view and subconical inlateral view; toes with broad lateral fringes; toewebbing absent; discs on toes equal to thoseon fingers, most prominent on Toes IV–V, toediscs well defined by circumferential grooves;relative lengths of toes: 1 , 2 , 3 , 5 , 4(Fig. 10D); Toe V much longer than Toe III(disc on Toe III not reaching distal subar-ticular tubercle on Toe IV, disc on Toe Vextending to distal subarticular tubercle onToe IV).

Measurements (in mm) of holotype: SVL34.1; TL 16.4; FL 16.2; HL 12.5; HW 13.1;ED 3.4; TY 1.6; IOD 4.8; EW 2.4; IND 2.3;E–N 3.9.

Coloration of holotype in preservative.—Dorsum cream with dark brown blotches;longitudinal, dark brown streak on each sideof neck; narrow, discontinuous, dark browninterorbital bar; dark brown canthal andsupratympanic stripes; flanks cream; few palebrown blotches present on arms and thighs;shanks and outer surface of feet with manyblotches; belly grayish cream, other ventralsurfaces uniformly cream.

Coloration of holotype in life.—Dorsumyellowish tan with dark brownish-purpleblotches and streaks as described above; darkbrownish-purple blotches on extremities;flanks white; chest and belly white with palegray interspaces between granules; otherventral surfaces yellow to flesh-colored; irisbrownish red, with horizontal dark brownband and fine black reticulations.

Variation.—Individuals differ in the num-ber of dorsal blotches and streaks. In males,the dorsum is less intensively blotched thanin females; moreover, the anterior half ofdorsum in males has many blotches, whereasthe posterior half has fewer blotches. Maleshave white, nonspinous nuptial pads dorsolat-erally on the bases of the thumbs. SeeTables 8, 9 for measurements of the typeseries; see Table 10 for ranges and propor-tions.

Etymology.—The specific name is the di-minutive of the Greek noun pardos, meaningleopard or panther. The name refers to thedorsal pattern of dark blotches on a yellowish-tan ground color, which reminds us ofa leopard.


Distribution and ecology.—Eleutherodacty-lus pardalinus is known only from the typelocality, next to the village Huasahuasi(Fig. 4), which belongs to the Puna Ecore-

gion. All specimens were found during theearly afternoon in terrestrial bromeliads sur-rounded by the Peruvian Feather Grass (Stipaichu) on a hillside. No other anurans werefound there. See Figure 5 for comparativealtitudinal distributions.

COMMENTS ON CENTRAL

PERUVIAN ELEUTHERODACTYLUS

Eleutherodactylus bipunctatus Duellman andHedges 2005

Eleutherodactylus bipunctatus in the E.conspicillatus Group was described on thebasis of one adult female; seven juveniles werereferred to the species. The following data arefrom 36 specimens (Appendix) collectedbetween 1997 and 2004. SVL in adult females35.0–41.5 mm (n 5 10), in adult males 22.6–28.8 (n 5 9); the smallest of 17 juveniles(SMF 80400) is 11.4 mm. In addition to thesmall tubercles in the scapular region notedby Duellman and Hedges (2005), all speci-mens have a pair of black scapular warts.Duellman and Hedges (2005) mentioned only

TABLE 8.—Measurements (in mm) of selected females of Eleutherodactylus pardalinus.

Character MHNSM 19806 MHNSM 19807 MTD 45666 MTD 45667 MTD 45668 MHNSM 19808 MTD 45669


TABLE 9.—Measurements (in mm) of selected males of Eleutherodactylus pardalinus.

Character MTD 45672 MHNSM 19809 MHNSM 19810 MHNSM 19812 MHNSM 19813 MTD 45673 MHNSM 19811


TABLE 10.—Ranges (in mm) and proportions of adultEleutherodactylus pardalinus; ranges followed by means

and one standard deviation in parentheses.

Characters Females (n 5 6) Males (n 5 8)

SVL 29.4–34.1 (31.9 6 1.5) 21.7–26.7 (23.6 6 1.9)TL 16.2–17.1 (16. 6 7 0.4) 12.0–13.4 (13.0 6 0.6)FL 14.5–16.4 (15.5 6 0.7) 10.1–12.3 (11.3 6 0.9)HL 11.0–12.5 (11.9 6 0.6) 7.8–9.6 (8.7 6 0.6)HW 11.0–13.1 (12.1 6 0.7) 8.1–10.1(9.1 6 0.7)ED 2.7–3.4 (3.0 6 0.2) 2.1–2.9 (2.4 6 0.2)TY 1.2–1.7 (1.5 6 0.2) 0.7–1.0 (1.0 6 0.1)IOD 4.1–4.8 (4.5 6 0.2) 2.9–3.9 (3.4 6 0.3)EW 2.2–2.6 (2.4 6 0.1) 1.7–2.1 (1.9 6 0.2)IND 2.2–3.3 (2.2 6 0.1) 1.7–2.1 (1.9 6 0.1)E–N 2.1–2.3 (3.6 6 0.2) 2.4–3.1 (2.8 6 0.2)TL/SVL 0.48–0.58 0.50–0.61FL/SVL 0.47–0.51 0.45–0.52HL/SVL 0.35–0.40 0.34–0.41HW/SVL 0.35–0.40 0.37–0.41HW/HL 1.00–1.05 1.00–1.07E–N/ED 1.10–1.30 0.96–1.33EW/IOD 0.46–0.63 0.47–0.66TY/ED 0.44–0.57 0.32–0.50


a pair of black dots, but not the presence ofwarts. Males have vocal slits and a dorsalnuptial pad near the base of the thumb(nuptial pads absent according to Duellmanand Hedges, 2005). The thumb is remarkablythickened in several specimens without ap-parently correlation to climatic season (dis-cussed below).

Distribution and ecology.—Previously knownonly from the type locality at 2120 m inDepartamento de Pasco (Duellman andHedges, 2005). The species is widely distribut-ed in lowland and cloud forests (230–2320 m)in central Peruvian Departamentos Pasco,Junın, and Ucayali. Specimens from Pozuzoand Bolognesi previously were misidentified asE. peruvianus (Lehr, 2002, 2005). Individualswere found at night on vegetation 5–200 cmabove ground (Lehr, 2002). Five males (collect-ing dates in parentheses) have thickenedthumbs: MHNSM 20375 (22 Jan. 1999),MTD 45920 (06 Dec. 2003), 45694 (25 May2003), 46484 (11 Dec. 2004), SMF 80398 (23Jan. 1999); the following males lack thickenedthumbs: MTD 46472 (27 Nov. 2004), MHNSM19828 (25 May 2003), MHNSM 20371 (16 Jul.1998), SMF 80394 (12 Jul. 1998). A female(MTD 46483, 11 Dec. 2004) had 63 ovarianeggs (27 left, 36 right) with an average diameterof 2.6 6 0.12 mm (n 5 10). Sympatric speciesof Eleutherodactylus at lower elevations(230 m) include E. eurydactylus and E. rhab-docnemus, and those at higher elevations(above 1500 m) include E. cruciocularis, E.flavobracatus, and E. rhabdocnemus.

Eleutherodactylus bromeliaceus Lynch 1979

Morales and Icochea (2000) recordedEleutherodactylus bromeliaceus in northernand central Peru, and Duellman and Pramuk(1999) in northern Peru. In central Peru, wefound E. bromeliaceus in arboreal bromeliadsat elevations of 2180–2780 m. Males havea SVL of 18.0–22.6 mm (n 5 8), and onefemale (MHNSM 18660) has a SVL of18.1 mm. The coloration in life of MHNSM18660 was recorded by E. Lehr on 13February 2003: ‘‘Dorsally pale olive, withbrown blotches dorsolaterally; broad, salm-on-colored band between eyes borderedposteriorly and anteriorly by a dark brownstripe, dark brown canthal and supratympanic

stripe; upper lip dark brown, above creamstripe from tympanum to naris; throat, chest,and belly pale gray, extremities pale olive; irisreddish with black reticulations.’’ SympatricEleutherodactylus in central Peru include E.rhabdocnemus and E. sagittulus.

Eleutherodactylus eurydactylus Hedges andSchluter 1992

This species was described from Panguana,Departamento de Huanuco, Peru, 200 m andreported from up to 1300 m elevation in theSerranıa de Sira (Hedges and Schluter, 1992).A specimen (MHNSM 20374) from Bolognesi(10u 069 130 S, 73u 499 020 W, 230 m) is thefirst record of this species from Departamentode Ucayali. This individual previously wasmisidentified as E. diadematus (Lehr 2001a,2002). Sympatric Eleutherodactylus include E.bipunctatus and E. rhabdocnemus.

Eleutherodactylus mendax Duellman 1978

Morales and Icochea (2000) supplementedthe distributional information about Eleuther-odactylus mendax in Peru. In the Departa-mento de Pasco, we found E. mendax (E.lacrimosus in Lehr, 2005) at elevations of2850–3325 m. Individuals were found by dayin arboreal or terrestrial bromeliads, andmales were found calling at night frombamboo. Males have a SVL of 21.6–23.3 mm(n 5 4), whereas females have a SVL of 22.3–26.0 mm (n 5 3). The coloration in life ofa male (MTD 45900) was recorded by E. Lehron 10 December 2003: ‘‘Dorsally and laterallyyellow, pale greenish yellow behind eyes,dorsum with fine black blotches, tips offingers and toes pale red, belly white, irisreddish copper with black reticulations.’’Sympatric anurans include Bufo multiverru-cosus, E. lundbergi, E. ornatus, and E.platydactylus, Gastrotheca stictopleura, Phry-nopus juninensis, and Phyllonastes duellmani.

Eleutherodactylus platydactylus(Boulenger 1903)

Several authors (De la Riva, 1997; De laRiva et al., 2000; Kohler and Lotters, 1999)have reported that Eleutherodactylus platy-dactylus is highly polymorphic and mayrepresent several sibling species. The speciesis known at elevations of 950–3470 m from


central Peru to Bolivia (Kohler, 2000). De laRiva (1997) redescribed E. platydactylus,noted 10 distinct color morphs, and diagnosedthree putative new taxa for Bolivia. Onespecies (E. llojsintuta), syntopic with E.platydactylus, was described as new mainlybased on distinct advertisement calls (Kohlerand Lotters, 1999).

Herein we present the northernmost distri-butional records for the species in the centralPeruvian Departamento de Pasco where wefound E. platydactylus at elevations of 2500–3275 m. At night, they were on low vegetationand on moist moss. By day they were foundunder dry leaves on the ground and interrestrial bromeliads. Females have SVLs of24.4–35.3 mm (n 5 11), and males have SVLsof 20.0–23.8 mm (n 5 8). Another series ofspecimens tentatively referred to E. platydac-tylus was obtained from localities in Departa-mento de Junın (Pampa Hermosa, and belowPampa Hermosa near Rıo Ulcumayo), whereit was abundant at night on low vegetation inlower montane forests at elevations of 1240–1540 m. These frogs are smaller than speci-mens from Pasco—SVL in females 28.6–20.6 mm (n 5 11); SVL in males 20.0–16.0 mm (n 5 12). The dorsum is yellowishbrown, in contrast to grayish brown in E.platydactylus from Pasco. Sympatric anuranswith E. platydactylus include Bufo multi-verrucosus, Eleutherodactylus lundbergi, E.mendax, E. ornatus, Gastrotheca stictopleura,Phrynopus juninensis and Phyllonastes duell-mani; sympatric with E. cf. platydactylus areE. bipunctatus, E. cruciocularis, and Gastro-theca atympana.

Eleutherodactylus rhabdocnemus Duellmanand Hedges 2005

Duellman and Hedges (2005) described E.rhabdocnemus based on two adult femalesfrom Departamento de Pasco, 2.9 km N and5.5 km E Oxapampa, 2600 m. Herein, werefer additional specimens to E. rhabdocne-mus obtained from Departamentos Pasco andUcayali (see Appendix). Eleutherodactylusrhabdocnemus is highly variable in color andpattern, and we recognize remarkable mor-phological similarities with E. lirellus. Furtherresearch is necessary to assess the validity ofE. rhabdocnemus.

Sympatric anurans at the localities inDepartamento de Pasco include Adenomerasp., Atelopus sp., Bufo cf. margaritifer,Eleutherodactylus bipunctatus, E. flavobraca-tus, Gastrotheca testudinaea, Hyalinobatra-chium bergeri, Hybsiboas cf. melanopleura,Leptodactylus rhodonotus, and Oreobatesquixensis.

Eleutherodactylus stictogaster Duellman andHedges 2005

This species was described by Duellmanand Hedges (2005) based on a single subadultfemale with a SVL of 20.5 mm from nearOxapampa, 2600 m, We report the secondknown specimen (MTD 46474), an adultfemale with ovarian eggs, from Marıa Teresa(10u 429 05.60 S, 75u 279 22.20 W, 1470 m),with the following measurements: SVL 22.0;TL 11.9; FL 9.7; HL 9.5; HW 8.8; ED 2.5;IOD 2.8; EW 1.4; IND 2.7; E–N 2.0.Morphologically, the specimen is like theholotype. In life, the iris was gold with blackreticulations, and the groin and anteriorsurfaces of thighs pale red. Males of thisspecies are unknown. Sympatric anurans atMarıa Teresa include Bufo cf. margaritifer, E.bipunctatus, E. rhabdocnemus, Gastrothecatestudinaea, Hyalinobatrachium bergeri, Hyb-siboas cf. melanopleura, Oreobates quixensis,and Leptodactylus rhodonotus.

DISCUSSION

Currently, 91 species of Eleutherodactylusare known from Peru (updated from Frost[2004]); these represent about 25% of the 376species of anurans known to inhabit Peru(AmphibiaWeb, 2005). Because of extensivefieldwork in the Departamento de Pasco(localities around Oxapampa, Pozuzo, andPaucartambo), the species diversity of thisarea is relatively well known. However,because herpetological surveys have not beenconducted in many areas in Peru, the numberof species of Eleutherodactylus undoubtedlywill continue to increase including additionallocalities and new species. Species descrip-tions of Eleutherodactylus should be detailedand include photos (in life), as well asdrawings of the head (at least in lateral view),foot, and hand in ventral aspect so as to


facilitate further studies, comparisons andidentifications of the species.

In central Peru, Andean species ofEleutherodactylus (Fig. 5) can be foundbetween 200 and 3325 m elevation; the singlespecies of Phyllonastes occurs at 2900 m, andPhrynopus occur between 2600 and 4390 m.Two species, E. bipunctatus and E. rhabdoc-nemus, have relatively wide elevational (230–2320 m in E. bipunctatus; 230–2600 m in E.rhabdocnemus) and geographical distribu-tions, but most species of Eleutherodactylushave limited vertical and spatial distributions.Except for Phrynopus kauneorum and P.montium, most of the species of Phrynopushave even more limited vertical and spatialdistribution and in many cases are known onlyfrom their type localities. Between 1000 and2600 m the number of species of eleuther-odactyline frogs increases from four at1000 m, five at 1400 m, six at 2200 m, toa maximum of eight at 2600 m. Above 2600 melevation the species number decreases exceptfor 3250 m where the species number slightlyincreases to six from five at 3000 m. Thegreatest density of species of Eleutherodacty-lus has been recorded by Lynch and Duell-man (1997) between 2500 and 2700 in theEastern Andes of Ecuador. Thus, this eleva-tional range in the eastern Andean slopesseems to provide optimal life conditions (e. g.humidity) for Eleutherodactylus.

Among the 29 eleutherodactyline frogsreferred to in Figure 5, 16 species (55%) lacka tympanum. These include all species ofPhrynopus (Lehr et al., 2005a, b), the singlePhyllonastes, and three species of Eleuther-odactylus. The absence of a tympanum incentral Peruvian eleutherodactyline frogsseems to be associated with increasing eleva-tion—25% lack a tympanum at elevationsbelow 1000 m, 33% at 1000–2000 m, 36% at2000–3000 m, 77% at 3000–4000 m, and100% above 4000 m. The way these frogscommunicate or hear, if at all, is unknown. Nocalls have been recorded during fieldwork.Other Andean anurans such as Bufo spinulo-sus (Bufonidae) and Pleurodema marmorata(Leptodactylidae) are known from high eleva-tions (up to 4175 m in B. spinulosus; up to4480 m in P. marmorata) in the Andes ofcentral Peru (Lehr, 2002) and both species

have a tympanum and vocalization. But B.spinulosus and P. marmoratum produce freeswimming-tadpoles whereas eleutherodacty-line frogs have direct development. Thus,reproductive mode may be associated with theacoustic abilities.

Secondary sex characters in male Eleutheo-dactylus include vocal apparatus (vocal slitsand vocal sac) and nuptial pads (Lynch andDuellman, 1997). Depending on the species,male Eleutherodactylus have nuptial padspresent or absent. If present, nuptial padsare usually located basally on the first fingercovering its dorsal and lateral surfaces in-cluding outer parts of the inner palmar andsubarticular tubercles (e.g. as in many speciesof the E. conspicillatus Group). In somespecies (e.g. E. bipunctatus) nuptial pads arethickened on Finger I. Compared to otheranurans (e.g. bufonids or hylids) males ofwhich have brown and spinous nuptial padsthat are clearly distinct from the surroundingskin, nuptial pads in Eleutherodactylus areusually nonspinous and glandular, usuallywhite or cream colored and less pigmentedthan the surrounding skin. Thus, nuptial padsin Eleutherodactylus can easily be overseen,especially in badly preserved or old material.The use of a microscope with high magnifica-tion often is necessary to detect nuptial pads.It is unknown whether nuptial pads arepermanent in adult male Eleutherodactylusor occur only during reproductive seasons.This phenomenon needs study of large seriesof specimens from throughout the year. Atleast in the specimens of E. bipunctatusstudied herein, nuptial pads seem to bepermanently present, possibly indicating thatthis species reproduces throughout the year.Therefore, nuptial pads are good characters toseparate sexes and are helpful to distinguishspecies.

RESUMEN

Se describen cuatro especies nuevas proce-dentes de bosques montanos en los Departa-mentos de Huanuco, Junın y Pasco en Perucentral, con elevaciones que van desde los1330–3000 m en la Cordillera Oriental. Tresde las especies nuevas se asignan al grupo deEleutherodactylus unistrigatus. Dos de ellascarecen de tımpano, pero presentan una


coloracion y patron de iris diferentes: unatiene la ingle y los miembros posterioresparcialmente coloreados de anaranjado a rojo,y el iris con una raya vertical parda y otrahorizontal negra formando una cruz, mientrasque la otra tiene la ingle y las patas posterioresparcialmente amarillas, y el iris con una rayahorizontal parda y otra vertical negra desde lapupila al margen inferior del ojo, formandouna ‘‘T.’’ La tercera especie, asignada a la seriede Eleutherodactylus conspicillatus, es similara E. rhabdolaemus, pero difiere de esta encaracteres morfologicos y de coloracion. Lacuarta especie asignada a la agrupacion de E.lacrimosus tiene el dorso tabaco amarillentocon manchas y rayas oscuras pardo purpureo,con flancos y vientre blancos, habita brome-liaceas terrestres, y es comparada con E.schultei del norte de Peru. Se anaden datosmorfologicos y de distribucion para las sieteespecies recientemente descritas de Eleuther-odactylus del Peru central. Se compara ladistribucion altitudinal entre los Eleuthero-dactylus, Phyllonastes, y Phrynopus de Perucentral.

Acknowledgments.—For comments on the manuscriptwe are grateful to R. Diaz, W. E. Duellman, and L. Trueb.We thank J. R. McCranie, I. De la Riva, and oneanonymous reviewer for helpful comments on themanuscript. Specimens were loaned by J. H. Cordova(MHNSM), U. Fritz (MTD), G. Kohler (SMF), and L.Trueb (KU). EL thanks D. Frost (AMNH), D. Kizirian(AMNH) and the American Museum of Natural History,New York, for their support and for providing a CollectionStudy Grant. C. W. Myers (AMNH) provided photos anda color in life description for a specimen referred to E.cruciocularis. The research was supported by a post-doctoral grant given to E. Lehr by the Alexander vonHumboldt-Foundation. We thank R. Acero for collectingand export permits issued by the Ministerio de Agricul-tura (INRENA), Lima, Peru.

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LEHR, E., G. FRITZSCH, AND A. MULLER. 2005b. An analysisof Andes frogs (Phrynopus, Leptodactylidae, Anura)phylogeny based on 12S and 16S mitochondrial rDNAsequences. Zoologica Scripta 34:593–603.

LEHR, E., G. KOHLER, AND E. PONCE. 2000. A newspecies of Phrynopus from Peru (Amphibia, Anura,Leptodactylidae). Senckenbergiana biologica 80:205–212.

LEHR, E., M. LUNDBERG, AND C. AGUILAR. 2005a. Threenew species of Phrynopus from central Peru (Amphib-ia, Anura, Leptodactylidae). Copeia 2005:479–491.

LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E.DAWSON. 1985. Standards in herpetology and ichthyol-ogy: part I. Standard symbolic codes for institutionalresource collections in herpetology and ichthyology.Copeia 1985:802–832.

LYNCH, J. D. 1979. Leptodactylid frogs of the genusEleutherodactylus from the Andes of southern Ecua-dor. The University of Kansas, Museum of NaturalHistory, Miscellaneous Publication 66:1–62.

———. 1980. A taxonomic and distributional synopsis ofthe Amazonian frogs of the genus Eleutherodactylus.American Museum Novitates 2696:1–24.

LYNCH, J. D., AND W. E. DUELLMAN. 1980. TheEleutherodactylus of the Amazonian slopes of theEcuadorian Andes (Anura: Leptodactylidae). TheUniversity of Kansas, Museum of Natural History,Miscellaneous Publication 69:1–86.

———. 1997. Frogs of the genus Eleutherodactylus inwestern Ecuador. Systematics, ecology, and biogeogra-phy. The University of Kansas, Natural HistoryMuseum Special Publication 23:1–236.

MORALES, V. R., AND J. ICOCHEA. 2000. Review of the typematerial of Eleutherodactylus mendax and a new recordof Eleutherodactylus bromeliaceus from Peru. Journalof Herpetology 34:158–160.

APPENDIX I

Comparative Specimens Examined

Eleutherodactylus altamazonicus.—ECUADOR: NAPO:Santa Cecilia, 340 m: KU 146060–62, 148722–84; PERU:HUANUCO: Finca Panguana, Rıo Llullapichis, 4–5 kmupstream from Rıo Pachitea, ca. 200 m: KU 154759;MADRE DE DIOS: 15 km E Puerto Maldonado, 200 m: KU209956, 215459–60.

Eleutherodactylus aniptopalmatus.—PERU: PASCO:2.9 km north and 5.5 km east (airline) of Oxapampa,2600 m: KU 291626 (holotype), 291627, 291631 (para-types); PASCO: ca. 12 km east of Oxapampa, Cumbre deOllon, 2300 m: KU 206102 (paratype).

Eleutherodactylus bipunctatus.—PERU: PASCO: 0.7 kmS, 4.5 km E (airline) Oxapampa: KU 291638 (holotype);PASCO: Provincia de Pasco, Distrito de Paucartambo: km85.2 (near Milagro) on road from Paucartambo to Yaupi,10u 439 42.80 S, 75u 359 51.60 W, 1800 m: MHNSM 18014,18019, 18021, MTD 45920; PASCO: Provincia de Oxa-pampa, Distrito de Chontabamba: km 24 on road fromOxapampa to Yaupi, 10u 439 39.00 S, 75u 279 57.60 W,1530 m: MHNSM 19827–31, 19885, MTD 45694–97;PASCO: Provincia de Oxapampa, Distrito de Chontabamba:Marıa Teresa, 10u 429 05.60 S, 75u 279 22.20 W, 1470 m:MTD 46335–36, 46472–73, 46483–84; PASCO: Provinciade Oxapampa: on road to Laguna San Daniel, 2320 m,near Parque Nacional Yanachaga-Chemillen: MTD46343; PASCO: Provincia de Oxapampa, Distrito dePalcazu: Iscozacın (Cordillera San Matıas), 10u 109 410S, 75u 079 540 W, 410 m: SMF 80400; PASCO: Provincia deOxapampa, Distrito de Pozuzo: Pozuzo, 10u 049 100 S, 75u319 420 W, 940–1000 m: MHNSM 20371–73, SMF80392–95; PASCO: Provincia de Oxapampa, Distrito dePozuzo: Pozuzo (Yulitunqui), 10u 109 230 S, 75u 359 130 W,1000 m: SMF 80396–97; JUNIN: Provincia de Tarma,Distrito de Huasahuasi: Pampa Hermosa, 10u 599 33.30 S,75u 259 58.00 W, 1540 m: MTD 45645, 46366; UCAYALI:Provincia de Atalaya, Distrito de Tahuania: Bolognesi(Campamento), 10u 069 130 S, 73u 499 020 W, 230 m:MHNSM 20375–76, SMF 80398.

Eleutherodactylus bromeliaceus.—ECUADOR: MORO-

NA-SANTIAGO: Mirador: KU 174524 (paratype); MORONA-SANTIAGO: El Cruzado: KU 174525 (paratype); PERU: SAN

MARTIN: Rioja, E slope Abra Pardo de Miguel, 2180 m:KU 212213–4; PASCO: 2.9 km N, 5.5 km E (airline)Oxapampa: KU 291657–58, 291701–03; PASCO: 2.5 kmN, 11 km E (airline) Oxapampa: KU 291704–05; PASCO:ca. 8 km SW La Suiza on E slope: KU 291706; PASCO:Oxapampa: Antenna de Pajonal, 10u 399 300 S, 75u 179 520W, 2780 m: MTD 46309–11; PASCO: Oxapampa: SanAlberto, ca. 10u 329 530 S, 75u 229 170 W, 2200 m:MHNSM 18660, 19945–46, MTD 45210–11, 45987.

Eleutherodactylus colodactylus.—PERU: PIURA: sum-mit cordillera between Canchaque and Huancabamba,3100 m: KU 135494, 135496–501; PIURA: 31 km WHuancabamba, 3080 m: KU 181262–64; PIURA: 33 km WHuancabamba, ca. 3050 m: KU 196443–61.

Eleutherodactylus diadematus.—PERU: PASCO: Po-zuzo, 10u 049 100 S, 75u 319 420 W, 970 m: MHNSM20383–84, SMF 80410; PASCO: Pozuzo, 10u 089 560 S, 75u309 550 W, 1110 m: MHNSM 20392; PASCO: Pozuzo:Yulitunqui, 10u 109 230 S, 75u 359 130 W, 930 m: MHNSM20393.


Eleutherodactylus eurydactylus.—PERU: HUANUCO:Panguana, 200 m: KU 218292 (paratopotype); UCAYALI:Bolognesi (Campamento), 10u 069 130 S, 73u 499 020 W,230 m MHNSM 20374.

Eleutherodactylus imitatrix.—PERU: MADRE DE DIOS:Puerto Maldonado, 200 m: KU 205139–41, 207709–14,215474–76, 215478.

Eleutherodactylus lacrimosus.—ECUADOR: NAPO:Isla Anaconda, Rıo Napo, 25 km down Lago Agrio: KU202406, 297131; PASTAZA: Mera, 1140 m: KU 119513–17,119522–23, 120247–49, 132631–32, 177361–82; PASTAZA:Rıo Alpayacu, 1 km E Merca, 1100 m: 119518–21;PASTAZA: Puyo, 960 m: KU 146100; PERU: CUZCO:40 km SE Quincemil: KU 175100–1;

Eleutherodactylus lirellus.—PERU: SAN MARTIN: Rioja,Rıo Cerranayacu, 76 km NW Roja, 1200 m: KU 212226–31, 212238–9 (all paratypes); SAN MARTIN: Lamas: W slopeAbra Tangarana, 7 km NW San Juan Pacaysapa, 1080 m:KU 212240 (holotype), KU 212241, 212250 (paratypes);HUANUCO: Rıo Llullapichis, 4–5 km upstream from RıoPachitea, 200 m: KU 171893–94.

Eleutherodactylus mendax.—PERU: CUZCO: Pistipata,Rıo Umasbamba, 12 km SE Huyro, 1820 m: KU 173234(holotype), KU 173235 (paratype); CUZCO: 4 km SW SantaIsabel, 1700 m: KU 162290 (paratype); PASCO: Auquimarca,Puqiopata forest, 10u 43,759 S, 75u 42,809 W, 3310 m: MTD44329; PASCO: Auquimarca, Puqiopata forest, 10u 43,789 S,75u 42,839 W, 3325 m: MTD 44330; PASCO: Auquimarca:Cillapata, 2900 m: MTD 45080; PASCO: near road fromUchuhuerta to Auquimarca, 10u 399 16.70 S, 75u 499 32.50W, 2850 m: MHNSM 19865; PASCO: near road fromUchuhuerta to Auquimarca, 10u 429 12.50 S, 75u 449 33.90W, 3000 m: MHNSM 19866, MTD 45899–00; PASCO:Paucartambo, 2932 m: MTD 44381–82, 44773.

Eleutherodactylus platydactylus.—PERU: PASCO: Pau-cartambo: Puquiopata, 10u 43, 709 S, 75u 42,759 W,3275 m: MHNSM 20691; PASCO: Paucartambo: Auqui-marca, 2800 m: MTD 44763; PASCO: Paucartambo:Cillapata, 10 u499 31.00 S, 75u 389 57.70 W, 2900 m:MHNSM 20662, MTD 44764, 45077; PASCO: Huachon:Puagmaray, 10u 389 55.70 S, 75u 469 40.10 W, 2540 m:MTD 45083; PASCO: Paucartambo: near Auquimarca,2900 m: MTD 45718; PASCO: Paucartambo: near Auqui-marca, 2650 m: MHNSM 19861, MTD 45723; PASCO:Paucartambo: near road from Uchuhuerta to Auquimarca,10u 409 25.80 S, 75u 459 46.90 W, 2850 m: MTD 45893;PASCO: Paucartambo: near road from Uchuhuerta toAuquimarca, 10u 419 05.60 S, 75u 459 28.70 W, 2900 m:MHNSM 19862–64, MTD 45894–96; PASCO: Paucar-tambo: near road from Auquimarca to Uchuhuerta, 10u439 56.70 S, 75u 429 00.40 W, 2840 m: MTD 45901; PASCO:Huachon: Uchuhuerta, 2500 m: KU 291685, MHNSM19874, MTD 45911; PASCO: Paucartambo: Chinche,3000 m: MHNSM 20663.

Eleutherodactylus cf. platydactylus.—PERU: JUNIN:Tarma: Huasahuasi: Pampa Hermosa, ca. 10u 599 33.30S, 75u 259 58.00 W, 1540 m: MHNSM 18693–97, 19801,

20692–98, MTD 45645–59; JUNIN: Chanchamayo: belowPampa Hermosa, near Rıo Ulcumayo, ca. 11u 009 30.40 S,75u 259 36.00 W, 1240 m: MHNSM 19802–03, MTD45660–61.

Eleutherodactylus rhabdocnemus.—PERU: PASCO: 2.9 kmnorth and 5.5 km east (airline) of Oxapampa, 10u 329

380 S, 75u 219 100 W, 2600 m: KU 291646 (holotype),291651 (paratype), 291647–291650, 291652–291656;PASCO: Oxapampa: Chontabamba, km 24 on road fromOxapampa–Yaupi, 10u 439 39.00 S, 75u 279 57.60 W,1530 m: MHNSM 19834, 19835, 19836–39, 19841–46,19851–55, 19884, MTD 45700–02, 45703, 45704, 45705,45706–13, 45719–22, 46318, 46320–21; PASCO: Oxapampa:km 34 on road from Oxapampa–Yaupi, 1770 m: MHNSM19847, 19849–50, MTD 45714–15; PASCO: Oxapampa:between Oxapampa and Yaupi, 10u 449 530 S, 75u 289 370

W, 1600 m: MTD 45222; PASCO: Oxapampa, on trail toLaguna San Daniel, 2400 m: MTD 46344; PASCO:Oxapampa: San Alberto, ca. 10u 329 53.40 S, 75u 229

16.60 W, 2200 m: MHNSM 18657, 18659, MTD 45205,45212; PASCO: Oxapampa: Chontabamba: Marıa Teresa,10u 429 05.60 S, 75u 279 22.20 W, 1470 m: MHNSM 19551,MTD 46470–71, 46475–82; PASCO: Oxapampa: km 53 onroad from Oxapampa–Pozuzo, 10u 139 38.30 S, 75u 349

49.20 W, 1290 m: MTD 46317, 46323–25; PASCO:Oxapampa: km 59 on road from Oxapampa–Pozuzo, 10u119 00.50 S, 75u 349 21.10 W, 1050 m: MTD 46322; PASCO:Pozuzo, 10u 049 100 S, 75u 319 420 W, 970 m: MHNSM20378–82, 20388, 20390–91, MTD 41698, SMF 80401–03, 80405–06, 80408–09, 80415–16; PASCO: Pozuzo:Yulitunqui, 10u 109 230 S, 75u 359 130 W, 930 m: MHNSM20394, SMF 8417; PASCO: Pozuzo, 10u 089 560 S, 75u 309

550 W, 1110 m: SMF 80411–14; UCAYALI: CampamentoBolognesi, 10u 069 130 S, 73u 499 020 W, 230 m: SMF80399.

Eleutherodactylus rhabdolaemus.—BOLIVIA: COCHA-

BAMBA: 68.8 km SW Villa Tumari, 1860 m: KU 183009;PERU: CUZCO: 7 km Santa Isabel, 1900 m: KU 13887(paratype); AYACUCHO: Huanhuachayocc, on Tambotrail: KU 175082–83 (paratypes); AYACUCHO: betweenMitupucuru and Estero Ruana: KU 175084 (paratype);CUZCO: Buenos Aires, 2400 m: KU 173237–53 (5paratypes of E. pharangobates); CUZCO: Huyro, 1720 m:KU 175086–88.

Eleutherodactylus schultei.—PERU: AMAZONAS: 5 kmN Levanto: 06u 179 s, 77u 519, 2850 m: KU 212222(holotype), KU 209496–97 (paratypes), KU 212220–24(paratypes); AMAZONAS: ca. 5 km NW Mendoza, ca.2400 m: KU 209498–504 (paratypes).

Eleutherodactylus stictogaster.—PERU: PASCO: 2.9 kmN, 5.5 km E (airline) Oxapampa: KU 291659 (holotype);PASCO: Oxapampa: Chontabamba: Marıa Teresa, 10u 429

05.60 S, 75u 279 22.20 W, 1470 m: MTD 46474.Eleutherodactylus toftae.—PERU: HUANUCO: Rıo Llu-

llapichis, 4–5 km upstream from Rıo Pachitea, ca. 200 m:KU 154806–19, 171852–62, 171864–66.


Species Altitudinal range (m above sea level) Literature

E. aniptopalmatus 2300–2600 Duellman and Hedges, 2005E. bipunctatus 230–2320 this paperE. bromeliaceus 2180–2780 this paperE. cruciocularis 1540 this paperE. diadematus 930–1110 this paperE. eurydactylus 200–1300 Hedges and Schluter, 1992E. flavobracatus 1770 this paperE. lundbergi 1800–2760 Lehr, 2005E. mendax 2850–3325 this paperE. ornatus 2550–3000 this paperE. pardalinus 2640 this paperE. platydactylus 2500–3275 this paperE. cf. platydactlus 1240–1540 this paperE. rhabdocnemus 230–2600 Duellman and Hedges, 2005; this paperE. sagittulus 2050–2200 Lehr et al., 2004bE. stictogaster 1470–2600 Duellman and Hedges, 2005; this paperPhyllonastes duellmani 2900 Lehr et al., 2004aPhrynopus barthlenae 3425–3680 Lehr and Aguilar, 2002; this paperP. bracki 2600 Hedges, 1990P. bufoides 3850–4100 Lehr et al., 2005aP. dagmarae 3020–3380 Lehr et al., 2002aP. heimorum 3420–3430 Lehr, 2001bP. horstpauli 3030–3430 Lehr et al., 2000P. juninensis 3650 Cannatella, 1985P. kauneorum 2735–3380 Lehr et al., 2002a; Lehr et al., 2002bP. montium 2950–3660 Duellman, 2000P. paucari 3600 Lehr et al., 2005aP. pesantesi 4280–4390 Lehr et al., 2005aP. tautzorum 3770 Lehr and Aguilar, 2003

APPENDIX II

Elevational range in Eleutherodactylus, Phyllonastes and Phrynopus from Andean central Peru.


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