anura: leptodactylidae - BioOne

Herpetological Monographs, 20, 2006, 105–128 E 2006 by The Herpetologists’ League, Inc.

NEW SPECIES OF ELEUTHERODACTYLUS (ANURA: LEPTODACTYLIDAE) FROM THE EASTERN ANDES OF CENTRAL PERU WITH COMMENTS ON CENTRAL PERUVIAN ELEUTHERODACTYLUS EDGAR LEHR1,4,5, MIKAEL LUNDBERG1, CESAR AGUILAR2,

AND

RUDOLF

VON

MAY3

1

Staatliche Naturhistorische Sammlungen Dresden, Museum fu¨r Tierkunde, Ko¨nigsbru¨cker Landstrasse 159, D-01109 Dresden, Germany 2 Museo de Historia Natural, Departamento de Herpetologıá, Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Jesu´s Marıá, Ap. 14-0434, Lima, Peru 3 Department 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 of 1330–3000 m in the Cordillera Oriental in the central Peruvian Departamentos Huańuco, Junıń and Pasco, Three of the new species are assigned to the Eleutherodactylus unistrigatus Group. Two of them lack a tympanum, but have distinct colorations and iris patterns: one has the groin and hind limbs partly orange to red and the iris with a brown vertical and black horizontal streak forming a cross, whereas the other has the groin and hind limbs partly yellow and the iris with a brown horizontal streak and a black vertical streak from the pupil to the lower margin of the eye forming a ‘‘T’’. The third species, assigned to the Eleutherodactylus conspicillatus Group, is similar to E. rhabdolaemus, but differs from that in certain morphological characters and coloration. The fourth species, assigned to the E. lacrimosus assemblage, has a yellowish-tan dorsum with dark brownish-purple blotches and streaks and white flanks and venter. It inhabits terrestrial bromeliads and is compared with E. schultei from northern Peru. Morphological and distributional data are provided for seven previously described Eleutherodactylus from central Peru. The elevational distributions of Andean Eleutherodactylus, Phyllonastes, and Phrynopus from central Peru is compared. Key words: Andes; Anura; Eleutherodactylus; Leptodactylidae; New species; Peru.

KNOWLEDGE OF ELEUTHERODACTYLINE FROGS in central Peru (Departamentos Huańuco, Pasco, Junıń, and southern part of San Martıń) is limited. The species composition of Phrynopus seems to be relatively well known (Hedges, 1990; Duellman, 2000; Lehr, 2002; Lehr et al., 2005a), but only one species of Phyllonastes (Lehr et al., 2004a) and 10 species of Eleutherodactylus have been described for central Peru within the last 10 years (Dwyer, 1995; Lehr et al., 2004b; Lehr 2005; Duellman and Hedges, 2005). Intensive fieldwork since 1998 on the lower and upper forested slopes of the eastern Andes in central Peru has resulted in the collection of several, mostly unidentified, species of Eleutherodactylus, two of which were recently described by Lehr et al. (2004b) and Lehr (2005). The taxonomy of Eleutherodactylus is 4 PRESENT ADDRESS: Natural History Museum and Biodiversity Research Center, Division of Herpetology, The University of Kansas, Jayhawk Boulevard 1345, Lawrence, KS 66045-7561, USA 5 CORRESPONDENCE: e-mail, [email protected]

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complex because of the high number of species (more than 700; Frost, 2004), high interspecific similarity, high intraspecific polymorphism, and the difficulty of identifying diagnostic characters (coloration, skin texture) in poorly preserved specimens. Access to scientific collections that harbor many type specimens as well as identified species of Eleutherodactylus is extremely helpful as a source of comparative material for species descriptions. This was made possible when the senior author visited the Division of Herpetology at The University of Kansas Natural History Museum and Biodiversity Research Center. The examination of recently collected specimens and comparison with museum material, led to the recognition of four undescribed species. Herein, we describe the new species, add information about morphology and distribution for seven other central Peruvian Eleutherodactylus, and compare their elevational distributions with those of Phyllonastes and Phrynopus in the Andes of central Peru.

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HERPETOLOGICAL MONOGRAPHS

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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) and ventral (D) views, photos by M. Lundberg.

MATERIALS AND METHODS The format for the diagnoses follows that of Lynch and Duellman (1997), with inclusion of a description of a holotype. Field notes on the new species were recorded by E. Lehr in August 2003, and by M. Lundberg in March 2000, March 2002, and January 2004. Additionally, photographs were used for descriptions of coloration in life. Definitions of the Peruvian ecoregions follow Brack (1986). Specimens collected between 2000 and 2002 were preserved in 4% formalin; those collected in 2003 and 2004 were preserved in 96% ethanol; all were stored in 70% ethanol. If secondary sex characters were present, specimens were sexed externally; otherwise, they were dissected to determine the presence or absence of ovaries. We follow the definition of a tympanum by Lynch and Duellman (1997). The otic region was dissected in order to evaluate the condition of the tympanic annulus. Measurements taken with digital calipers and rounded to the nearest 0.1 mm are: SVL (snout–vent length), TL (tibia length), FL (foot length, distance from proximal margin of inner metatarsal tubercle to tip of Toe IV), HL (head length, from

angle of jaw to tip of snout), HW (head width, at level of angle of jaw), ED (eye diameter), IOD (interorbital distance), EW (upper eyelid width), IND (internarial distance), E–N (eye–nostril distance, straight line distance between anterior corner of orbit and posterior margin of external nares), TY (tympanum diameter, horizontal). Lengths of Toes III and V were determined when both were adpressed against Toe IV; lengths of Fingers I and II were determined when adpressed against each other. All drawings were made by the senior author using a stereomicroscope with drawing tube attachment (Nikon SMZ 1000). Specimens were deposited in the herpetological collections of the Museo de Historia Natural Universidad Nacional Mayor de San Marcos (MHNSM) in Lima, Peru, and the Museum fu¨r Tierkunde Dresden (MTD), in Dresden, Germany. Codes for museum collections follow those of Leviton et al. (1985) and Frost (2004). For specimens examined, see Appendix I. Eleutherodactylus cruciocularis sp. nov. Holotype.—MHNSM 18685 (Figs. 1A, B), an adult female obtained from Pampa Her-

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mosa (10u 599 33.30 S, 75u 259 58.00 W) at 1540 m between 28 and 29 August 2003, Distrito de Huasahuasi, Provincia de Tarma, Departamento de Junıń, Peru, by E. Lehr. Paratopotypes.—All collected with the holotype 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 (MHNSM 19988, MTD 46832–33), and five juveniles (MHNSM 19985–87, MTD 46834– 35), all obtained from San Alberto (10u 349 38.50 S, 75u 239 32.20 W) at 1850 m on 25 June 2005, Distrito de Oxapampa, Provincia de Oxapampa, Departamento de Pasco, Peru, by M. Lundberg; one female (AMNH 91548) obtained from about 30 km (airline) NE Tingo Marıá, Cordillera Azul, [5 5 km by road SW high point (1640 m) on Tingo MarıáPucallpa Road] at 1330 m on 3 November 1974, Departamento de Huańuco, Peru, by C. W. Myers and J. Daly. Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having the following combination of characters: (1) Skin on dorsum shagreen with small scattered spicules, that on venter areolate; discoidal fold not evident; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus absent; (3) snout short, rounded in dorsal and lateral views; (4) upper eyelid with small tubercles; width of upper eyelid narrower than IOD; cranial crests absent; (5) vomerine teeth small; (6) males lacking vocal sac, vocal slits, and nuptial pads; (7) Finger I shorter than Finger II; discs on outer fingers broadly expanded, rounded; (8) fingers without lateral fringes; (9) ulnar and tarsal tubercles present; (10) heel with single tubercle; inner tarsal fold absent; (11) inner metatarsal tubercle moderate, ovoid, 33 as large as outer; outer metatarsal tubercle small, rounded; few low, diffuse supernumerary plantar tubercles present; (12) toes without lateral fringes; toe webbing absent; Toe V much longer than Toe III; toe discs slightly smaller than those on fingers; (13) dorsum and venter dark grayish brown; groin and anterior surfaces of thighs orange to red; iris gold, with fine, black reticulations and a dark brown,

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broad horizontal streak across pupil and a narrow black vertical streak across pupil forming a cross; (14) SVL in adult females 18.7–21.8 mm (n 5 7), in adult males 11.4– 15.4 mm (n 5 7). Eleutherodactylus cruciocularis is readily distinguished from all other species of the genus by the combination of its lack of a tympanic annulus and membrane, possession of an iris with a cruciform mark and the partly orange to red groin and hind limbs. Five other species of Eleutherodactylus (E. colodactylus, E. flavobracatus, E. imitatrix, E. lirellus, and E. rhabdocnemus) from the eastern Andes and Amazonian lowlands of central Peru lack a tympanum, but none has an iris with a cruciform mark. Eleutherodactylus cruciocularis differs from the similarsized E. imitatrix (females to 20 mm [Duellman and Hedges, 2005] vs. up 21.8 mm in E. cruciocularis) by having vomerine teeth (absent in E. imitatrix). Eleutherodactylus cruciocularis 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 pale brown spots (pale belly with brown flecks and small brown spots). Eleutherodactylus cruciocularis is most similar to E. flavobracatus, but differs from it in the following features (characters of E. flavobracatus in parentheses): maximum SVL in females 21.8 mm (23.4) and in males 15.4 mm (19.6), groin and hind limbs with orange or red blotches (yellow), posterior surface of thighs brown with small, dark orange spots (brown with large yellow longitudinal stripe extending to shanks), upper eyelid tubercles low (prominent), supratympanic fold absent (weak), and eye with dark brown horizontal and narrow, black vertical streak forming a cross (dark brown horizontal streak present, black vertical streak from pupil to venterolateral margin of eye). Eleutherodactylus cruciocularis shares with E. altamazonicus a nearly uniform dark venter, and orange to red blotches in groin and anterior surfaces of thighs, but differs as follows (characters of E. altamazonicus in parentheses): maximum SVL in females 21.8 mm (33.9 mm; Lynch 1980), tympanum absent (present, although partly concealed in some

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

individuals), and inner tarsal fold absent (present). Description of the holotype.—Head as wide as body and nearly as wide as long; head width 35% of SVL; head length 33% of SVL; snout short, rounded in dorsal and in lateral view (Figs. 2A, B); eye diameter larger than eye– nostril distance (eye–nostril distance 83% of length of eye); nostrils slightly protuberant, directed dorsolaterally; canthus rostralis slightly curved in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelid bearing small, low tubercles; upper eyelid width 74% of IOD; supratympanic fold, tympanic annulus and tympanic membrane absent; enlarged postrictal tubercles absent. Choanae small, ovoid, not concealed by palatal shelf of maxilla; vomerine teeth minute, concealed by palatal mucosa, situated posteromedial to choanae, each vomer bearing four teeth in an oblique row, narrowly separated; tongue 1.83 as long as wide (length 6.8 mm, width at midlength of tongue 3.7 mm), not notched posteriorly, posterior one-half free. Skin on dorsum smooth with few, scattered spicules; dorsolateral folds absent; skin on

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flanks smooth with few, scattered spicules; skin on belly and chest areolate, skin on other ventral surfaces smooth; discoidal fold not evident; cloacal sheath short; large tubercles absent in cloacal region. Ulnar and tarsal tubercles present, both low; palmar tubercles slightly elevated, outer palmar tubercle bifid, approximately 2.53 the size of ovoid, inner palmar tubercle; supernumerary tubercles below Fingers III–IV, round, low, slightly smaller than subarticular tubercles; subarticular tubercles well defined, round in dorsal view and subconical in lateral view, most prominent on base of Finger IV; fingers without lateral fringes; Finger I shorter than Finger II; discs on fingers expanded, most prominent on Fingers III–IV, rounded terminally; discs on Finger I slightly expanded; all fingers having ventral pads well defined by circumferential grooves (Fig. 3A). Hind limbs slender, tibia length 50% of SVL; foot length 39% of SVL; upper surfaces of hind limbs smooth; posterior and ventral surfaces of thighs weakly areolate; heel bearing single, round tubercle; tarsus with row of small, low tubercles; tarsal fold absent; inner metatarsal tubercle elevated, elliptical, about 33 subconical outer metatarsal tubercle; few, low plantar supernumerary tubercles; subarticular tubercles well defined, round in dorsal view and subconical in lateral view; toes without lateral fringes; toe webbing absent; discs on toes about equal in size to those on fingers, most prominent on Toes IV–V, toe discs well defined by circumferential grooves; relative lengths of toes: 1 , 2 , 3 , 5 , 4 (Fig. 3B); Toe V much longer than Toe III (disc on Toe III not reaching distal subarticular tubercle on Toe IV, tip of the disc on Toe V extending to distal border of distal subarticular tubercle on Toe IV). Measurements (in mm) of holotype: SVL 21.8; TL 11.0; FL 8.5; HL 7.2; HW 7.6; ED 2.9; IOD 2.7; EW 2.0; IND 1.8; E–N 2.4. Coloration of holotype in preservative.— Dorsum grayish brown with dark brown interorbital bar and blotches in scapula and sacral regions; limbs with dark brown transverse bars; three dark brown labial bars; narrow, dark brown stripe extending from posterior margin of eye diagonally to forearm insertion; venterolateral posterior half of

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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).

flanks dark brown with pale cream blotches; groin and anterior surface of thighs brown with pale cream blotches, posterior surface of thighs brown with small pale cream spots; throat pale gray, chest and belly grayish brown with pale gray spots; posterior part of belly and anterior half of thighs with a pale cream blotch, separated medially by a narrow, grayish-brown stripe; outer surface of left shank with three pale cream blotches, right shank with five pale cream blotches; tips of fingers and toes pale gray; iris pale gray with fine, black reticulations and black horizontal and vertical streaks that form a cross in the iris across the pupil.

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Coloration of holotype in life.—Dorsum pale brown with small red spots on back, dark brown interorbital bar, dark brown blotches in scapula and sacral regions; limbs with dark brown transverse bars; three dark brown labial bars; narrow, dark brown supratympanic stripe extending from posterior margin of eye diagonally to insertion of forearm; posteroventral half of flanks black with pale cream blotches; groin with dark orange blotches; anterior surfaces of thighs brown with dark orange blotches, posterior surfaces of thighs brown with small dark orange spots; ventral surfaces dark gray with pale cream spots; posterior part of belly and anterior half of thighs with a dark orange blotch, separated medially by a narrow, dark gray stripe; outer surface of left shank with three dark orange blotches, right shank with five orange blotches; tips of fingers pale yellow, of toes, pale red; iris gold with fine, black reticulations and dark brown horizontal and black vertical streaks that form a cross in the iris across the pupil. Variation.—The paratypes differed in dorsal and lateral coloration in life. MHNSM 18684, 18688, MTD 45638, and 45642 had tan flanks and a dark brown dorsum. In MHNSM 18682, 18691, MTD 45635, and 45644, the dorsum was pale brown with a dark brown middorsal stripe and two dark brown lateral stripes extending from about the naris to the cloaca; the flanks were dark brown. In MTD 45643, there was a single, narrow, pale brown middorsal stripe and a pale brown stripe on the throat. Also, the specimens differ in the numbers and sizes of the orange blotches. Coloration in life of a female (AMNH 91548) was noted by C. W. Myers as follows: ‘‘Pattern of contrasting, brown markings above, with vivid bright orange flash mark in groin. Rear thigh black with gray flecking and few dots of pale orange; some small pale orange spots on shank.’’ See Tables 1, 2 for measurements of the type series; see Table 3 for ranges and proportions. Etymology.—The specific name is derived from the Latin noun crux, meaning cross, and the Latin noun oculatus, meaning ‘‘having eyes.’’ The name refers to the cruciform iris pattern of this species. Distribution and ecology.—Eleutherodactylus cruciocularis is known from three localities

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

SVL TL FL HL HW ED IOD EW IND E–N

MHNSM 18685

MHNSM 18684

MTD 45637

MTD 45638

MHNSM 18683

MHNSM 18682

MTD 45634

21.8 11.0 8.5 7.2 7.6 2.9 2.7 2.0 1.8 2.4

21.6 10.9 8.6 7.6 8.0 2.6 2.9 1.7 1.8 2.2

20.8 10.5 7.6 7.9 7.9 2.8 2.9 1.4 1.7 1.9

20.7 11.1 7.7 7.2 7.6 2.7 2.7 1.5 2.0 1.9

19.6 10.6 7.5 7.4 7.5 2.5 2.7 1.5 1.9 2.0

18.9 9.7 7.2 6.9 7.1 2.6 2.7 1.9 1.6 2.1

18.7 9.6 7.5 7.6 7.4 2.6 2.8 1.4 1.6 2.2

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

SVL TL FL HL HW ED IOD EW IND E–N

MHNSM 18697

MTD 45635

MTD 45644

MTD 45640

MTD 45641

MTD 45639

MHNSM 18690

15.4 8.0 6.5 6.6 5.8 2.0 2.0 1.2 1.4 2.0

14.7 8.0 6.2 5.6 5.4 2.4 2.0 1.5 1.2 1.8

14.1 7.7 5.7 5.1 5.2 2.2 2.2 1.2 1.1 2.2

13.7 7.2 5.5 5.5 4.8 1.7 1.7 1.2 1.4 1.5

12.9 6.8 5.3 4.6 4.7 1.9 1.9 1.2 1.2 1.4

12.4 6.7 5.1 4.6 4.7 1.5 1.9 1.3 1.4 1.4

11.4 6.2 4.6 4.7 4.7 1.7 1.5 1.2 1.3 1.5

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

E. flavobracatus

Character

Females (n 5 7)

Males (n 5 7)

Females (n 5 2)

Males (n 5 2)

SVL TL FL HL HW ED IOD EW IND E–N TL/SVL FL/SVL HL/SVL HW/SVL HW/SVL HW/HL E–N/ED EW/IOD

18.7–21.8 (20.3 6 1.2) 9.6–11.1 (10.5 6 0.6) 7.5–8.6 (7.8 6 0.5) 6.9–7.9 (7.4 6 0.3) 7.1–8.0 (7.6 6 0.3) 2.5–2.9 (2.7 6 0.1) 2.7–2.9 (2.8 6 0.1) 1.4–2.0 (1.6 6 0.2) 1.6–2.0 (1.8 6 0.1) 1.9–2.4 (2.1 6 0.2) 0.52–0.54 0.40–0.42 0.36–0.43 0.35–0.41 0.35–0.41 0.87–1.02 0.74–1.00 0.55–0.80

11.4–15.4 (13.5 6 1.2) 6.2–8.0 (7.2 6 0.6) 4.6–6.5 (5.6 6 0.6) 4.6–6.6 (5.2 6 0.7) 4.7–5.8 (5.0 6 0.4) 1.5–2.4 (1.9 6 0.3) 1.5–2.2 (1.9 6 0.2) 1.2–1.5 (1.3 6 0.1) 1.1–1.4 (1.3 6 0.1) 1.4–2.2 (1.7 6 0.3) 0.50–0.54 0.37–0.40 0.33–0.41 0.35–0.40 0.35–0.40 0.97–1.06 0.68–0.85 0.48–0.74

21.5–23.4 (22.5 6 1.0) 11.3–11.4 (11.4 6 0.0) 8.8–9.1 (9.0 6 0.2) 8.1–8.5 (8.3 6 0.2) 7.7–8.4 (8.1 6 0.4) 2.5–2.6 (2.6 6 0.1) 2.6–2.7 (2.7 6 0.1) 1.7–1.8 (1.8 6 0.0) 1.6–1.9 (1.8 6 0.2) 2.4–2.5 (2.5 6 0.0) 0.48–0.53 0.39–0.41 0.36–0.38 0.36 0.36 0.99–1.08 0.96 0.65–0.67

18.2–19.6 (18.9 6 0.7) 10.3–10.4 (10.4 6 0.0) 8.2–8.4 (8.3 6 0.1) 8.0–8.1 (8.1 6 0.0) 7.0–7.1 (7.1 6 0.0) 2.2–2.3 (2.2 6 0.0) 2.8–2.9 (2.9 6 0.1) 1.6–1.8 (1.7 6 0.1) 1.4–1.6 (1.5 6 0.1) 1.9–2.2 (2.1 6 0.2) 0.53–0.56 0.42–0.46 0.41–0.45 0.36 0.36–0.38 0.86–0.89 0.68–0.76 0.57–0.62

(Pampa Hermosa, San Alberto, and 30 km [airline] NE Tingo Marıá) at elevations of 1330–1850 m in the Selva Alta Ecoregion of central Andean Peru. The type locality (Fig. 4) is near the Chanchamayo Valley and

supports undisturbed primary forest. All specimens were found at night on vegetation approximately 1 m above the ground or on the ground. Sympatric anurans include Eleutherodactylus bipunctatus, E. bromeliaceus, E. cf.

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

platydactylus, E. sagittulus, Hybsiboas cf. melanopleura, and Gastrotheca atympana. See Figure 5 for comparative altitudinal distributions. Eleutherodactylus flavobracatus sp. nov. Holotype.—MHNSM 19871 (Figs. 1C, D), an adult female obtained at Km 34 on road

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from Oxapampa to Yaupi (10u 449 44.40 S, 75u 309 02.29 W) at 1770 m on 18 October 2003, Distrito de Chontabamba, Provincia de Oxapampa, Departamento de Pasco, Peru, by M. Lundberg. Paratopotypes.—Four, all collected by M. Lundberg: 1 female (MTD 45908, on 18 October 2003), 2 males (MHNSM 19848, MTD 45716, on 25 May 2004), 1 juvenile (MTD 45717, on 1 June 2003). Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having: (1) skin on dorsum shagreen with small scattered tubercles, that on venter areolate; discoidal fold not evident; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus absent; (3) snout short, rounded in dorsal and lateral views; (4) upper eyelid with two tubercles, one at midlength of eyelid near inner margin, the other at its posterior end; upper eyelid narrower than IOD; cranial crests absent; (5) vomerine teeth small; (6) males lacking vocal sac, vocal slits, and nuptial pads; (7) Finger I shorter than Finger II; discs on outer fingers broadly expanded, rounded; (8) fingers without lateral fringes; (9) ulnar and tarsal tubercles present; (10) heel with single tubercle; inner tarsal fold

FIG. 5.—Range of altitudinal distribution in Andean Eleutherodactylus, Phyllonastes, and Phrynopus (excluding species 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 of distribution and precise distributional data, see Appendix II.

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absent; (11) inner metatarsal tubercle moderate, ovoid, 23 as large as outer; outer metatarsal tubercle small, rounded; few supernumerary plantar tubercles present, low, diffuse; (12) toes without lateral fringes; toe webbing absent; Toe V much longer than Toe III; discs on toes slightly smaller than those on fingers; (13) dorsum and venter dark brown; groin and anterior and posterior surfaces of thighs yellow; shank yellow ventrally; iris gold with fine, black reticulations and a broad, dark brown horizontal streak across pupil and a narrow, black vertical streak from pupil to lower margin of eye forming a ‘‘T;’’ (14) SVL in adult females 21.5–23.4 mm (n 5 2), in adult males 18.2–19.6 mm (n 5 2). Eleutherodactylus flavobracatus is readily distinguished from all species of the genus by the combination of lacking a tympanum and possessing an iris with a dark brown horizontal streak and a dark brown vertical streak from pupil to lower margin of eye forming a ‘‘T,’’ and having the groin and hind limbs partly yellow. Five other species of Eleutherodactylus (E. colodactylus, E. cruciocularis, E. imitatrix, E. rhabdocnemus, and E. sp. nov.) from the eastern Andes and Amazonian lowlands of Peru lack a tympanum, but in none does the eye have a T–shaped mark. Furthermore, E. flavobracatus is slightly larger than E. imitatrix (SVL to 23.4 mm in female E. flavobracatus vs. 20 mm in female E. imitatrix [Duellman and Hedges, 2005]) and has vomerine teeth (absent in E. imitatrix). Eleutherodactylus flavobracatus differs from E. rhabdocnemus (which has a similar size, 24.5 mm in females [Duellman and Hedges, 2005]) by having prominent tubercles on the upper eyelid (absent in E. rhabdocnemus), and by having a brown belly with dark brown spots (belly pale with brown flecks and small brown spots). Eleutherodactylus flavobracatus is most similar to E. cruciocularis; for comparison of the two species see diagnosis of E. cruciocularis. Description of the holotype.—Head as wide as body, slightly longer than wide; head width 36% of SVL; head length 38% of SVL; snout short, rounded in dorsal and lateral views (Figs. 2C, D); eye diameter barely larger than eye–nostril distance (eye–nostril distance 96% of length of eye); nostrils slightly protuberant,

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directed dorsolaterally; canthus rostralis curved in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelid bearing two prominent tubercles, one located at midlength of eyelid near outer margin, the other at its posterior end; upper eyelid width 65% of IOD; supratympanic fold weak, tympanic annulus and tympanic membrane absent; enlarged postrictal tubercles absent. Choanae small, ovoid, not concealed by palatal shelf of maxillary arch; vomerine teeth small, concealed by surrounding tissues, situated posteromedially to choanae, each vomer bearing four teeth in an oblique row and narrowly separated; tongue 1.23 as long as wide (length 5.2 mm, width at midlength of tongue 4.5 mm), not notched posteriorly, posterior one-third free. Skin on dorsum smooth with few, scattered spicules; dorsolateral folds absent; skin on flanks smooth with few, scattered spicules; skin on belly and chest weakly areolate, skin on other ventral surfaces smooth; discoidal fold not evident; cloacal sheath short; large tubercles in cloacal region absent. Ulnar and tarsal tubercles present, both low; palmar tubercles slightly elevated, outer palmar tubercle bifid, approximately 23 size of ovoid, inner palmar tubercle; supernumerary tubercles below Fingers II–IV, round, low, slightly smaller than subarticular tubercles; subarticular tubercles well defined, round in dorsal view and subconical in lateral view, most prominent on base of Finger I; fingers without lateral fringes; Finger I shorter than Finger II; discs on fingers expanded, most prominent on Fingers III–IV, rounded terminally; disc on Finger I slightly expanded; all fingers having ventral pads well defined 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 hind limbs smooth; posterior and ventral surfaces of thigh areolate; heel bearing single, slightly elevated, round tubercle; tarsus with row of small, low tubercles; tarsal fold absent; inner metatarsal tubercle elevated, elliptical, about 23 subconical outer metatarsal tubercle; few, low plantar supernumerary tubercles present; subarticular tubercles well defined, round in dorsal view and subconical in lateral view; toes without lateral fringes; toe webbing absent;

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

Sex SVL TL FL HL HW ED IOD EW IND E–N

MTD 45908

MHNSM 19871

MHNSM 19848

MTD 45716

MTD 45717

Female 23.4 11.3 9.1 8.5 8.4 2.5 2.7 1.8 1.9 2.4

Female 21.5 11.4 8.8 8.1 7.7 2.6 2.6 1.7 1.6 2.5

Male 19.6 10.4 8.2 8.0 7.1 2.2 2.8 1.6 1.4 1.9

Male 18.2 10.3 8.4 8.1 7.0 2.3 2.9 1.8 1.6 2.2

Juvenile 14.7 8.8 6.8 5.6 5.6 1.9 2.0 1.3 1.6 1.7

discs on toes slightly smaller than those on fingers, most prominent on Toes IV–V, well defined by circumferential grooves; relative lengths of toes: 1 , 2 , 3 , 5 , 4 (Fig. 3D); Toe V much longer than Toe III (disc on Toe III not reaching distal subarticular tubercle on Toe IV, tip of the disc on Toe V extending to distal border of distal subarticular tubercle on Toe IV). Measurements (in mm) of holotype: SVL 21.5; TL 11.4; FL 8.8; HL 8.1; HW 7.7; ED 2.6; IOD 2.6; EW 1.7; IND 1.6; E–N 2.5. Coloration of holotype in preservative.— Dorsum dark brown with tan blotch anterior to nares; hind limbs with transverse dark brown bars; ill–defined, dark brown labial bars present; narrow, dark brown supratympanic stripe extending from posterior margin of eye diagonally to insertion of forearm; flanks tan dorsolaterally, dark brown ventrolaterally; groin with a white blotch; anterior surfaces of thighs white, bordered by dark brown bars, posterior surfaces of thighs with a transverse white streak; ventral surfaces brown with dark brown spots, except for posterior part of belly, proximal half of thighs, and ventral surfaces of shanks which are white; tarsus white dorsally; finger tips and toe tips pale gray. Iris black. Coloration of holotype in life.—As described above with the exception that the white areas were yellow, and the iris gold with fine, black reticulations, a dark brown horizontal streak across pupil, and a black vertical streak from pupil to lower margin of eye forming a ‘‘T.’’ Variation.—Variation exists in the dorsal and lateral coloration, and in the amount of

yellow coloration. In MTD 45908, the dorsum and flanks were dark brown, and there was a yellow blotch in the groin. The holotype had the most yellow coloration in the groin and on the hind limbs. See Table 4 for measurements of the type series; see Table 3 for ranges and proportions. Etymology.—The specific name is derived from the Latin noun flavus, meaning yellow, and the Latin adjective bracatus, meaning ‘‘wearing trousers.’’ The name refers to the yellow coloration of groin and parts of the hind limbs of this species. Distribution and ecology.—Eleutherodactylus flavobracatus is known only from the type locality (Fig. 4), which is in the Selva Alta Ecoregion. All specimens were found at night at 1900–2200 hr on vegetation 1–3 m above ground, next to a road. The roadside is nearly vertical and covered with secondary vegetation, which is cut several times a year as part of road maintenance. There are several small creeks near the type locality, and a roadside ditch contained running water. At night the leaves were covered with water drops even when it was not raining. No calls were heard. Sympatric anurans include Atelopus sp., Eleutherodactylus bipunctatus, E. rhabdocnemus, and Hyalinobatrachium bergeri. See Figure 5 for comparative altitudinal distributions. Eleutherodactylus ornatus sp. nov. Holotype.—MHNSM 20664 (Figs. 6A, B), an adult female obtained at Cillapata (ca. 1.5 km NNE Auquimarca, approximately 10u 439 520 S; 75u 429 480 W) at 2900 m on 11 March 2002, Distrito de Paucartambo, Pro-

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FIG. 6.—Eleutherodactylus ornatus (MTD 45897, female, SVL 21.1 mm), in lateral (A) and ventral (B) views, photos by E. Lehr; Eleutherodactylus rhabdolaemus (KU 173237, male, SVL 22.7 mm) in lateral view (C), photo by W. E. Duellman.

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

[No. 20

Paucartambo: Cillapata (approximately 1.5 km north-northeast of Auquimarca), approximately 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 (March 2002), MTD 45073 (1 June 2002); Paucartambo: near road from Uchuhuerta to Auquimarca (10u 419 05.60 S; 75u 459 28.70 W, 2900 m): MTD 45897–98 (10 December 2003); Huachoń: Uchuhuerta, 2400 m: MHNSM 19883 (22 October 2003); Huachoń: km 77 at the road Ninacaca-Huachoń-Puagmaray-Oxapampa, 2550 m: MTD 46319 (27 January 2004). Diagnosis.—A member of the Eleutherodactylus (Eleutherodactylus) conspicillatus Group having: (1) skin on dorsum finely shagreen with small scattered tubercles, that on venter smooth; discoidal fold present; dorsolateral folds present; (2) tympanic membrane and tympanic annulus present, upper and posterolateral margins concealed by supratympanic fold; (3) snout long, subacuminate in dorsal view, round in lateral view; (4) upper eyelid without tubercles; width of upper eyelid much narrower than IOD; cranial crests absent; (5) vomerine teeth small, oblique; (6) males with vocal slits, vocal sac not visible externally, nuptial pads absent; (7) Finger I shorter than Finger II; discs on outer fingers broadly expanded, rounded; (8) fingers without lateral fringes; (9) ulnar and tarsal tubercles absent; (10) heel without tubercles; inner tarsal fold absent; (11) inner metatarsal tubercle moderate, ovoid, 23 as large as outer; outer metatarsal tubercle small, rounded; few supernumerary plantar tubercles present, low; (12) toes without lateral fringes and webbing; Toe V slightly longer than Toe III; toe discs smaller than those on fingers; (13) dorsum tan with dark brown blotches forming two irregular chevrons; extremities with irregular dark brown blotches; dark brown canthal and supratympanic stripes; venter whitish gray with brown blotches on chest and throat; groin and anterior and posterior surfaces of thighs reddish brown; lower one-half of iris dark brown, upper onehalf pale gold with fine, black reticulations; (14) SVL in adult females 20.7–27.3 mm (n 5 11), in adult males 16.7–20.5 mm (n 5 7).

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

Eleutherodactylus ornatus is most similar to E. rhabdolaemus (Fig. 6C), a member of the E. conspicillatus Series according to Lynch and Duellman (1997), which is known from the Peruvian Departamentos Huańuco, Ayacucho, and Cusco, southwards to Departamento Santa Cruz in Bolivia at elevations of 1000–2700 m (Frost, 2004). Both E. ornatus and E. rhabdolaemus have dorsolateral folds and dark brown canthal and supratympanic stripes (broader in E. ornatus), but these species can be distinguished by the following features (characters of E. rhabdolaemus in parentheses): maximum SVL in females 27.3 mm (31.7; Duellman, 1978a) and in males 20.5 mm (24.0; Duellman, 1978a), dorsolateral fold weakly defined (clearly distinct), diameter of eye approximately 33 the diameter of tympanum (approximately 23), diameter of tympanum equal to eye–tympanum distance, Figs. 7A, B (one-half of tympanum diameter, Figs. 7C, D), males with vocal sac not visible externally (subgular vocal sac visible), vomerine teeth in a clump (oblique row), scapular tubercles absent (present), venter smooth (coarsely areolate), supratympanic fold prominent (weak), terminal discs on fingers not broadly expanded,

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

1.23 width of digit, Fig. 8A (terminal discs on fingers broadly expanded, 23 width of digit, Fig. 8C), small tubercles on heel absent (present), Toe III slightly shorter than Toe V, Fig. 8B (much shorter, Fig. 8D); Toe III not reaching distal subarticular tubercle on Toe IV (reaching), Toe V reaching lower margin of distal subarticular tubercle on Toe IV (overlapping distal subarticular tubercle), irregularly shaped, dark brown blotches on body forming ‘‘interrupted’’ chevrons (clearly defined dark chevrons), extremities with dark brown blotches, rarely presenting diagonal bars (clearly defined diagonal bars present). Superficially, Eleutherodactylus ornatus can be confused with E. toftae, a member of

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the E. unistrigatus Group, which is restricted to forests of the Amazonian lowlands to lower Andean foothills of southern Peru to Cochabamba, Bolivia (Frost, 2004). These species are similar in size (maximum SVL 27.4 mm in females of E. toftae; Duellman, 2005) and both have a long snout, dark brown canthal and supratympanic stripes, and dorsolateral folds (absent in E. toftae according to the original species description and a recent species account by Duellman [1978b, 2005]). However, E. ornatus has fingers and toes without lateral fringes (present in E. toftae), tan flanks with brown blotches (dark brown in E. toftae), and lacks a labial stripe (present in E. toftae). Eleutherodactylus ornatus and the central Peruvian E. aniptopalmatus have similar elevational distributions, smooth ventral skin, and dark brown canthal and supratympanic stripes, but E. ornatus is larger (maximum SVL in females 27.3 mm vs. 22.0 mm in E. aniptopalmatus; Duellman and Hedges, 2005), has a long snout (moderately long in E. aniptopalmatus) and dorsolateral folds (absent in E. aniptopalmatus). Juvenile E. ornatus can be confused with juvenile E. bipunctatus, a member of the E. conspicillatus Group. Both species have long snouts, dark brown canthal and supratympanic stripes, dorsolateral folds, and lack lateral fringes, 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 is restricted to lower elevations (230–2320 m), whereas E. ornatus occurs at higher elevations (2400–3000 m). Description of the holotype.—Head as wide as body, as wide as long; head width 36.4% of SVL; head length 36.8% of SVL; snout long, subacuminate in dorsal view, round in lateral view (Figs. 7A, B); eye diameter nearly equal to as eye–nostril distance; nostrils slightly protuberant, directed dorsolaterally; canthus rostralis straigth in dorsal view, rounded in profile; loreal region concave; lips rounded; upper eyelid without tubercles; width of upper eyelid much narrower than IOD (upper eyelid width 58.6% of IOD); supratympanic fold prominent; tympanic membrane and tympanic annulus present, tympanum length 33% of eye length, separated from eye by

[No. 20

distance equal to tympanum length; tympanum slightly higher than long, its upper and posterolateral margin concealed by supratympanic fold; two enlarged, conical postrictal tubercles present. Choanae small, ovoid, not concealed by palatal shelf of maxilla; vomerine teeth small, oval, situated posteromedially to choanae, each vomer bearing five teeth, vomers widely separated; tongue 1.33 as long as wide (length 6.1 mm, width at midlength of tongue 4.6 mm), not notched posteriorly, posterior half free. Skin on dorsum finely shagreen with small, scattered tubercles; weakly defined dorsolateral fold on each side of body; skin on flanks finely shagreen with scattered tubercles slightly larger than those on dorsum; skin on proximal posteroventral surfaces of thighs weakly areolate, other ventral surfaces smooth; discoidal fold present; cloacal sheath short; large tubercles absent in cloacal region. Ulnar and tarsal tubercles or folds absent; palmar tubercles slightly elevated, outer palmar tubercle bifid, approximately 2.53 size of ovoid, inner palmar tubercle; few supernumerary tubercles present, round, low, slightly smaller than subarticular tubercles; subarticular tubercles well defined, round in dorsal view and subconical in lateral view, most prominent on bases of fingers; fingers without lateral fringes; Finger I shorter than Finger II; discs on fingers expanded, most prominent on Fingers III–IV, disc on Finger III nearly truncate terminally, other discs on fingers rounded, disc on Finger I slightly expanded; all fingers with ventral pads well defined by circumferential grooves (Fig. 8A). Hind limbs slender, tibia length 58.1% of SVL; foot length 52.6% of SVL; upper surface of hind limbs smooth; posterior and ventral surfaces of thighs weakly areolate; heel without tubercles; tarsus without tubercles and fold; inner metatarsal tubercle elevated, elliptical, about 23 subconical outer metatarsal tubercle; few, low plantar supernumerary tubercles present; subarticular tubercles well defined, round in dorsal view, subconical in lateral view; toes without lateral fringes and webbing; discs on toes slightly smaller than those on fingers, most prominent on Toes IV–V; toe discs well defined by circumferential grooves; relative lengths of

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

SVL TL FL HL HW ED TY IOD EW IND E–N

MHNSM 20665

MTD 44768

MTD 45073

MTD 44767

MHNSM 20667

MTD 45897

MTD 45898

27.3 15.2 13.4 10.3 9.8 2.8 1.0 3.0 1.9 2.9 2.9

26.3 14.3 13.1 9.4 9.5 2.4 1.1 3.2 1.8 2.8 2.8

26.1 14.9 13.8 9.9 9.9 2.7 1.1 3.2 1.6 3.0 2.9

25.3 14.3 12.6 9.2 9.7 2.3 0.9 3.2 1.7 2.7 2.6

23.7 13.9 13.1 8.4 9.5 2.4 0.8 2.9 1.8 2.7 2.6

21.1 12.6 11.4 8.4 8.3 2.2 1.0 3.2 1.2 2.3 2.5

20.9 12.3 10.4 8.5 8.4 2.2 0.8 3.0 1.4 2.7 2.4

toes: 1 , 2 , 3 , 5 , 4 (Fig. 8C); Toe V slightly longer than Toe III (tip of the disc on Toe III reaches middle of disc on Toe V; disc on Toe III not reaching penultimate subarticular tubercle on Toe IV, tip of the disc on Toe V reaching middle of penultimate tubercle on Toe IV). Measurements (in mm) of holotype: SVL 25.3; TL 14.7; FL 13.3; HL 9.3; HW 9.2; ED 2.4; TY 0.8; IOD 2.9; EW 1.7; IND 3.3; E–N 2.3. Coloration of holotype in preservative.— Dorsum grayish tan with dark brown blotches arranged in three ill-defined, interrupted (without contact at midline) chevrons, and narrow, dark brown interorbital bar; dark brown blotches on lower arm and outer surface of hand; anterior surfaces of arms with a dark brown longitudinal stripe; hind limbs with dark brown blotches, and thighs with ill-defined, transverse, dark brown bars; dark brown canthal and supratympanic stripes; flanks slightly paler than dorsum, with dark brown blotches forming three irregular diagonal bars; groin pale cream with small,

pale brown spots; anterior and posterior surfaces of thighs grayish tan with small pale brown spots; ventral surfaces cream with small dark brown spots, most densely on throat and chest; iris black. Coloration of holotype in life.—Unknown. Variation.—Coloration in life for female MTD 45897 is based on photos. Dorsum tan and brown with blackish-brown blotches forming interrupted chevrons; canthal and supratympanic stripes blackish brown; upper half of flanks pale tan with blackish-brown blotches forming three irregular diagonal bars; lower half of flanks white; groin, anterior and posterior surfaces of thigh reddish brown; belly, chest, and throat whitish gray with grayish-brown blotches more dense on throat; upper half of iris pale gold with black reticulations, lower part with a broad, dark brown, horizontal stripe. See Tables 5, 6 for measurements of the type series; see Table 7 for ranges and proportions. Etymology.—The specific name is derived from the Latin noun ornatus, meaning decorated. 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

20.5 12.2 10.9 8.0 8.3 21.0 1.0 2.4 1.4 2.6 2.1

20.3 12.1 9.9 8.3 8.1 2.2 0.7 2.1 1.2 2.4 2.3

20.1 11.1 10.4 7.5 7.9 2.4 0.9 2.5 1.5 2.4 2.1

19.9 11.3 9.9 7.4 7.6 1.9 0.8 2.2 1.5 2.3 2.3

19.8 11.3 10.3 7.4 7.9 1.9 0.8 2.1 1.2 2.6 2.1

18.4 10.9 8.9 6.7 7.0 2.0 0.8 2.1 1.4 2.2 2.0

16.7 10.7 8.6 7.3 6.7 1.8 0.8 2.1 1.3 2.2 2.0

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[No. 20

TABLE 7.—Ranges (in mm) and proportions of adult Eleutherodactylus ornatus; ranges followed by means and one standard deviation in parentheses. Characters

Females (n 5 11)

SVL 20.7–27.3 (24.4 6 2.3) TL 12.3–15.2 (14.1 6 0.9) FL 10.4–13.8 (12.6 6 1.0) HL 8.4–10.3 (9.3 6 0.7) HW 8.3–9.9 (9.3 6 0.6) ED 2.2–2.8 (2.5 6 0.2) TY 0.8–1.2 (1.0 6 0.2) IOD 2.9–3.3 (3.1 6 0.2) EW 1.2–1.9 (1.6 6 0.2) IND 2.2–3.3 (2.8 6 0.2) E–N 2.3–2.9 (2.6 6 0.2) TL/SVL 0.53–0.68 FL/SVL 0.47–0.59 HL/SVL 0.35–0.49 HW/SVL 0.33–0.48 HW/HL 0.88–1.08 E–N/ED 0.89–1.17 EW/IOD 0.38–0.64 TY/ED 0.33–0.48

Males (n 5 7)

16.7–20.5 (19.4 6 1.3) 10.7–12.2 (11.4 6 0.5) 8.6–10.9 (9.8 6 0.8) 6.7–8.3 (7.5 6 0.5) 6.7–8.3 (7.6 6 0.5) 1.8–2.4 (2.0 6 0.2) 0.7–1.0 (0.8 6 0.1) 2.1–2.5 (2.2 6 0.2) 1.2–1.5 (1.4 6 0.1) 2.2–2.6 (2.4 6 0.2) 2.1–2.3 (2.1 6 0.1) 0.57–0.64 0.48–0.53 0.36–0.44 0.63–0.70 0.92–1.07 0.88–1.21 0.57–0.68 0.32–0.50

on the dorsum; these are reminiscent of emblems (chevrons) used on military uniforms. Distribution and ecology.—This species is known from several localities at elevations of 2400–3000 m in the Provincia de Pasco (Fig. 4). Eleutherodactylus ornatus was found at night between 1900–2300 hr on the ground or on vegetation up to 1 m above the ground. During the day, specimens were found among leaf litter and under stones. One pair (female MHNSM 20665, male MTD 44771) was in axillary amplexus on 5 March 2002. Sympatric anurans include Bufo multiverrucosus, E. lundbergi, E. mendax, E. platydactylus, Gastrotheca stictopleura, Phrynopus juninensis, and Phyllonastes duellmani. See Figure 5 for comparative altitudinal distributions. Eleutherodactylus pardalinus sp. nov. Holotype.—MHNSM 19806 (Fig. 9), an adult female obtained at Huasahuasi (11u 169 06.10 S, 75u 399 28.30 W) at 2640 m on 1 September 2003, Distrito de Huasahuasi, Provincia de Tarma, Departamento de Junıń, Peru, by E. Lehr. Paratopotypes.—15, all collected with the holotype by C. Aguilar, E. Lehr, and R. von May: 7 females (MHNSM 19807–08, MTD 45666–69, 45671), 8 males (MHNSM 19809– 13, MTD 45670, 45672–73).

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

Diagnosis.—A member of the Eleutherodactylus lacrimosus Assemblage of the Eleutherodactylus (Eleutherodactylus) unistrigatus Group having: (1) skin on dorsum shagreen with small tubercles, that on venter coarsely areolate; discoidal fold not evident; dorsolateral folds absent; (2) tympanic membrane and tympanic annulus distinct, upper and posterolateral margins of tympanum concealed by supratympanic fold; (3) snout acuminate in dorsal view, rounded in lateral view; (4) upper eyelid with small tubercles and usually one prominent tubercle at center of eyelid; width of upper eyelid narrower than IOD; cranial crests absent; (5) vomerine teeth widely separated, in oblique clump; (6) males with vocal slits and nuptial pads; (7) Finger I shorter than Finger II; discs on outer fingers broadly expanded, rounded; (8) fingers with broad lateral fringes; (9) ulnar and tarsal tubercles present; (10) heel with small, low

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tubercles and one prominent tubercle; inner tarsal fold absent; (11) inner metatarsal tubercle ovoid, 33 outer; conical outer metatarsal tubercle; many supernumerary plantar tubercles present, low, diffuse; (12) toes with broad lateral fringes; toe webbing absent; Toe V much longer than Toe III; toe discs equal in size to those on fingers; (13) dorsum yellowish tan with dark, brownish-purple blotches; flanks white; venter white; groin, anterior and posterior surfaces of thighs yellowish tan; iris brownish red or tan with horizontal, dark brown band and with fine, black reticulations; (14) SVL in females 26.1– 34.1 mm (n 5 8), in males 21.1–26.7 mm (n 5 8). With respect to habitus and coloration, Eleutherodactylus pardalinus resembles other bromeliad-inhabiting species in the Cordillera Oriental, these are E. bromeliaceus, E. lacrimosus, E. mendax, E. olivaceus, and E. schultei. Eleutherodactylus pardalinus is slightly smaller than E. schultei, but larger than the other species: the maximum SVL of female E. pardalinus is 34.1 mm, whereas that of female E. bromeliaceus is 28.1 mm (Lynch, 1979), E. olivaceus 24.2 mm (Ko¨hler, 2000), E. lacrimosus 32.5 mm (Lynch, 1980), and E. mendax 28.1 mm (Duellman, 1990). Furthermore, in E. pardalinus the dorsum is shagreen, whereas the dorsum is smooth in E. bromeliaceus. Eleutherodactylus pardalinus has robust hind limbs, whereas the hind limbs are slender in E. lacrimosus (Lynch and Duellman, 1980). Eleutherodatylus pardalinus has white flanks, whereas E. mendax has yellow flanks (Duellman, 1978c). Eleutherodactylus pardalinus has the dorsum yellowish tan with brownish-purple blotches, whereas the dorsum is uniform olive-green in E. olivaceus. Eleutherodactylus pardalinus is most similar to E. schultei, which is known at elevations of 2400–2850 m (Duellman, 1990) at two localities in northern Peru (Departamento de Amazonas). Both species are about the same size (males: 21.1–26.7 mm in E. pardalinus and 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) and have white bellies. However, they can be distinguished by the following features (char-

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

acters of E. schultei in parentheses): body and extremities robust (slender); fingers and toes with broad lateral fringes (narrow); males with nuptial pads (absent); disc on Toe III reaches above distal subarticular tubercle on Toe IV (not reaching above); 3–5 vomerine teeth (2– 4, or absent [Duellman, 1990]); dorsal coloration yellowish tan (dull green, greenish yellow, creamy tan, pale yellowish tan, or reddish brown [Duellman, 1990]); dorsum with large purple blotches and streaks (dorsum uniform, bordered by black stripes, or with small dark flecks [Duellman, 1990]); flanks white (tan to greenish yellow [Duellman, 1990]). Description of the holotype.—Head narrower than body, slightly wider than long; head width 38.4% of SVL; head length 36.7% of SVL; snout moderately short, acuminate in dorsal 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 dorsolaterally; canthus rostralis straight in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; upper eyelid bearing small tubercles and one prominent tubercle at

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center of eyelid; width of upper eyelid 50% of IOD; supratympanic fold prominent; tympanic annulus and tympanic membrane present, tympanum round, its length 47.1% of eye length, separated from eye by distance about one-half length of tympanum, its upper margin concealed by supratympanic fold; one enlarged, conical postrictal tubercles on each side. Choanae ovoid, not concealed by palatal shelf of maxillary arch; vomerine teeth small, situated posteromedially to choanae, right vomer bearing four teeth (vomerine teeth on left side completely concealed by buccal tissues) in an oblique row, widely separated; tongue 1.13 as long as wide (length 6.3 mm, width at midlength of tongue 5.6 mm), slightly notched posteriorly, posterior third free. Skin on dorsum shagreen with small tubercles; dorsolateral folds absent; skin on upper half of flanks same as dorsum, venterolaterally coarsely areolate; skin on thighs, belly, and chest coarsely areolate, skin on other ventral surfaces smooth; discoidal fold not evident; cloacal sheath short; no large tubercles in cloacal region. Venterolateral surface of forearm bearing row of low tubercles; palmar tubercles slightly elevated, outer palmar tubercle bifid, approximately 43 the size of narrow, ovoid, inner palmar tubercle; numerous low round supernumerary tubercles, slightly to much smaller than subarticular tubercles; subarticular tubercles well defined, round in dorsal view and subconical in lateral view; fingers with broad lateral fringes, outer fringe of Finger IV extends to outer edge of palm with its shape slightly undulated; Finger I shorter than Finger II; discs on fingers broadly expanded, most prominent on Fingers III–IV, rounded terminally; all fingers with ventral pads well defined by circumferential grooves (Fig. 10C). Hind limbs robust, tibia length 48.1% of SVL; foot length 47.5% of SVL; upper surfaces of hind limbs shagreen with small tubercles; posterior and ventral surfaces of thighs coarsely areolate; heel bearing small, low, round tubercles, two on each heel more prominent; inner ventral surface of tarsus with row of low tubercles partially coalesced into fold; inner metatarsal tubercle elevated, elongate, about 2.33 subconical outer metatarsal tubercle; numerous, low plantar supernumer-

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ary tubercles; subarticular tubercles well defined, round in dorsal view and subconical in lateral view; toes with broad lateral fringes; toe webbing absent; discs on toes equal to those on fingers, most prominent on Toes IV–V, toe discs 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 subarticular tubercle on Toe IV, disc on Toe V extending to distal subarticular tubercle on Toe IV). Measurements (in mm) of holotype: SVL 34.1; 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 side of neck; narrow, discontinuous, dark brown interorbital bar; dark brown canthal and supratympanic stripes; flanks cream; few pale brown blotches present on arms and thighs; shanks and outer surface of feet with many blotches; belly grayish cream, other ventral surfaces uniformly cream. Coloration of holotype in life.—Dorsum yellowish tan with dark brownish-purple blotches and streaks as described above; dark brownish-purple blotches on extremities; flanks white; chest and belly white with pale gray interspaces between granules; other ventral surfaces yellow to flesh-colored; iris brownish red, with horizontal dark brown band and fine black reticulations. Variation.—Individuals differ in the number of dorsal blotches and streaks. In males, the dorsum is less intensively blotched than in females; moreover, the anterior half of dorsum in males has many blotches, whereas the posterior half has fewer blotches. Males have white, nonspinous nuptial pads dorsolaterally on the bases of the thumbs. See Tables 8, 9 for measurements of the type series; see Table 10 for ranges and proportions. Etymology.—The specific name is the diminutive of the Greek noun pardos, meaning leopard or panther. The name refers to the dorsal pattern of dark blotches on a yellowishtan ground color, which reminds us of a leopard.

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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

34.1 16.4 16.2 12.5 13.1 3.4 1.6 4.8 2.4 2.3 3.9

33.4 16.2 15.7 12.1 12.2 3.0 1.7 4.4 2.3 2.3 3.2

32.0 17.0 16.4 12.2 12.3 3.0 1.5 4.8 2.2 2.2 3.9

31.5 17.1 15.0 12.5 12.5 2.9 1.4 4.1 2.6 2.1 3.6

31.1 16.2 15.2 11.0 11.0 2.9 1.5 4.4 2.4 2.1 3.4

29.4 17.0 14.5 11.0 11.2 2.7 1.2 4.5 2.2 2.3 3.5

26.1 14.5 12.7 9.9 10.2 2.9 1.2 3.9 2.2 2.1 3.2

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

26.7 13.4 12.3 9.6 10.1 2.9 1.0 3.9 1.9 2.1 3.1

26.3 14.3 12.9 9.5 10.1 2.4 1.2 3.9 2.2 1.8 3.2

24.8 12.4 11.2 8.5 9.1 2.5 1.0 3.2 2.1 2.0 2.8

22.7 12.9 11.0 8.9 8.9 2.5 1.0 3.3 1.7 1.9 2.9

22.4 12.8 10.7 7.8 8.4 2.5 0.9 3.6 1.7 1.7 2.4

22.1 12.0 10.6 8.1 8.1 2.1 0.8 3.2 1.7 1.8 2.6

21.7 13.2 10.1 8.8 8.8 2.2 0.7 3.3 1.8 1.8 2.8

Distribution and ecology.—Eleutherodactylus pardalinus is known only from the type locality, next to the village Huasahuasi (Fig. 4), which belongs to the Puna EcoreTABLE 10.—Ranges (in mm) and proportions of adult Eleutherodactylus pardalinus; ranges followed by means and one standard deviation in parentheses. Characters

SVL TL FL HL HW ED TY IOD EW IND E–N TL/SVL FL/SVL HL/SVL HW/SVL HW/HL E–N/ED EW/IOD TY/ED

gion. All specimens were found during the early afternoon in terrestrial bromeliads surrounded by the Peruvian Feather Grass (Stipa ichu) on a hillside. No other anurans were found there. See Figure 5 for comparative altitudinal distributions.

COMMENTS ON CENTRAL PERUVIAN ELEUTHERODACTYLUS 29.4–34.1 (31.9 6 1.5) 21.7–26.7 (23.6 6 1.9) 16.2–17.1 (16. 6 7 0.4) 12.0–13.4 (13.0 6 0.6) Eleutherodactylus bipunctatus Duellman and Hedges 2005 14.5–16.4 (15.5 6 0.7) 10.1–12.3 (11.3 6 0.9) 11.0–12.5 (11.9 6 0.6) 7.8–9.6 (8.7 6 0.6) Eleutherodactylus bipunctatus in the E. 11.0–13.1 (12.1 6 0.7) 8.1–10.1(9.1 6 0.7) conspicillatus Group was described on the 2.7–3.4 (3.0 6 0.2) 2.1–2.9 (2.4 6 0.2) 1.2–1.7 (1.5 6 0.2) 0.7–1.0 (1.0 6 0.1) basis of one adult female; seven juveniles were 4.1–4.8 (4.5 6 0.2) 2.9–3.9 (3.4 6 0.3) referred to the species. The following data are 2.2–2.6 (2.4 6 0.1) 1.7–2.1 (1.9 6 0.2) from 36 specimens (Appendix) collected 2.2–3.3 (2.2 6 0.1) 1.7–2.1 (1.9 6 0.1) between 1997 and 2004. SVL in adult females 2.1–2.3 (3.6 6 0.2) 2.4–3.1 (2.8 6 0.2) 0.48–0.58 0.50–0.61 35.0–41.5 mm (n 5 10), in adult males 22.6– 0.47–0.51 0.45–0.52 28.8 (n 5 9); the smallest of 17 juveniles 0.35–0.40 0.34–0.41 (SMF 80400) is 11.4 mm. In addition to the 0.35–0.40 0.37–0.41 small tubercles in the scapular region noted 1.00–1.05 1.00–1.07 1.10–1.30 0.96–1.33 by Duellman and Hedges (2005), all speci0.46–0.63 0.47–0.66 mens have a pair of black scapular warts. 0.44–0.57 0.32–0.50 Duellman and Hedges (2005) mentioned only Females (n 5 6)

Males (n 5 8)

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a pair of black dots, but not the presence of warts. Males have vocal slits and a dorsal nuptial pad near the base of the thumb (nuptial pads absent according to Duellman and Hedges, 2005). The thumb is remarkably thickened in several specimens without apparently correlation to climatic season (discussed below). Distribution and ecology.—Previously known only from the type locality at 2120 m in Departamento de Pasco (Duellman and Hedges, 2005). The species is widely distributed in lowland and cloud forests (230–2320 m) in central Peruvian Departamentos Pasco, Junıń, and Ucayali. Specimens from Pozuzo and Bolognesi previously were misidentified as E. peruvianus (Lehr, 2002, 2005). Individuals were found at night on vegetation 5–200 cm above ground (Lehr, 2002). Five males (collecting dates in parentheses) have thickened thumbs: MHNSM 20375 (22 Jan. 1999), MTD 45920 (06 Dec. 2003), 45694 (25 May 2003), 46484 (11 Dec. 2004), SMF 80398 (23 Jan. 1999); the following males lack thickened thumbs: MTD 46472 (27 Nov. 2004), MHNSM 19828 (25 May 2003), MHNSM 20371 (16 Jul. 1998), SMF 80394 (12 Jul. 1998). A female (MTD 46483, 11 Dec. 2004) had 63 ovarian eggs (27 left, 36 right) with an average diameter of 2.6 6 0.12 mm (n 5 10). Sympatric species of Eleutherodactylus at lower elevations (230 m) include E. eurydactylus and E. rhabdocnemus, and those at higher elevations (above 1500 m) include E. cruciocularis, E. flavobracatus, and E. rhabdocnemus. Eleutherodactylus bromeliaceus Lynch 1979 Morales and Icochea (2000) recorded Eleutherodactylus bromeliaceus in northern and central Peru, and Duellman and Pramuk (1999) in northern Peru. In central Peru, we found E. bromeliaceus in arboreal bromeliads at elevations of 2180–2780 m. Males have a SVL of 18.0–22.6 mm (n 5 8), and one female (MHNSM 18660) has a SVL of 18.1 mm. The coloration in life of MHNSM 18660 was recorded by E. Lehr on 13 February 2003: ‘‘Dorsally pale olive, with brown blotches dorsolaterally; broad, salmon-colored band between eyes bordered posteriorly and anteriorly by a dark brown stripe, dark brown canthal and supratympanic

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stripe; upper lip dark brown, above cream stripe from tympanum to naris; throat, chest, and belly pale gray, extremities pale olive; iris reddish with black reticulations.’’ Sympatric Eleutherodactylus in central Peru include E. rhabdocnemus and E. sagittulus. Eleutherodactylus eurydactylus Hedges and Schlu¨ter 1992 This species was described from Panguana, Departamento de Huańuco, Peru, 200 m and reported from up to 1300 m elevation in the Serranıá de Sira (Hedges and Schlu¨ter, 1992). A specimen (MHNSM 20374) from Bolognesi (10u 069 130 S, 73u 499 020 W, 230 m) is the first record of this species from Departamento de Ucayali. This individual previously was misidentified as E. diadematus (Lehr 2001a, 2002). Sympatric Eleutherodactylus include E. bipunctatus and E. rhabdocnemus. Eleutherodactylus mendax Duellman 1978 Morales and Icochea (2000) supplemented the distributional information about Eleutherodactylus mendax in Peru. In the Departamento de Pasco, we found E. mendax (E. lacrimosus in Lehr, 2005) at elevations of 2850–3325 m. Individuals were found by day in arboreal or terrestrial bromeliads, and males were found calling at night from bamboo. 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 of a male (MTD 45900) was recorded by E. Lehr on 10 December 2003: ‘‘Dorsally and laterally yellow, pale greenish yellow behind eyes, dorsum with fine black blotches, tips of fingers and toes pale red, belly white, iris reddish copper with black reticulations.’’ Sympatric anurans include Bufo multiverrucosus, E. lundbergi, E. ornatus, and E. platydactylus, Gastrotheca stictopleura, Phrynopus juninensis, and Phyllonastes duellmani. Eleutherodactylus platydactylus (Boulenger 1903) Several authors (De la Riva, 1997; De la Riva et al., 2000; Ko¨hler and Lo¨tters, 1999) have reported that Eleutherodactylus platydactylus is highly polymorphic and may represent several sibling species. The species is known at elevations of 950–3470 m from

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central Peru to Bolivia (Ko¨hler, 2000). De la Riva (1997) redescribed E. platydactylus, noted 10 distinct color morphs, and diagnosed three putative new taxa for Bolivia. One species (E. llojsintuta), syntopic with E. platydactylus, was described as new mainly based on distinct advertisement calls (Ko¨hler and Lo¨tters, 1999). Herein we present the northernmost distributional records for the species in the central Peruvian Departamento de Pasco where we found E. platydactylus at elevations of 2500– 3275 m. At night, they were on low vegetation and on moist moss. By day they were found under dry leaves on the ground and in terrestrial bromeliads. Females have SVLs of 24.4–35.3 mm (n 5 11), and males have SVLs of 20.0–23.8 mm (n 5 8). Another series of specimens tentatively referred to E. platydactylus was obtained from localities in Departamento de Junıń (Pampa Hermosa, and below Pampa Hermosa near Rıó Ulcumayo), where it was abundant at night on low vegetation in lower montane forests at elevations of 1240– 1540 m. These frogs are smaller than specimens 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 yellowish brown, in contrast to grayish brown in E. platydactylus from Pasco. Sympatric anurans with E. platydactylus include Bufo multiverrucosus, Eleutherodactylus lundbergi, E. mendax, E. ornatus, Gastrotheca stictopleura, Phrynopus juninensis and Phyllonastes duellmani; sympatric with E. cf. platydactylus are E. bipunctatus, E. cruciocularis, and Gastrotheca atympana. Eleutherodactylus rhabdocnemus Duellman and Hedges 2005 Duellman and Hedges (2005) described E. rhabdocnemus based on two adult females from Departamento de Pasco, 2.9 km N and 5.5 km E Oxapampa, 2600 m. Herein, we refer additional specimens to E. rhabdocnemus obtained from Departamentos Pasco and Ucayali (see Appendix). Eleutherodactylus rhabdocnemus is highly variable in color and pattern, and we recognize remarkable morphological similarities with E. lirellus. Further research is necessary to assess the validity of E. rhabdocnemus.

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Sympatric anurans at the localities in Departamento de Pasco include Adenomera sp., Atelopus sp., Bufo cf. margaritifer, Eleutherodactylus bipunctatus, E. flavobracatus, Gastrotheca testudinaea, Hyalinobatrachium bergeri, Hybsiboas cf. melanopleura, Leptodactylus rhodonotus, and Oreobates quixensis. Eleutherodactylus stictogaster Duellman and Hedges 2005 This species was described by Duellman and Hedges (2005) based on a single subadult female with a SVL of 20.5 mm from near Oxapampa, 2600 m, We report the second known specimen (MTD 46474), an adult female with ovarian eggs, from Marıá 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 the holotype. In life, the iris was gold with black reticulations, and the groin and anterior surfaces of thighs pale red. Males of this species are unknown. Sympatric anurans at Marıá Teresa include Bufo cf. margaritifer, E. bipunctatus, E. rhabdocnemus, Gastrotheca testudinaea, Hyalinobatrachium bergeri, Hybsiboas cf. melanopleura, Oreobates quixensis, and Leptodactylus rhodonotus. DISCUSSION Currently, 91 species of Eleutherodactylus are known from Peru (updated from Frost [2004]); these represent about 25% of the 376 species of anurans known to inhabit Peru (AmphibiaWeb, 2005). Because of extensive fieldwork in the Departamento de Pasco (localities around Oxapampa, Pozuzo, and Paucartambo), the species diversity of this area is relatively well known. However, because herpetological surveys have not been conducted in many areas in Peru, the number of species of Eleutherodactylus undoubtedly will continue to increase including additional localities and new species. Species descriptions of Eleutherodactylus should be detailed and include photos (in life), as well as drawings of the head (at least in lateral view), foot, and hand in ventral aspect so as to

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facilitate further studies, comparisons and identifications of the species. In central Peru, Andean species of Eleutherodactylus (Fig. 5) can be found between 200 and 3325 m elevation; the single species of Phyllonastes occurs at 2900 m, and Phrynopus occur between 2600 and 4390 m. Two species, E. bipunctatus and E. rhabdocnemus, have relatively wide elevational (230– 2320 m in E. bipunctatus; 230–2600 m in E. rhabdocnemus) and geographical distributions, but most species of Eleutherodactylus have limited vertical and spatial distributions. Except for Phrynopus kauneorum and P. montium, most of the species of Phrynopus have even more limited vertical and spatial distribution and in many cases are known only from their type localities. Between 1000 and 2600 m the number of species of eleutherodactyline frogs increases from four at 1000 m, five at 1400 m, six at 2200 m, to a maximum of eight at 2600 m. Above 2600 m elevation the species number decreases except for 3250 m where the species number slightly increases to six from five at 3000 m. The greatest density of species of Eleutherodactylus has been recorded by Lynch and Duellman (1997) between 2500 and 2700 in the Eastern Andes of Ecuador. Thus, this elevational range in the eastern Andean slopes seems to provide optimal life conditions (e. g. humidity) for Eleutherodactylus. Among the 29 eleutherodactyline frogs referred to in Figure 5, 16 species (55%) lack a tympanum. These include all species of Phrynopus (Lehr et al., 2005a, b), the single Phyllonastes, and three species of Eleutherodactylus. The absence of a tympanum in central Peruvian eleutherodactyline frogs seems to be associated with increasing elevation—25% lack a tympanum at elevations below 1000 m, 33% at 1000–2000 m, 36% at 2000–3000 m, 77% at 3000–4000 m, and 100% above 4000 m. The way these frogs communicate or hear, if at all, is unknown. No calls have been recorded during fieldwork. Other Andean anurans such as Bufo spinulosus (Bufonidae) and Pleurodema marmorata (Leptodactylidae) are known from high elevations (up to 4175 m in B. spinulosus; up to 4480 m in P. marmorata) in the Andes of central Peru (Lehr, 2002) and both species

[No. 20

have a tympanum and vocalization. But B. spinulosus and P. marmoratum produce free swimming-tadpoles whereas eleutherodactyline frogs have direct development. Thus, reproductive mode may be associated with the acoustic abilities. Secondary sex characters in male Eleutheodactylus include vocal apparatus (vocal slits and vocal sac) and nuptial pads (Lynch and Duellman, 1997). Depending on the species, male Eleutherodactylus have nuptial pads present or absent. If present, nuptial pads are usually located basally on the first finger covering its dorsal and lateral surfaces including outer parts of the inner palmar and subarticular tubercles (e.g. as in many species of the E. conspicillatus Group). In some species (e.g. E. bipunctatus) nuptial pads are thickened on Finger I. Compared to other anurans (e.g. bufonids or hylids) males of which have brown and spinous nuptial pads that are clearly distinct from the surrounding skin, nuptial pads in Eleutherodactylus are usually nonspinous and glandular, usually white or cream colored and less pigmented than the surrounding skin. Thus, nuptial pads in Eleutherodactylus can easily be overseen, especially in badly preserved or old material. The use of a microscope with high magnification often is necessary to detect nuptial pads. It is unknown whether nuptial pads are permanent in adult male Eleutherodactylus or occur only during reproductive seasons. This phenomenon needs study of large series of specimens from throughout the year. At least in the specimens of E. bipunctatus studied herein, nuptial pads seem to be permanently present, possibly indicating that this species reproduces throughout the year. Therefore, nuptial pads are good characters to separate sexes and are helpful to distinguish species. RESUMEN Se describen cuatro especies nuevas procedentes de bosques montanos en los Departamentos de Huańuco, Junıń y Pasco en Peru´ central, con elevaciones que van desde los 1330–3000 m en la Cordillera Oriental. Tres de las especies nuevas se asignan al grupo de Eleutherodactylus unistrigatus. Dos de ellas carecen de tı´mpano, pero presentan una

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coloracioń y patroń de iris diferentes: una tiene la ingle y los miembros posteriores parcialmente coloreados de anaranjado a rojo, y el iris con una raya vertical parda y otra horizontal negra formando una cruz, mientras que la otra tiene la ingle y las patas posteriores parcialmente amarillas, y el iris con una raya horizontal parda y otra vertical negra desde la pupila al margen inferior del ojo, formando una ‘‘T.’’ La tercera especie, asignada a la serie de Eleutherodactylus conspicillatus, es similar a E. rhabdolaemus, pero difiere de e´sta en caracteres morfolo´gicos y de coloracioń. La cuarta especie asignada a la agrupacioń de E. lacrimosus tiene el dorso tabaco amarillento con manchas y rayas oscuras pardo purpu´reo, con flancos y vientre blancos, habita bromeliaćeas terrestres, y es comparada con E. schultei del norte de Peru´. Se anãden datos morfolo´gicos y de distribucioń para las siete especies recientemente descritas de Eleutherodactylus del Peru´ central. Se compara la distribucioń altitudinal entre los Eleutherodactylus, Phyllonastes, y Phrynopus de Peru´ central. Acknowledgments.—For comments on the manuscript we are grateful to R. Diaz, W. E. Duellman, and L. Trueb. We thank J. R. McCranie, I. De la Riva, and one anonymous reviewer for helpful comments on the manuscript. Specimens were loaned by J. H. Co´rdova (MHNSM), U. Fritz (MTD), G. Ko¨hler (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 Collection Study Grant. C. W. Myers (AMNH) provided photos and a color in life description for a specimen referred to E. cruciocularis. The research was supported by a postdoctoral grant given to E. Lehr by the Alexander von Humboldt-Foundation. We thank R. Acero for collecting and export permits issued by the Ministerio de Agricultura (INRENA), Lima, Peru.

LITERATURE CITED AMPHIBIA WEB: Information on amphibian biology and conservation. [web application]. 2005. Berkeley, California: AmphibiaWeb. Available at: http://amphibiaweb. org/. (Accessed: Jun 17, 2005). BRACK, A. 1986. Las Ecoregiones del Peru´. Boletıń de Lima 44:57–70. CANNATELLA, D. C. 1985. The systematic status of Syrrhophus juninensis Shreve (Anura: Letodactylidae). Proceedings of the Biological Society of Washington 98:774–777. DE LA RIVA, I. 1997. Redescription, variation, biology, and distribution of Eleutherodactylus platydactylus (Boulenger, 1903) (Amphibia: Anura: Leptodactylidae), taxonomic status of E. bockermanni Donoso-Barros,

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1970, and comments on polymorphism in anurans. Revista Espanõla de Herpetologıá 11:71–90. DE LA RIVA, I., J. KO¨HLER, S. LO¨TTERS, AND S. REICHLE. 2000. Ten years of research on Bolivian amphibians: updated checklist, distribution, taxonomic problems, literature and iconography. Revista Espanõla de Herpetologıá 14:19–164. DUELLMAN, W. E. 1978a. Two new species of Eleutherodactylus (Anura: Leptodactylidae) from the Peruvian Andes. Transactions of the Kansas Academy of Science 81:65–71. ———. 1978b. Three new species of Eleutherodactylus from Amazonian Peru´ (Amphibia: Anura: Leptodactylidae). Herpetologica 34:264–270. ———. 1978c. New species of leptodactylid frogs of the genus Eleutherodactylus from the Cosn˜ipata Valley, Peru´. Proceedings of the Biological Society of Washington 91:418–430. ———. 1990. A new species of Eleutherodactylus from the Andes of Northern Peru (Anura: Leptodactylidae). Journal of Herpetology 24:348–350. ———. 2000. Leptodactylid frogs of the genus Phrynopus in northern Peru with descriptions of three new species. Herpetologica 56:273–285. ———. 2005. Cusco Amazońico. The lives of amphibians and reptiles in an Amazonian rainforest. Cornell University Press, Ithaca and London. DUELLMAN, W. E., AND S. B. HEDGES. 2005. Eleutherodactyline frogs (Anura: Leptodactylidae) from the Cordillera de Yanachaga in Central Peru. Copeia 2005:526–538. DWYER, C. M. 1995. A new species of Eleutherodactylus from Peru´ (Anura: Leptodactylidae). Amphibia-Reptilia 16:245–256. FROST, D. R. 2004. Amphibian Species of the World: an Online Reference. Version 3.0 (22 August, 2004). Electronic Database accessible at http://research.amnh. org/herpetology/amphibia/index.html. American Museum of Natural History, New York, New York, U.S.A. HEDGES, S. B. 1990. A new species of Phrynopus (Anura: Leptodactylidae) from Peru. Copeia 1990:108–112. HEDGES, S. B., AND A. SCHLU¨TER. 1992. Eleutherodactylus eurydactylus, a new species of frog from central Amazonian Peru´ (Anura: Leptodactylidae). Copeia 1992:1002–1006. KO¨HLER, J. 2000. Amphibian diversity in Bolivia: a study with special reference to montane forest regions. Bonner Zoologische Monographien 48:1–243. KO¨HLER, J., AND S. LO¨TTERS. 1999. New species of the Eleutherodactylus unistrigatus Group (Amphibia: Leptodactylidae) from Montane Rain Forest of Bolivia. Copeia 1999:422–427. LEHR, E. 2001a. New records for amphibians and reptiles from Departamentos Pasco and Ucayali, Peru. Herpetological Review 32:130–132. ———. 2001b. A new species of Phrynopus (Anura: Leptodactylidae) from the eastern Andean slopes of central Peru. Salamandra 37:11–20. ———. 2002. Amphibien und Reptilien in Peru: Die Herpetofauna entlang des 10. Breitengrades von Peru: Arterfassung, Taxonomie, o¨kologische Bemerkungen und biogeographische Beziehungen. Natur und Tier Verlag (NTV Wissenschaft), Mu¨nster, Germany.

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———. 2005. A new species of the Eleutherodactylus nigrovittatus group (Anura: Leptodactylidae) from Andean Peru. Herpetologica 66:199–208. LEHR, E., AND C. AGUILAR. 2002. A new species of Phrynopus (Amphibia, Anura, Leptodactylidae) from the puna of Maraypata (Departamento de Huańuco, Peru). Zoologische Abhandlungen (Dresden) 52:57– 64. ———. 2003. A new species of Phrynopus (Amphibia, Anura, Leptodactylidae) from the puna of Maraypata (Departamento de Huańuco, Peru). Zoologische Abhandlungen (Dresden) 53:87–92. LEHR, E., C. AGUILAR, AND G. KO¨HLER. 2002a. Two sympatric new species of Phrynopus (Anura: Leptodactylidae) from a cloud forest in the Peruvian Andes. Journal of Herpetology 36:208–216. LEHR, E., C. AGUILAR, AND J. H. CO´RDOVA. 2002b. Morphological and ecological remarks on Phrynopus kauneorum (Amphibia, Anura, Leptodactylidae). Zoologische Abhandlungen (Dresden) 52:71–75. LEHR, E., C. AGUILAR, AND M. LUNDBERG. 2004a. A new species of Phyllonastes from Peru (Anura: Leptodactylidae). Journal of Herpetology 38:214–218. LEHR, E., C. AGUILAR, AND W. E. DUELLMAN. 2004b. A striking new species of Eleutherodactylus from Andean Peru (Anura: Leptodactylidae). Herpetologica 60:275– 280. LEHR, E., G. FRITZSCH, AND A. MU¨LLER. 2005b. An analysis of Andes frogs (Phrynopus, Leptodactylidae, Anura) phylogeny based on 12S and 16S mitochondrial rDNA sequences. Zoologica Scripta 34:593–603. LEHR, E., G. KO¨HLER, AND E. PONCE. 2000. A new species of Phrynopus from Peru (Amphibia, Anura, Leptodactylidae). Senckenbergiana biologica 80:205– 212. LEHR, E., M. LUNDBERG, AND C. AGUILAR. 2005a. Three new species of Phrynopus from central Peru (Amphibia, Anura, Leptodactylidae). Copeia 2005:479–491. LEVITON, A. E., R. H. GIBBS, JR., E. HEAL, AND C. E. DAWSON. 1985. Standards in herpetology and ichthyology: part I. Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802–832. LYNCH, J. D. 1979. Leptodactylid frogs of the genus Eleutherodactylus from the Andes of southern Ecuador. The University of Kansas, Museum of Natural History, Miscellaneous Publication 66:1–62. ———. 1980. A taxonomic and distributional synopsis of the Amazonian frogs of the genus Eleutherodactylus. American Museum Novitates 2696:1–24. LYNCH, J. D., AND W. E. DUELLMAN. 1980. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). The University of Kansas, Museum of Natural History, Miscellaneous Publication 69:1–86. ———. 1997. Frogs of the genus Eleutherodactylus in western Ecuador. Systematics, ecology, and biogeography. The University of Kansas, Natural History Museum Special Publication 23:1–236. MORALES, V. R., AND J. ICOCHEA. 2000. Review of the type material of Eleutherodactylus mendax and a new record of Eleutherodactylus bromeliaceus from Peru. Journal of Herpetology 34:158–160.

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APPENDIX I Comparative Specimens Examined Eleutherodactylus altamazonicus.—ECUADOR: NAPO: Santa Cecilia, 340 m: KU 146060–62, 148722–84; PERU: HUAŃUCO: Finca Panguana, Rıó Llullapichis, 4–5 km upstream from Rıó Pachitea, ca. 200 m: KU 154759; MADRE DE DIOS: 15 km E Puerto Maldonado, 200 m: KU 209956, 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 (paratypes); PASCO: ca. 12 km east of Oxapampa, Cumbre de Olloń, 2300 m: KU 206102 (paratype). Eleutherodactylus bipunctatus.—PERU: PASCO: 0.7 km S, 4.5 km E (airline) Oxapampa: KU 291638 (holotype); PASCO: Provincia de Pasco, Distrito de Paucartambo: km 85.2 (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 Oxapampa, Distrito de Chontabamba: km 24 on road from Oxapampa 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: Marıá Teresa, 10u 429 05.60 S, 75u 279 22.20 W, 1470 m: MTD 46335–36, 46472–73, 46483–84; PASCO: Provincia de Oxapampa: on road to Laguna San Daniel, 2320 m, near Parque Nacional Yanachaga-Chemilleń: MTD 46343; PASCO: Provincia de Oxapampa, Distrito de Palcazu´: Iscozacıń (Cordillera San Matıás), 10u 109 410 S, 75u 079 540 W, 410 m: SMF 80400; PASCO: Provincia de Oxapampa, Distrito de Pozuzo: Pozuzo, 10u 049 100 S, 75u 319 420 W, 940–1000 m: MHNSM 20371–73, SMF 80392–95; PASCO: Provincia de Oxapampa, Distrito de Pozuzo: Pozuzo (Yulitunqui), 10u 109 230 S, 75u 359 130 W, 1000 m: SMF 80396–97; JUNIŃ: 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: MORONA-SANTIAGO: Mirador: KU 174524 (paratype); MORONASANTIAGO: 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 km N, 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 179 520 W, 2780 m: MTD 46309–11; PASCO: Oxapampa: San Alberto, ca. 10u 329 530 S, 75u 229 170 W, 2200 m: MHNSM 18660, 19945–46, MTD 45210–11, 45987. Eleutherodactylus colodactylus.—PERU: PIURA: summit cordillera between Canchaque and Huancabamba, 3100 m: KU 135494, 135496–501; PIURA: 31 km W Huancabamba, 3080 m: KU 181262–64; PIURA: 33 km W Huancabamba, ca. 3050 m: KU 196443–61. Eleutherodactylus diadematus.—PERU: PASCO: Pozuzo, 10u 049 100 S, 75u 319 420 W, 970 m: MHNSM 20383–84, SMF 80410; PASCO: Pozuzo, 10u 089 560 S, 75u 309 550 W, 1110 m: MHNSM 20392; PASCO: Pozuzo: Yulitunqui, 10u 109 230 S, 75u 359 130 W, 930 m: MHNSM 20393.

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Eleutherodactylus eurydactylus.—PERU: HUAŃUCO: 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 DIÓS: Puerto Maldonado, 200 m: KU 205139–41, 207709–14, 215474–76, 215478. Eleutherodactylus lacrimosus.—ECUADOR: NAPO: Isla Anaconda, Rıó Napo, 25 km down Lago Agrio: KU 202406, 297131; PASTAZA: Mera, 1140 m: KU 119513–17, 119522–23, 120247–49, 132631–32, 177361–82; PASTAZA: Rıó 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 MARTIŃ: Rioja, Rıó Cerranayacu, 76 km NW Roja, 1200 m: KU 212226– 31, 212238–9 (all paratypes); SAN MARTIŃ: Lamas: W slope Abra Tangarana, 7 km NW San Juan Pacaysapa, 1080 m: KU 212240 (holotype), KU 212241, 212250 (paratypes); HUAŃUCO: Rıó Llullapichis, 4–5 km upstream from Rıó Pachitea, 200 m: KU 171893–94. Eleutherodactylus mendax.—PERU: CUZCO: Pistipata, Rıó Umasbamba, 12 km SE Huyro, 1820 m: KU 173234 (holotype), KU 173235 (paratype); CUZCO: 4 km SW Santa Isabel, 1700 m: KU 162290 (paratype); PASCO: Auquimarca, Puqiopata forest, 10u 43,759 S, 75u 42,809 W, 3310 m: MTD 44329; 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 from Uchuhuerta to Auquimarca, 10u 399 16.70 S, 75u 499 32.50 W, 2850 m: MHNSM 19865; PASCO: near road from Uchuhuerta to Auquimarca, 10u 429 12.50 S, 75u 449 33.90 W, 3000 m: MHNSM 19866, MTD 45899–00; PASCO: Paucartambo, 2932 m: MTD 44381–82, 44773. Eleutherodactylus platydactylus.—PERU: PASCO: Paucartambo: Puquiopata, 10u 43, 709 S, 75u 42,759 W, 3275 m: MHNSM 20691; PASCO: Paucartambo: Auquimarca, 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: Huachoń: 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 Auquimarca, 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 to Auquimarca, 10u 419 05.60 S, 75u 459 28.70 W, 2900 m: MHNSM 19862–64, MTD 45894–96; PASCO: Paucartambo: near road from Auquimarca to Uchuhuerta, 10u 439 56.70 S, 75u 429 00.40 W, 2840 m: MTD 45901; PASCO: Huachoń: Uchuhuerta, 2500 m: KU 291685, MHNSM 19874, MTD 45911; PASCO: Paucartambo: Chinche, 3000 m: MHNSM 20663. Eleutherodactylus cf. platydactylus.—PERU: JUNIŃ: Tarma: Huasahuasi: Pampa Hermosa, ca. 10u 599 33.30 S, 75u 259 58.00 W, 1540 m: MHNSM 18693–97, 19801,

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20692–98, MTD 45645–59; JUNIŃ: Chanchamayo: below Pampa Hermosa, near Rıó Ulcumayo, ca. 11u 009 30.40 S, 75u 259 36.00 W, 1240 m: MHNSM 19802–03, MTD 45660–61. Eleutherodactylus rhabdocnemus.—PERU: PASCO: 2.9 km north 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 from Oxapampa–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: MHNSM 19847, 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 to Laguna 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: Marıá Teresa, 10u 429 05.60 S, 75u 279 22.20 W, 1470 m: MHNSM 19551, MTD 46470–71, 46475–82; PASCO: Oxapampa: km 53 on road 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, 10u 119 00.50 S, 75u 349 21.10 W, 1050 m: MTD 46322; PASCO: Pozuzo, 10u 049 100 S, 75u 319 420 W, 970 m: MHNSM 20378–82, 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: MHNSM 20394, SMF 8417; PASCO: Pozuzo, 10u 089 560 S, 75u 309 550 W, 1110 m: SMF 80411–14; UCAYALI: Campamento Bolognesi, 10u 069 130 S, 73u 499 020 W, 230 m: SMF 80399. Eleutherodactylus rhabdolaemus.—BOLIVIA: COCHABAMBA: 68.8 km SW Villa Tumari, 1860 m: KU 183009; PERU: CUZCO: 7 km Santa Isabel, 1900 m: KU 13887 (paratype); AYACUCHO: Huanhuachayocc, on Tambo trail: KU 175082–83 (paratypes); AYACUCHO: between Mitupucuru and Estero Ruana: KU 175084 (paratype); CUZCO: Buenos Aires, 2400 m: KU 173237–53 (5 paratypes of E. pharangobates); CUZCO: Huyro, 1720 m: KU 175086–88. Eleutherodactylus schultei.—PERU: AMAZONAS: 5 km N 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 km N, 5.5 km E (airline) Oxapampa: KU 291659 (holotype); PASCO: Oxapampa: Chontabamba: Marıá Teresa, 10u 429 05.60 S, 75u 279 22.20 W, 1470 m: MTD 46474. Eleutherodactylus toftae.—PERU: HUAŃUCO: Rıó Llullapichis, 4–5 km upstream from Rıó Pachitea, ca. 200 m: KU 154806–19, 171852–62, 171864–66.

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

Altitudinal range (m above sea level)

E. aniptopalmatus E. bipunctatus E. bromeliaceus E. cruciocularis E. diadematus E. eurydactylus E. flavobracatus E. lundbergi E. mendax E. ornatus E. pardalinus E. platydactylus E. cf. platydactlus E. rhabdocnemus E. sagittulus E. stictogaster Phyllonastes duellmani Phrynopus barthlenae P. bracki P. bufoides P. dagmarae P. heimorum P. horstpauli P. juninensis P. kauneorum P. montium P. paucari P. pesantesi P. tautzorum

2300–2600 230–2320 2180–2780 1540 930–1110 200–1300 1770 1800–2760 2850–3325 2550–3000 2640 2500–3275 1240–1540 230–2600 2050–2200 1470–2600 2900 3425–3680 2600 3850–4100 3020–3380 3420–3430 3030–3430 3650 2735–3380 2950–3660 3600 4280–4390 3770

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