South American Journal of Herpetology, 4(2), 2009, 125-138 © 2009 Brazilian Society of Herpetology

THREE NEW SPECIES OF (ANURA: ) FROM THE REGION OF CUSCO, PERU

Edgar Lehr1,3 and Alessandro Catenazzi2

1 Senckenberg Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstrasse 159, D‑01109 Dresden, Germany. 2 Department of Integrative Biology, University of California at Berkeley. 3060 Valley Life Sciences Bldg #3140, Berkeley CA 94720, USA. Email: [email protected] 3 Corresponding Author: [email protected]

Abstract. We describe three new species of Bryophryne from the Region of Cusco in southern Peru, increasing the number of currently known Bryophryne to six. One of the new species differs from all species of Bryophryne in having a tympanic annulus and tympanic membrane. Males of this species have vocal slits, a vocal sac, and produce a call, which we analyze herein. This species is found at San Luis, a montane cloud forest along the road from Abra Málaga to Quillabamba at elevations between 3272 and 3354 m. The second new species has an orange throat and groin and is found along the Ericsson trail that connects Acjanaco to Pillahuata in Manu National Park and near Abra Acjanaco along the Paucartambo-Pilcopata road at elevations between 3266 and 3430 m. The third new species has the throat and chest mottled pale gray and tan, whereas the belly is black with white flecks. This species is only known from the upper Marcapata valley along the road from Abra Huallahualla to Quincemil at elevations between 3129 and 3285 m. Ecological observations for all new species are described and a map showing the type localities of all currently known species of Bryophryne is presented.

Keywords. Anura, Bryophryne, new species, Strabomantidae, vocalization.

Introduction odontophores (Duellman et al., 2006). We follow the definition of conditions of the tympanum by Lynch The genus Bryophryne was described by Hedges and Duellman (1997). Specimens were preserved in et al. (2008). Based on DNA sequences, their phy- 10% formalin and stored in 70% ethanol. Specimens logenetic analysis of strabomantid recognized were dissected to determine sex and maturity, and a distinct clade among Holodeninae that consisted the otic region was dissected in order to determine of the single species B. cophites (formerly Phryno‑ condition of the tympanic annulus. We measured the pus cophites Lynch, 1975). bustamantei following variables to the nearest 0.1 mm with dig- Chaparro, De la Riva, Padial, Ochoa, and Lehr, 2007, ital calipers under a microscope: snout-vent length was assigned to Bryophryne by Hedges et al. (2008) (SVL), tibia length (TL), foot length (FL, distance based on morphological similarities with B. cophites. from proximal margin of inner metatarsal tubercle to Lehr and Catenazzi (2008) recently described B. nu‑ tip of Toe IV), head length (HL, from angle of jaw bilosus from the Region of Cusco in southern Peru. to tip of snout), head width (HW, at level of angle of Both authors hypothesized that the deep valley of the jaw), eye diameter (ED), tympanum diameter (TY), Rio Apurimac is a biogeographic border separating interorbital distance (IOD), upper eyelid width (EW), Phrynopus from Bryophryne (Lehr and Catenazzi, internarial distance (IND), eye-nostril distance (E‑N, 2008), and pointed out that new species will likely be straight line distance between anterior corner of or- discovered when fieldwork continues. Herein we de- bit and posterior margin of external nares). We deter- scribe three new species of Bryophryne recently dis- mined comparative lengths of Toes III and V by ad- covered in the Region of Cusco southeast of the Rio pressing both toes against Toe IV; lengths of Fingers Apurimac valley, thus doubling the number of species I and II were determined by adpressing the fingers within the genus. against each other. Drawings were made by the senior author using a stereomicroscope with drawing tube attachment. Photographs taken by A. Catenazzi were Materials and Methods used for descriptions of color in life and are available for all types at the Calphoto online database (http:// follows Hedges et al. (2008) and the calphotos.berkeley.edu). format for the description follows that of Lynch and We recorded advertisement calls for one of the Duellman (1997), except that we used the term den- new species with a Sony TCM‑150 audio tape-re- tigerous processes of vomers instead of vomerine corder equipped with a Azden SMX‑10 microphone. 126 Three new species of Bryophryne

We used Cool Edit version 96 (Syntrillium Software pads absent; (7) Finger I shorter than Finger II; tips Corporation) and Raven Lite, version 1.0 (Cornell of digits rounded; (8) fingers with lateral fringes; Laboratory of Ornithology) to digitalize calls and (9) ulnar and tarsal tubercles present; (10) heel with analyze sonograms. We digitized and edited vocaliza- small tubercles; inner tarsal fold absent; (11) in- tions at a sampling frequency of 44 KHz, FFT with ner metatarsal tubercle ovoid, about twice as large 512 points, and 16‑bit resolution. as rounded outer metatarsal tubercle; supernumer- Locality names follow the spelling of the US ary plantar tubercles indistinct; (12) toes with lateral Board on Geographic Names (http://gnswww.nga. fringes; basal webbing present; Toe V shorter than mil) and, for localities not listed in this database, ac- Toe III; toe tips rounded, about as large as those on cording to Cartas Nacionales “Calca” (Hoja 27‑s), fingers; (13) in life, dorsum reddish brown, grayish “Corani” (28‑u), “Ocongate” (28‑t) and “Urubamba” brown, purplish brown, or dark gray, with narrow tan (27‑r), Instituto Geográfico Nacional, Lima (except middorsal stripe, venter dark brown, tan, or reddish for Acjanaco = Acanaco, Acanacu). We deposited brown with pale gray flecks; (14) SVL in adult fe- specimens in the herpetological collections of the Mu- males 16.0‑22.2 mm (n = 10), in males 16.7‑19.3 mm seo de Historia Natural, Universidad Nacional Mayor (n = 10). de San Marcos (MUSM) in Lima, Peru, Museum für Tierkunde Dresden (MTD), Germany, the Museum of Bryophryne gymnotis is readily distinguished Vertebrate Zoology, University of California Berke- from the other currently known five species (includ- ley (MVZ), and the Musée d’Histoire Naturelle de la ing the ones described herein) in having a tympanic Ville de Genève, Geneva, Switzerland (MHNG). For membrane and tympanic annulus, and a smooth (are- specimens examined, see Appendix. olate in all others) venter. Furthermore, it differs from all its congeners in having the dorsal color uniform reddish brown, gray, dark brown, or purplish brown, Bryophryne gymnotis sp. nov. (Figs. 1 and 2) with a narrow, tan middorsal stripe, and a ventral

Holotype – MUSM 24543. Adult male, collected 1 km east of San Luis (13°04’32.1”S, 72°22’55.3”W) at elevations of 3272‑3354 m, Distrito de Huayopata, Provincia de La Convención, Región Cusco, Peru, on 8 March 2008 by A. Catenazzi, I. Chinipa, J. C. Jahu- anchi, and A. Machaca.

Paratypes – 29: 10 females (MHNG 2710.28, MTD 46860‑61, 47297, MUSM 24546‑50, MVZ 258407), 8 males (MTD 47288, 47291‑92, MUSM 24541‑42, 24544‑45, MVZ 258408‑09), 11 juveniles (MNNG 2710.29, MTD 46862‑64, MUSM 24551‑56, MVZ 258410), all collected with the holotype on 8 March 2008 by A. Catenazzi, I. Chinipa, J. C. Jahuanchi, and A. Machaca.

Diagnosis – A medium sized species of Bryophryne having the following combination of characters: (1) skin on dorsum shagreen, skin on venter smooth; discoidal fold absent, thoracic fold present; nar- row, discontinuous dorsolateral fold; (2) tympanic membrane and tympanic annulus present; (3) snout rounded in dorsal and lateral views; (4) upper eye- lid without enlarged tubercles; width of upper eyelid narrower than IOD; cranial crests absent; (5) dentig- Figure 1. Holotype of Bryophryne gymnotis in life (MUSM erous processes of vomers present, minute, oblique; 24543, male) in lateral (A), and ventral (B) views. Photos by A. (6) males with vocal sac and vocal slits, but nuptial Catenazzi. Lehr, E. and Catenazzi, A. 127 color of tan, reddish brown, or dark brown with pale one half free; vocal slits straight, located at posterior gray flecks. Table 1 compares selected characters half of mouth floor between tongue and margin of among species of Bryophryne. jaw; vocal sac distinct.

Description of holotype – Head narrower than body, Skin on dorsum shagreen, short occipital fold slightly wider than long, HW 102.8% of HL; HW merging into narrow dorsolateral fold extending to 40.2% of SVL; HL 39.1% of SVL; snout short, acute- sacral region, anterior part continuous, posterior part ly rounded in dorsal view, rounded in lateral view discontinuous; skin on flanks slightly tuberculate (Figs. 2A, B), ED larger than E‑N distance; nostrils with tubercles coalescing into long, narrow fold on slightly protuberant, directed dorsolaterally; canthus upper part on both side of flanks; skin on throat, chest, rostralis straight in dorsal view, rounded in profile; and belly smooth; discoidal fold absent, thoracic fold loreal region slightly concave; lips rounded; upper present; cloacal sheath absent; large tubercles absent eyelid without enlarged tubercles; EW narrower than in cloacal region. Outer surface of forearm with four IOD (EW 58.3% of IOD); supratympanic fold nar- minute tubercles (= ulnar tubercles, only two are vis- row, slightly curved, extending from posterior corner ible in Fig. 2C) arranged in a longitudinal row; outer of eye to insertion of arm; tympanic membrane and and inner palmar tubercles low, ovoid, about equal tympanic annulus present, weakly defined, upper and in size; supernumerary tubercles distinct, low, ovoid, posterior margin concealed by supratympanic fold; about one third size of subarticular tubercles; sub- two enlarged postrictal tubercles on each side of head. articular tubercles prominent, ovoid in dorsal view, Choanae small, ovoid, not concealed by palatal shelf rounded in lateral view, most prominent on base of of maxilla; dentigerous processes of vomers present, fingers; fingers with lateral fringes; Finger I shorter minute, embedded in buccal mucosa, oblique; tongue than Finger II; tips of digits rounded lacking marginal 3x as long as wide, not notched posteriorly, posterior grooves (Fig. 2C).

Figure 2. Dorsal (A) and lateral (B) views of head and ventral views of left hand (C) and right foot (D) of Bryophryne gymnotis (MUSM 24543). Drawings by E. Lehr. 128 Three new species of Bryophryne

Table 1. Selected characters (+ = character present; — = character absent), character conditions, and elevational distribution among Bryophryne.

B. nubilosus B. bustamantei B. gymnotis B. hanssaueri B. zonalis B. cophites Lehr and Characters and source Chaparro et al., this paper this paper this paper Lynch, 1975 Catenazzi, 2008, 2007, this paper this paper Maximum SVL (mm) 22.2 24.6 24.4 23.4 29.3 21.9 16.0‑22.2 18.1‑24.6 22.2‑24.4 21.0‑23.4 19.2‑21.9 Female SVL (20.0 ± 2.2) (21.7 ± 1.8) (23.3 ± 1.1) (22.0 ± 1.2) 21.9‑29.3 (20.5 ± 1.0) n = 10 n = 12 n = 3 n = 3 n = 5 16.7‑19.3 12.3‑18.0 14.8‑17.6 19.0‑22.9 12.7‑18.9 Male SVL (18.1 ± 0.9) (15.5 ± 2.8) (16.0 ± 1.4) (21.0 ± 2.8) 18.0‑22.7 (14.9 ± 2.2) n = 10 n = 4 n = 3 n = 2 n = 5 Tympanic annulus + — — — — — Tympanic membrane + — — — — — Dentigerous processes of + — — — — — vomers Dorsolateral folds +, discontinuous +, discontinuous +, discontinuous +, continuous +, discontinuous +, discontinuous Skin on venter smooth areolate areolate areolate areolate areolate Vocal sac + — — + — — Vocal slits + — — + — — Lateral fringes + + + — — — Nuptial pads — — — — + — Finger I < II + + + + + + Elevational distribution [m] 3272‑3354 3266‑3430 3129‑3285 3555‑3950 3195‑3700 2350‑3215

Hind limbs short, robust, TL 37.0% of SVL; FL midventral tan stripe extending to chest; chest, belly, 41.3% of SVL; upper surface of hind limbs shagreen and extremities mottled brown and tan with white with small, scattered tubercles; posterior and ventral and dark brown flecks; concealed surfaces of hind surfaces of thighs areolate; heel without small tu- limbs with white flecks; Fingers I‑III, Toes I‑III, and bercles; outer surface of tarsus with few minute tu- metatarsal tubercles tan, remaining ventral surfaces bercles; metatarsal tubercles prominent, inner meta- of hands and feet gray; upper half of iris pale bronze, tarsal tubercle ovoid, elevated, about twice the size lower half dark brown, pupil encircled by pale orange of round, low outer metatarsal tubercle; few plantar ringlet. supernumerary tubercles indistinct; subarticular tu- bercles low, ovoid in dorsal view; toes with lateral Color of holotype in preservative – Dorsum mottled fringes, basal webbing present; toe tips rounded, dark gray and dark brown with narrow pale gray mid- lacking marginal grooves, about as large as those on dorsal stripe extending from snout to cloaca; narrow fingers; relative lengths of toes: 1 < 2 < 5 < 3 < 4; Toe pale gray stripe extending on both sides from upper V shorter than Toe III (Fig. 2D). margin of cloaca diagonally on posterior surface of thigh to inside of knee; dark brown canthal and su- Measurements of holotype (in mm) – SVL 18.4; TL pratympanic stripes, latter more distinct; dark brown 6.8; FL 7.6; HL 7.2; HW 7.4; ED 1.9; TY 1.1; IOD bar below eye; upper and lower lips dark brown with 2.4; EW 1.4; IND 1.7; E‑N 1.5. pale gray spots; flanks as dorsum but slightly paler; tips of postrical tubercles white; throat mottled dark Color of holotype in life (Fig. 1) – Dorsum purplish brown and tan with white spots and midventral tan brown with pale gray spots and reddish brown flecks; stripe extending to chest; chest, belly, and extremi- narrow purplish gray middorsal stripe extending from ties mottled brown and tan with white and dark brown snout to cloaca; narrow purplish gray stripe extending flecks; concealed surfaces of hind limbs with white on both sides from upper margin of cloaca diagonally flecks; Fingers I‑III, Toes I‑III, and metatarsal tuber- on posterior surface of thigh to inside of knee; dark cles pale gray, remaining ventral surfaces of hands brown canthal and supratympanic stripes; dark brown and feet gray; iris gray. bar below eye; upper and lower lips dark brown with white spots; tips of postrical tubercles white; throat Variation – In life, all specimens have a tan middorsal mottled dark brown and tan with white spots and stripe but differ in dorsal ground color, which varies Lehr, E. and Catenazzi, A. 129

from reddish brown (MTD 47288), brown (MTD Table 2. Measurements (in mm) and proportions of Bryophryne 47292), grayish brown (MUSM 24547), and purplish gymnotis; ranges followed by means and one standard deviation brown (MUSM 24545, 24548) to dark gray (MVZ in parentheses. 258408). Two females (MUSM 24549‑50) have a tan B. gymnotis Characters dorsal band and a paler tan middorsal stripe. One juve- Females (n = 10) Males (n = 10) 16.0‑22.2 16.7‑19.3 nile (MUSM 24554) has the dorsum nearly uniformly SVL tan; two juveniles (MHNG 2710.29, MUSM 24553) (20.0 ± 2.2) (18.1 ± 0.9) have the dorsum pale purplish gray with dark brown TL 6.0‑7.5 (6.9 ± 0.4) 6.2‑6.8 (6.6 ± 0.2) blotches and dark brown bars on extremities. Ven- FL 7.0‑9.2 (8.1 ± 0.7) 6.9‑7.9 (7.5 ± 0.3) HL 6.4‑8.3 (7.5 ± 0.6) 6.7‑7.4 (7.0 ± 0.2) tral ground color varies from tan (MHNG 2710.28, HW 6.4‑8.5 (7.8 ± 0.7) 6.6‑7.9 (7.2 ± 0.4) MTD 47291) to brownish orange (MTD 47288, ED 1.8‑2.5 (2.1 ± 0.2) 1.8‑2.1 (1.9 ± 0.1) MUSM 24542) to dark brown (MUSM 24546, MVZ TY 0.7‑1.5 (1.0 ± 0.3) 0.7‑1.1 (1.9 ± 0.1) 258408). Two females (MUSM 24549‑50) and two IOD 2.2‑2.9 (2.6 ± 0.2) 2.3‑2.6 (2.4 ± 0.1) juveniles (MTD 46862, MUSM 24551) have a mid- EW 1.3‑1.7 (1.6 ± 0.1) 1.3‑1.7 (1.4 ± 0.1) ventral tan stripe crossed by a longitudinal tan stripe IND 1.5‑2.0 (1.7 ± 0.1) 1.3‑1.9 (1.7 ± 0.2) ventrally on arms and legs. Ventral surfaces of juve- E‑N 1.6‑1.9 (1.7 ± 0.1) 1.5‑1.8 (1.7 ± 0.1) niles are nearly uniformly dark brown with pale gray TL/SVL 0.32‑0.39 0.34‑0.38 spots. Males are smaller than females. The smallest FL/SVL 0.37‑0.46 0.40‑0.44 HL/SVL 0.34‑0.41 0.35‑0.41 juvenile is 10.3 mm SVL. Table 2 shows ranges and HW/SVL 0.38‑0.41 0.38‑0.41 proportions of the type series. HW/HL 0.96‑1.13 0.97‑1.10 E‑N/ED 0.72‑1.00 0.79‑0.95 Advertisement call – Several males of Bryophryne EW/IOD 0.50‑0.73 0.54‑0.68 gymnotis were calling during our visit to the type lo- TY/ED 0.38‑0.71 0.37‑0.58 cality between 1000 h and 1300 h on 8 March 2008. The recorded male (MUSM 24544, SVL 18.9 mm; call). Frequency modulation was slight, with the first T = 18.0°C) was found under moss and leaf litter on the part of the call having a fundamental frequency of ground. The call is a short, single note repeated at ir- 3.01 KHz (range 2.97‑3.02 KHz) and a harmonic at regular intervals and with a fundamental frequency of 4.45 KHz, and the second part having a fundamental 3.01 KHz (dominant) and one harmonic at 4.45 KHz frequency of 3.05 KHz (range 2.97‑3.07 KHz). (Fig. 3). The call duration ranged from 86 to 136 ms (mean = 121.0 ± 11.8 ms, n = 4). Calls were com- Distribution – The species is only known from the posed of two weakly modulated parts, the first part type locality of San Luis, a montane cloud forest having a generally ascendent amplitude modulation along the road from Abra Málaga to Quillabamba at and a duration of 54 to 82 ms (mean = 69.0 ± 7.6 ms, 3272‑3354 m. San Luis is 5.3 km from the closest n = 4), and the second part having a descendent am- locality of the congeneric B. bustamantei (Fig. 4). plitude modulation and a duration of 32 to 78 ms Based on available information, the elevational range (mean = 52.0 ± 9.7 ms, n = 4). Calls were produced of these two species does not seem to overlap. All every 29 to 75 s (mean = 50.0 ± 13.3 s, n = 3; time specimens were collected under moss, leaf litter, logs, from the end of one call to the beginning of the next and rocks on the ground during the day in the rainy

Figure 3. Waveform (above) and sonogram (below) of three advertisement calls (separated by intervals of 29‑75 s) of male Bryophryne gymnotis (MUSM 25444) recorded at the type locality (San Luis, Distrito de Huayopata, Provincia de La Convención, Región Cusco, Peru; elevation 3300 m) on 8 March 2008 at 1300 h (air temperature = 18.0°C). 130 Three new species of Bryophryne season. A clutch of B. gymnotis found at the type Peru. MUSM 24557, MVZ 258411 (13°11’29.4”S, locality contained 19 eggs ranging between 4.0 and 71°37’18.3”W) collected at 3381 m elevation on 4.8 mm in diameter. 23 February 2008; MTD 46865 (13°11’14.9”S, 71°37’18.6”W) at 3313 m elevation on 24 February Etymology – The specific name gymnotis is com- 2008; MTD 46866, MUSM 27568‑69, MVZ 258412 posed of the Greek adjective gymnos meaning “bare, (13°10’10.3”S, 71°37’51.2”W) at 3417 m elevation naked,” and the Greek noun “otos” meaning ear. The on 28 February 2008. Collected by A. Catenazzi, specific name refers to the character condition of the I. Chinipa, J. C. Jahuanchi, and A. Machaca. MTD new species, which currently is the only member of 46888, MUSM 27610 (13°11’28.3”S, 71°37’19.0”W) the genus having a tympanum. collected at 3360 m elevation on 23 October 2008. MHNG 2698.25, MUSM 27607, 27609, MVZ 258413 (13°11’15.5”S, 71°37’18.9”W) collected at Bryophryne hanssaueri sp. nov. (Figs. 5 and 6) 3320 m elevation on 23 October 2008. MTD 46889, MUSM 27608, 27611 (13°11’07.7”S, 71°37’16.9”W) Holotype – MUSM 27567. Adult female, collected at 3280 m elevation on 24 October 2008. MTD 46887 at Acjanaco, Manu National Park (13°10’58.4”S, (13°11’57.8”S, 71°36’27.3”W) at 3430 m on 25 Oc- 71°37’11.7”W) at 3266 m elevation, Distrito de tober 2008. Collected by A. Catenazzi, D. Cruz, J. C. Cosñipata, Provincia de Paucartambo, Región Cusco, Jahuanchi, and E. Luna. Peru, on 24 February 2008 by A. Catenazzi, I. Chini- pa, J. C. Jahuanchi, and A. Machaca. Diagnosis – A medium-sized species of Bryophryne having the following combination of characters: Paratypes – 17: 11 females (MHNG 2698.25, MTD (1) skin on dorsum shagreen with small scattered tu- 46865, 46887‑89, MUSM 24557, 27608‑11, MVZ bercles, skin on venter areolate; discoidal fold absent, 258411), 4 males (MTD 46866, MUSM 27568, thoracic fold present; short, irregularly shaped, dis- 27607, MVZ 258413), and two juveniles (MUSM continuous dorsolateral fold; (2) tympanic membrane 27569, MVZ 258412), all from near Acjanaco (= Aca- and tympanic annulus absent; (3) snout rounded in naco, Acanacu), Manu National Park, Distrito de dorsal and lateral views; (4) upper eyelid without en- Cosñipata, Provincia de Paucartambo, Región Cusco, larged tubercles; width of upper eyelid narrower than

Figure 4. Map illustrating the type localities of Bryophryne in Peru. Lehr, E. and Catenazzi, A. 131

IOD; cranial crests absent; (5) dentigerous processes life, dorsum pale grayish brown with a dark brown of vomers absent; (6) males without vocal sac, vocal X‑shaped or triangular blotch on head, with or without slits, and nuptial pads; (7) Finger I shorter than Finger narrow middorsal tan stripe, in females throat orange, II; tips of digits rounded; (8) fingers with lateral fring- chest and belly pale grayish white; in males venter es; (9) ulnar tubercles coalesced to narrow fold; tarsal brownish orange with grayish white flecks on chest tubercles present; (10) heel with small tubercles; in- and belly; (14) SVL in adult females 18.1‑24.6 mm ner tarsal fold absent; (11) inner metatarsal tubercle (n = 12), in males 12.3‑18.0 mm (n = 4). ovoid, twice as large as rounded outer metatarsal tubercle; supernumerary plantar tubercles indistinct; Bryophryne hanssaueri differs from the other (12) toes with lateral fringes; basal webbing present; currently known five species (including the ones Toe V usually longer than Toe III; toe tips slightly described herein) in that females have an orange pointed, about as large as those on fingers; (13) in (brownish or reddish orange in males) throat and or- ange (brownish or reddish orange in males) groin. Furthermore it differs from B. cophites in being smaller (maximum SVL 24.6 mm in B. hanssaueri vs. 29.3 mm: Lynch, 1975). Bryophryne hanssaueri dif- fers from B. gymnotis in lacking a tympanum. Male B. hanssaueri lack vocal slits and a vocal sac, both of which are present in B. gymnotis and B. bustamantei. Furthermore B. bustamantei has a continuous dorso- lateral fold (discontinuous, short in B. hanssaueri) and the flanks have prominent warts (small tuber- cles). Bryophryne hanssaueri and B. nubilosus both have flanks with tubercles and discontinuous dorsola- teral folds, but the folds are more prominent in B. nu‑ bilosus, and B. nubilosus has two slightly enlarged tubercles on the upper eyelid (absent in B. hanssau‑ eri). Bryophryne zonalis is larger than B. hanssaueri (max SVL 26.8 mm vs. 24.6 mm in B. hanssaueri) and female Bryophryne zonalis have a black belly with white flecks (belly pale grayish white in female B. hanssaueri). Table 1 compares selected characters among species of Bryophryne.

Description of holotype – Head narrower than body, wider than long, HW 115.4% of HL; HW 39.6% of SVL; HL 34.4% of SVL; snout short, rounded in dor- sal and lateral views (Figs. 6A, B), ED larger than E‑N; nostrils slightly protuberant, directed dorso- laterally; canthus rostralis straight in dorsal view, rounded in profile; loreal region slightly concave; lips rounded; EW without enlarged tubercles; EW nar- rower than IOD (EW 73.1% of IOD); supratympanic fold low, narrow; tympanic membrane and tympanic annulus absent; two enlarged postrictal tubercles on each side of head. Choanae small, ovoid, not con- cealed by palatal shelf of maxilla; dentigerous proc- esses of vomers absent; tongue 2x as long as wide, not notched posteriorly, posterior one third free.

Figure 5. Holotype of Bryophryne hanssaueri in life (MUSM 27567, female) in lateral (A), dorsal (B), and lateroventral (C) Skin on dorsum shagreen with small scattered tu- views. Photos by A. Catenazzi. bercles; dorsolateral fold from posterior margin of eye 132 Three new species of Bryophryne to sacral region, low, discontinuous; skin on flanks tubercle; plantar supernumerary tubercles indistinct; slightly tuberculate; skin on throat smooth, skin on subarticular tubercles low, ovoid in dorsal view, most chest and belly areolate; discoidal fold absent, tho- prominent on base of toes; toes with lateral fringes, racic fold present; cloacal sheath absent; large tuber- basal webbing present; toe tips slightly pointed, lack- cles absent in cloacal region. Outer surface of fore- ing marginal grooves, slightly smaller than those on arm (= ulnar) with a narrow fold extending from outer fingers; relative lengths of toes: 1 < 2 < 3 < 5 < 4; Toe margin of hand three-fourths distance to elbow; outer V longer than Toe III (Fig. 6D). and inner palmar tubercles low, outer ovoid, about twice size of inner elongate palmar tubercle; super- Measurements of holotype (in mm) – SVL 22.7; TL numerary tubercles indistinct; subarticular tubercles 7.9; FL 9.2; HL 7.8; HW 9.0; ED 2.4; IOD 2.6; EW prominent, ovoid in dorsal view, rounded in lateral 1.9; IND 2.2; E‑N 1.8. view, most prominent on base of fingers; fingers with lateral fringes; Finger I shorter than Finger II; tips Color of holotype in life (Fig. 5) – Dorsum pale gray- of digits slightly pointed, lacking marginal grooves ish brown with an irregularly shaped dark brown (Fig. 6C). blotch on head; flanks slightly paler, axilla mottled Hind limbs short, robust, TL 34.8% of SVL; FL pale gray and white; groin bright orange; dark brown 40.5% of SVL; upper surface of hind limbs shagreen canthal and supratympanic stripes, the latter less with small, scattered tubercles; posterior and ventral distinct; throat bright orange; chest, belly, and ex- surfaces of thighs areolate; heel with small tubercles; tremities (except hands and feet) pale grayish white outer surface of tarsus with few, minute tubercles; with grayish brown flecks; hands and feet ventrally metatarsal tubercles low, inner metatarsal tubercle brownish orange; iris greenish gold with fine black elongate, about twice size of ovoid outer metatarsal reticulations.

Figure 6. Dorsal (A) and lateral (B) views of head and ventral views of left hand (C) and left foot (D) of Bryophryne hanssaueri (MUSM 27567). Drawings by E. Lehr. Lehr, E. and Catenazzi, A. 133

Color of holotype in preservative – Dorsum grayish Paratypes – Nine: three females (MTD 46867, brown with an irregularly shaped brown blotch on MUSM 27572, MVZ 258414), three males (MTD head; dark brown canthal stripe; supratympanic stripe 46870, MUSM 27574, MVZ 258415), and three juve- indistinct; groin and throat pale orange; chest, belly, niles (MTD 46869, MUSM 27575, 27861). All were and extremities cream with grayish brown flecks; collected with the holotype (except MVZ 258415) on hands and feet ventrally pale gray; iris gray. 4 March 2008 by A. Catenazzi, I. Chinipa, J. C. Ja- huanchi, and A. Machaca. MVZ 258415 was collect- Variation – Adult females have a bright orange throat ed at Puente Coline (13°34’37.7”S, 71°00’23.8”W), and groin, whereas adult males have a brownish or 3285 m elevation, Distrito de Marcapata, Provincia reddish orange throat and groin. In juveniles, throat de Quipicanchis, Región Cusco, Peru, on 5 March and groin are tan, distinct dark brown canthal and su- 2008 by A. Catenazzi, I. Chinipa, J. C. Jahuanchi, and pratympanic stripes, a dark brown X‑shaped blotch A. Machaca. on head, and dark brown blotches on extremities on a tan dorsum are present. One juvenile (MUSM 27569) Diagnosis – A medium-sized species of Bryophryne has a dark brown diagonal stripe on both sides of having the following combination of characters: the flanks. Five adult females (MTD 46888, MUSM (1) skin on dorsum shagreen, skin on venter areolate; 24557, 27608‑09, MVZ 258411) and one male discoidal fold absent, thoracic fold present, weakly (MUSM 27607) have a narrow tan middorsal stripe, defined; dorsolateral fold usually discontinuous; and a narrow tan diagonal stripe on posterior surfaces (2) tympanic membrane and tympanic annulus ab- of the thighs and on concealed surfaces of the shanks. sent; (3) snout rounded in dorsal and lateral views; Two females (MTD 46865, MUSM 24557) have a (4) upper eyelid without enlarged tubercles; width X‑shaped blotch on the head. Four females (MTD of upper eyelid narrower than IOD; cranial crests 46888, MUSM 27608‑09, MVZ 258411) and three absent; (5) dentigerous processes of vomers absent; males (MTD 46866, MUSM 27568, 27607) have a (6) males without vocal sac, vocal slits, and nuptial triangular-shaped blotch on the head. Two females pads; (7) Finger I shorter than Finger II; tips of digits (MUSM 24557, MVZ 258411) and two males (MTD rounded; (8) fingers with lateral fringes; (9) ulnar and 46866, MUSM 27568) have a dark brown blotch tarsal tubercles present; (10) heel without tubercles; on upper lip below the eye. Males are smaller than inner tarsal fold absent; (11) inner metatarsal tuber- females and lack vocal slits, nuptial pads, and a vo- cle ovoid, one and a half the size of outer metatarsal cal sac. The smallest juvenile is 9.3 mm SVL (MVZ tubercle; outer metatarsal tubercle rounded; super- 258412). Table 3 shows ranges and proportions of the numerary plantar tubercles indistinct; (12) toes with type series. lateral fringes; basal webbing present; Toe V slightly longer or equal to Toe III; toe tips rounded, about as Distribution – Bryophryne hanssaueri has only been large as those on fingers; (13) in life, dorsum pale collected from the type locality (Fig. 4) at elevations gray, grayish brown or tan, throat and chest pale between 3266 and 3430 m along the Ericsson trail gray and tan mottled, belly black with white flecks; that connects Acjanaco to Pillahuata in Manu Nation- (14) SVL in adult females 12.2‑24.4 (n = 3), in males al Park and near Abra Acjanaco along the Paucartam- 14.8‑17.6 mm (n = 3). bo-Pilcopata road. Bryophryne zonalis differs from the other current- Etymology – The species is dedicated to Hans Sauer, ly known five species (including the ones described Germany, for supporting the BIOPAT initiative. herein) in having females that have a black belly with white flecks, a mottled pale gray and tan throat and chest, and partly reddish fingers and toes. Bryophryne Bryophryne zonalis sp. nov. (Figs. 7 and 8) zonalis (maximum SVL 24.4 mm) is much smaller than B. cophites (maximum SVL 29.3 mm, Lynch, Holotype – MUSM 27570. Adult female, collected 1975), larger than B. bustamantei (maximum SVL at Kusillochayoc (13°34’38.7”S, 71°00’08.6”W) at 23.4 mm, Chaparro et al., 2007), B. gymnotis (maxi- 3129 m elevation, Distrito de Marcapata, Provincia mum SVL 22.2 mm), B. nubilosus (maximum SVL de Quispicanchis, Región Cusco, Peru, on 4 March 21.9 mm), and about equal in SVL to B. hanssaueri 2008 by A. Catenazzi, I. Chinipa, J. C. Jahuanchi, and (maximum SVL 24.6 mm). Table 1 compares select- A. Machaca. ed characters among species of Bryophryne. 134 Three new species of Bryophryne

Table 3. Measurements (in mm) and proportions of Bryophryne hanssaueri and B. zonalis; ranges followed by means and one standard deviation in parentheses.

B. hanssaueri B. zonalis Characters Females (n = 12) Males (n = 4) Females (n = 3) Males (n = 3) SVL 18.1‑24.6 (21.7 ± 1.8) 12.3‑18.0 (15.5 ± 2.8) 22.2‑24.4 (23.3 ± 1.1) 14.8‑17.6 (16.0 ± 1.4) TL 6.5‑8.1 (7.5 ± 0.5) 4.7‑5.8 (5.4 ± 0.5) 8.7‑9.5 (9.0 ± 0.4) 6.5‑7.6 (7.1 ± 0.6) FL 7.2‑9.5 (8.7 ± 0.8) 5.2‑6.4 (5.9 ± 0.5) 9.6‑9.9 (9.7 ± 0.2) 6.9‑8.9 (7.8 ± 1.0) HL 6.9‑8.4 (7.7 ± 0.5) 5.2‑6.5 (5.9 ± 0.6) 7.9‑8.3 (8.0 ± 0.2) 5.5‑6.8 (6.2 ± 0.7) HW 6.9‑9.0 (8.0 ± 0.7) 4.5‑5.9 (5.3 ± 0.7) 7.5‑8.0 (7.8 ± 0.3) 5.2‑6.7 (5.9 ± 0.8) ED 1.8‑2.6 (2.3 ± 0.2) 1.5‑1.9 (1.8 ± 0.2) 2.2‑2.3 (2.3 ± 0.1) 1.6‑1.8 (1.7 ± 0.1) IOD 2.2‑3.1 (2.6 ± 0.2) 1.9‑2.1 (2.0 ± 0.1) 2.7‑2.8 (2.8 ± 0.1) 2.3‑2.4 (2.3 ± 0.1) EW 1.7‑2.0 (1.8 ± 0.1) 1.1‑1.6 (1.3 ± 0.3) 2.1 1.2‑1.4 (1.3 ± 0.1) IND 1.6‑2.2 (1.9 ± 0.2) 1.4‑1.6 (1.5 ± 0.1) 1.7‑2.2 (1.9 ± 0.3) 1.3‑1.7 (1.5 ± 0.2) E‑N 1.5‑2.2 (1.9 ± 0.2) 1.3‑1.8 (1.5 ± 0.2) 1.8‑2.0 (1.9 ± 0.1) 1.4‑1.8 (1.7 ± 0.2) TL/SVL 0.31‑0.39 0.32‑0.39 0.38‑0.40 0.43‑0.45 FL/SVL 0.38‑0.44 0.33‑0.46 0.39‑0.45 0.47‑0.51 HL/SVL 0.32‑0.40 0.35‑0.42 0.32‑0.37 0.37‑0.39 HW/SVL 0.34‑0.40 0.32‑0.37 0.32‑0.35 0.35‑0.38 HW/HL 0.96‑1.15 0.87‑0.97 0.94‑1.01 0.94‑0.99 E‑N/ED 0.73‑0.95 0.74‑0.95 0.82‑0.87 0.88‑1.06 EW/IOD 0.58‑0.77 0.52‑0.84 0.75‑0.78 0.52‑0.61

Description of holotype – Head narrower than body, Hind limbs short, slender, TL 38.9% of SVL; about as long as wide; HW32.8% of SVL; HL 32.4% FL 39.3% of SVL; upper surface of hind limbs sha- of SVL; snout short, acutely rounded in dorsal and lat- green with small, scattered tubercles; posterior and eral views (Figs. 8A, B), ED larger than E‑N; nostrils ventral surfaces of thighs areolate; heel without en- slightly protuberant, directed dorsolaterally; canthus larged tubercles; outer surface of tarsus with minute rostralis slightly curved in dorsal view, rounded in tubercles; metatarsal tubercles elevated, slightly profile; loreal region slightly concave; lips rounded; conical in lateral view, inner metatarsal tubercle EW without enlarged tubercles; EW narrower than elongate, about twice the size of rounded outer met- IOD (EW 75.0% of IOD); supratympanic fold short, atarsal tubercle; plantar supernumerary tubercles narrow; tympanic membrane and tympanic annulus indistinct; subarticular tubercles most prominent on absent, postrictal tubercles coalescing in two short, base of toes, low, ovoid in dorsal view, rounded in prominent ridges on each side of head. Choanae lateral view; toes with lateral fringes; basal web- small, ovoid, not concealed by palatal shelf of max- bing present; toe tips rounded, slightly pointed due illa; dentigerous processes of vomers absent; tongue to preservation, lacking marginal grooves, about 2.5x as long as wide, not notched posteriorly, poste- as large as those on fingers; relative lengths of rior one third free. toes: 1 < 2 < 3 < 5 < 4; Toe V longer than Toe III (Fig. 8D). Skin on dorsum shagreen, narrow, slightly elevated dorsolateral fold extending from posterior margin of Measurements of holotype (in mm) – SVL 24.4; TL upper eyelid to sacral region, posteriorly discontinuous; 9.5; FL 9.6; HL 7.9; HW 8.0; ED 2.3; IOD 2.8; EW skin on flanks slightly tuberculate; venter areolate; dis- 2.1; IND 2.2; E‑N 2.0. coidal fold absent, thoracic fold present; cloacal sheath short; large tubercles absent in cloacal region. Outer Color of holotype in life (Fig. 7) – Dorsum pale cin- surface of forearm with minute tubercles; palmar tu- namon with small dark brown flecks; canthal and su- bercles low, outer palmar ovoid, approximately 2x the pratympanic stripes ill defined, gray; margin of upper size of elongate, inner palmar tubercle; few supernu- and lower lips dark gray; throat mottled cream and merary tubercles, about one third size of subarticular pale gray; chest, ventral surfaces of arms and thighs tubercles; subarticular tubercles prominent, ovoid in pale gray with purpleish gray mottling; belly, groin, dorsal view, rounded in lateral view, most prominent concealed surfaces of shanks, and dorsal surface of on base of fingers; fingers with lateral fringes; Finger I tarsus dark gray with white flecks; fingers and toes shorter than Finger II; tips of digits rounded (Fig. 8C). partly reddish brown, otherwise dark gray; iris gold Lehr, E. and Catenazzi, A. 135 with fine black reticulations and a horizontal reddish 27574) has the dorsum mottled dark grayish brown brown streak through pupil. and reddish brown, postrictal ridges cream, and ven- ter mottled dark grayish brown, pale gray, and reddish Color of holotype in preservative – Dorsum gray with brown. Another male (MTD 46870) has the dorsum small dark gray flecks; throat and chest mottled cream mottled pale gray and reddish brown, the flanks pale and gray; ventral surfaces of hands and feet mottled gray, and the venter mottled blueish gray and red- cream and dark gray; iris mottled dark and pale gray; dish brown. One juvenile (MUSM 27575) has a tan otherwise as described below. dorsal band, dark gray dorsolateral folds, grayish tan flanks, and the venter mottled blueish gray and red- Variation – The dorsal color in females is more uni- dish brown. Another juvenile (MUSM 27861) has the form than in males, varying from pale cinnamon dorsum and flanks mottled pale reddish brown and to pale grayish tan to pale gray. One male (MUSM pale gray with dark brown blotches; dark brown bars on extremities; dark brown canthal and supratympan- ic stripes, a dark brown labial bar below eye, and tan postrictal ridges. Only females have a dark gray to black belly with white flecks. Females are larger than males and have more continuous, narrower, and less elevated dorsolateral folds. Males have more tuber- culate flanks than females. The smallest juvenile is 11.5 mm SVL (MUSM 27861). Table 3 shows ranges and proportions of the type series.

Distribution – Bryophryne zonalis is only known from the upper Marcapata valley, at elevations be- tween 3129 and 3285 m along the road from Abra Huallahualla to Quincemil (Fig. 4).

Etymology – The specific name zonalis is derived from the Latin noun “zona” meaning girdle or belt. The name refers to the contrasting pattern consisting of black with white flecks on the belly in females.

Ecological Results

All specimens of Bryophryne gymnotis occurred along a transect of ~400 x 2 m. Four people actively searched this transect for ground-dwelling anurans for 3 h. Frogs were found in damp soil under logs, moss, and rocks and in leaf litter. In some cases, up to three frogs were found under the same log. This species seems to have a patchy distribution, because no frogs were found in the first km of walking along a trail from San Luis to the type locality, followed by a high density of frogs for 400 m, and a subsequent drop in density above 3350 m. We found no sympat- ric anurans during our visit and during a nocturnal survey in San Luis, but Chaparro et al. (2007) report- ed Nannophryne corynetes, Gastrotheca excubitor, Pleurodema marmoratum, and Pristimantis rhabdol‑ Figure 7. Holotype of Bryophryne zonalis in life (MUSM 27570, female) in lateral (A), dorsal (B), and ventral (C) views. Photos aemus sympatric with B. bustamantei. In our survey by A. Catenazzi. at the nearby locality of Alfamayo (2490 m), we 136 Three new species of Bryophryne observed P. rhabdolaemus and Gastrotheca ochoai, B. cophites, B. nubilosus, usurpa‑ whereas at Abra de Málaga we found B. bustamantei tor, Gastrotheca excubitor, Gastrotheca marsupiata, and G. excubitor. Telmatobius timens and an undescribed species of Bryophryne hanssaueri inhabits leaf litter at the Telmatobius. transition of montane forest with the high-Andean Specimens of Bryophryne zonalis were found un- grassland or puna. Cano et al. (1995) discuss veg- der rocks and moss in fragments of elfin and montane etation of the area and Lehr and Catenazzi (2008) cloud forests. Much of the landscape of the upper list the most common plant species. The elevational Marcapata valley has been modified through defores- range of this species and relative density based on tation, cattle grazing, and cultivation; remnants of the 10 x 10 m² leaf litter quadrats has been illustrated by original forest habitats can be found along streams Lehr and Catenazzi (2009, Bryophryne sp. in Fig. 4). and in areas of difficult access. In addition to severe Bryophryne hanssaueri seems to be a rare species and habitat disturbance, the pathogenic fungus Batra‑ was not discovered until the rainy season of 2008, chochytrium dendrobatidis occurs this area (Seimon despite intensive sampling between 1996 and 1998 et al., 2006) and has been detected (pers. comm. A. and in the dry season of 2007. However, we found 12 Catenazzi) in one of the types of B. zonalis (MUSM individuals and a clutch in a single 10 x10 m2 quad- 27861). rat plot at 3415 m in the rainy season of 2009, sug- gesting that this species can be locally abundant. The clutch contained 19 eggs ranging between 4.2 and Discussion 4.8 mm in diameter and that were close to hatching. We found the clutch unattended under mosses, and The genus Bryophryne contains six species re- moved it to the field station until hatching occurred. stricted to the Cusco Region in southern Peru. Previ- Hatchlings measured 4.9‑5.2 mm in snout-vent length ously, one of the defining characters for Bryophryne (n = 8). The elevational range overlaps with that of was the absence of a tympanum, which was

Figure 8. Dorsal (A) and lateral (B) views of head and ventral views of right hand (C) and left foot (D) of Bryophryne zonalis (MUSM 27570). Drawings by E. Lehr. Lehr, E. and Catenazzi, A. 137 considered a synapomorphy. Bryophryne gymnotis is a ‘short whistle’ produced during the day in April, the first species of the genus with a tympanic������������� mem- but they did not provide the spectrogram and oscil- brane and a tympanic annulus, thus altering the��������� defi- logram of the call. The description of three new spe- nition of the genus provided by Hedges et al. (2008), cies of Bryophryne and the high endemism of Ande- as follows: …(2) �������������������������������tympanic membrane, tympanic an- an frogs (Duellman, 1999) suggest that many more nulus, columella, and cavum tympanicum are present Bryophryne species remain to be discovered from or absent. Even though the phylogenetic relationships other watersheds in southern Peru. among species of Bryophryne are unknown, we are confident of the generic assignment for B. gymnotis because of the overall similarity with other members Resúmen of the genus. We recognize a convergence in the state of the tympanum in Bryophryne and Phrynopus. Both Describimos tres nuevas especies de Bryophryne genera have high local species diversity and occur at de la Región de Cusco en el sur de Perú, incrementando high elevations, and the majority of the species lack a seis el numero de especies conocidas en el género. a tympanum. Una de las nuevas especies se diferencia de todos Bryophryne hanssaueri is sympatric with both los demás Bryophryne por la presencia de annulus y B. nubilosus and B. cophites in the upper Cosñipata membrana timpánicos. Los machos de esta especie valley, Manu National Park. This locality has been tienen un saco y aberturas vocales y producen intensively sampled over the past 12 years (Lehr and vocalizaciones que describimos en este artículo. Catenazzi, 2008, 2009; Catenazzi and Rodriguez, Esta especie ocurre en San Luis, un bosque nublado 2001). In the upper Umasbamba valley (near Abra de a elevaciones entre 3272 y 3354 m en la carretera Málaga), 80 km west of the upper Cosñipata valley, entre el Abra Málaga y Quillabamba. La segunda we found B. gymnotis approx. 5 km from the closest especie se caracteriza por tener garganta e ingles collecting site of B. bustamantei. These two species anaranjadas y ha sido encontrada en la parte alta del might be sympatric, but further field research is need- Parque Nacional del Manu (elevación 3266‑3430 m), ed to establish the elevational ranges of both species. en la trocha Ericsson entre Acjanaco y Pillahuata y a In the case of B. zonalis collected from the Marca- lo largo de la carretera Paucartambo-Pilcopata cerca pata valley (80 km southeast of the upper Cosñipata de Abra Acjanaco. La tercera especie tiene garganta valley), a currently undescribed species (identified as y pecho gris claros y gris oscuro moteado, mientras B. cophites in the KU catalogue, KU196596‑7, herein que el vientre es negro con manchas blancas. Este referred to as Bryophryne sp. A) might be sympatric Bryophryne se conoce únicamente de la parte alta del with B. zonalis. Our own collections of this unde- Valle de Marcapata a lo largo de la carretera entre Abra scribed species are limited to high elevations in the Huallahualla y Quincemil a elevaciones entre 3129 y high-Andean grassland, whereas B. zonalis was col- 3285 m. Incluimos observaciones sobre la ecología lected in fragments of montane cloud forest and elfin de cada especie y un mapa con las localidades tipo de forest. todas las especies conocidas de Bryophryne. In the three valleys (Umasbamba, Cosñipata, and Marcapata), only one species of Bryophryne seems to inhabit the high-Andean grassland: B. bustamantei Acknowledgments in Umasbamba, B. cophites in Cosñipata, and B. sp. A in Marcapata. One to two additional species live We thank an anonymous reviewer for his helpful comments in the elfin and montane forests immediately adja- and suggestions. Specimens were loaned by J. Córdova (MUSM). cent to the grasslands, usually around 3000‑3400 m AC was assisted in the field by J. Carrillo, D. Cruz, I. Chinipa, but down to 2300 m in the case of B. nubilosus in J. C. Jahuanchi, E. Luna, A. Machaca, and W. Qertehuari. AC thanks the staff of the Amazon Conservation Association in Cosñipata: B. gymnotis in Umasbamba, B. hanssau‑ Cusco and Wayqecha for field and logistic support; the National eri and B. nubilosus in Cosñipata, and B. zonalis in Institute for Natural Resources in Lima for research, collecting Marcapata. Interestingly, both species (B. bustaman‑ and export permits; the administration of Manu National Park in tei, B. gymnotis) from Umasbamba produce vocal- Cusco for research permit and logistic support. Fieldwork was izations and males have vocal slits and vocal sacs, funded by grants from the Amazon Conservation Association, the Chicago Board of Trade Endangered Species Fund, the Rufford whereas none of the Bryophryne species in the other Small Grants Foundation and the Fondation Matthey-Dupraz to watersheds is known to produce any call. Chaparro AC and by an Specialist Group Award to AC and et al. (2007) described the call of B. bustamantei as EL. AC was supported by a Post-doctoral fellowship from the 138 Three new species of Bryophryne

Swiss National Science Foundation (116305). EL thanks the Duellman, W. E., E. Lehr, and P. Venegas. 2006. Two new Humboldt-Foundation for an alumni grant to conduct research at species of Eleutherodactylus (Anura: Leptodactylidae) from the Natural History Museum and Research Center, northern Peru. Zootaxa, 1285:51‑64. The University of Kansas, USA. Hedges, S. B., W. E. Duellman, and M. P. Heinicke. 2008. New World direct developing frogs (Anura: Terrarana): molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737:1‑182. iterature ited L C Lehr, E. and A. Catenazzi. 2008. A new species of Bryophryne (Amphibia: Strabomantidae) from southern Peru. Zootaxa, Cano, A., K. R. Young, B. León, and R. B. Foster. 1995. 1784:1‑10. Composition and diversity of flowering plants in the upper Lehr, E. and A. Catenazzi. 2009. A new species of minute montane forest of Manu National Park, southern Peru; (Anura: Strabomantidae) from southern Peru: the pp. 271‑280. In: Churchill, S. P., H., Balslev, E., Forero, smallest of the Andes. Copeia, 2009:148‑156. and J. L. Luteyn (Eds.). Biodiversity and conservation of Lynch, J. D. 1975. A review of the Andean leptodactylid frog neotropical montane forests. New York Botanical Garden, genus Phrynopus. Occasional Papers of the Museum of New York. Natural History, The University Kansas, 35:1‑51. Catenazzi, A. and L. Rodríguez. 2001. Diversidad, distribución Lynch, J. D. and W. E. Duellman. 1997. Frogs of the genus y abundancia de anuros en la parte alta de la Reserva de Eleutherodactylus in western Ecuador. Systematics, ecology, Biosfera del Manu; pp. 53‑57. In: Rodriguez, L. (Ed.). El and biogeography. The University of Kansas Special Manu y otras experiencias de investigación y manejo de Publication, 23:1‑236. bosques neotropicales. Pro-Manu, Cusco, Peru. Seimon, T. A., A. Seimon, P. Daszak, S. R. P. Halloy, L. M. Chaparro, J. C., I. De la Riva, J. M. Padial, J. A. Ochoa, and E. Schloegel, C. A. Aguilar, P. Sowell, A. D. Hyatt, B. Lehr. 2007. A new species of Phrynopus from Departamento Konecky, and J. E. Simmons. 2006. Upward range extension of Cusco, southern Peru (Anura: Brachycephalidae). Zootaxa, Andean anurans and chytridiomycosis to extreme elevations 1618:61‑68. in response to tropical deglaciation. Global Change Biology, Duellman, W. E. 1999. Distribution patterns of 12:1‑12. in South America; pp. 255‑328. In: Duellman, W. E. (Ed.). Patterns of distribution of amphibians: a global perspective. Submitted 13 January 2009 The Johns Hopkins University Press, Baltimore, London. Accepted 09 April 2009

Appendix I

Bryophryne bustamantei – PERU: Cusco: Provincia La Convención: Abra de Málaga: MUSM 24537‑38.

Bryophryne cophites – PERU: Cusco: Provincia de Paucartambo: Distrito Cosñipata: S slope Abra Acanaco, 14 km NNE Paucartambo, 3400 m: KU 138884 (holotype); N slope Abra Acanaco, 27 km NNE Paucartambo, 3450 m: KU 138885‑908, 138911‑5 (all paratypes); 2 km NE of Abra Acanaco, 13°11.367’S, 71°36.223’W, 3280 m: MHNG 2698.24, Tres Cruces, 8.5 km N of Abra Acanaco, 13°07’18.3”S, 71°36’37.8”, 3590 m: MUSM 26283, 26267.

Bryophryne hanssaueri – PERU: Cusco: Provincia de Paucartambo: Distrito Cosñipata: Acjanaco, Manu Na- tional Park: 1 km south of the type locality, 13°11’30.0”S, 71°37’16.1”W, 3355 m: MUSM 27940; 900 m south of the type locality, 13°11’27.4”S, 71°37’15.2”W, 3390 m: MUSM 27941‑2; 1.9 km NW of the type locality, 13°10’07.9”S, 71°37’49.2”W, 3414 m: MUSM 27943‑54.

Bryophryne nubilosus – PERU: Cusco: Provincia de Paucartambo: Distrito de Cosñipata, 500 m NE of Esper- anza, 2712 m: MUSM 26310 (holotype), MUSM 26311; near the type locality, 13°11’33.21”S, 71°35’25.17”W, 3065 m: MTD 47294; near Hito Pillahuata, 2600 m: MUSM 20970; Quebrada Toqoruyoc, 3097 m: MUSM 26312, MTD 47293; Esperanza, 2800 m: MHNSM 26316‑17; 13°11’20.2”S, 71°35’07.3”W, 2900 m: MUSM 24539‑40.

Bryophryne sp. A – PERU: Cusco: Provincia de Quispicanchis: Distrito de Marcapata: Coline, 3672 m: MUSM 27571, 27573.