N.Orntates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

AMERICAN MUSEUM N.orntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3068, 15 pp., 12 figures, 1 table June 11, 1993 A New Poison Frog from Manu INational Park, Southeastern Peru (Dendrobatidae, Epipedobates) LILY RODRIGUEZ' AND CHARLES W. MYERS2 ABSTRACT Epipedobates macero is a new species of den- ilar to a few other species occurring along the An- drobatid poison frog from lowland rain forest of dean front in eastern Peru, namely E. petersi and the Manu National Park, in the upper Madre de E. cainarachi, which differ in details ofcoloration, Dios drainage ofsoutheastern Peru. It is most sim- morphology, and vocalization. RESUMEN Epipedobates macero, especie nueva, es un den- del llano amaz6nico al pie de los Andes orientales drobatido venenoso de la selva pluvial baja del peruanos, a saber, E. petersi y E. cainarachi, las Parque Nacional del Manu, en el drenaje del Rio cuales difieren en detalles de coloracion, morfo- Alto Madre de Dios, al sudeste del Peru. Es similar logla, y vocalizacion. a otras dos especies que ocurren en los bosques ' Field Associate, Department of Herpetology and Ichthyology, American Museum of Natural History. Investi- gadora Asociada: Asociaci6n Peruana para la Conservaci6n de la Naturaleza (APECO), Parque Jos6 de Acosta 187, Lima 17, Perfi; and Museo de Historia Natural de la Universidad Mayor de San Marcos, apartado 140434, Lima 14, Peru. 2 Curator, Department of Herpetology and Ichthyology, American Museum of Natural History. Copyright © American Museum of Natural History 1993 ISSN 0003-0082 / Price $3.90 2 AMERICAN MUSEUM NOVITATES NO. 3068 INTRODUCTION The western edge of Amazonia, at low to Epipedobates macero, new species moderate elevations along the Andes, may Figures 1-9 have been an important region of speciation or refuge3 for dendrobatid frogs of the genus HOLOTYPE: MHNSM 0726 (formerly Epipedobates. Eleven of 24 species recently MHNJP 2001), an adult female collected on assigned to this genus (Myers, 1987; Schulte, October 21,1985, by Lily Rodriguez, on west 1989; Jungfer, 1989) have fairly restricted side Rio Manu across from Cocha Cashu Bi- distributions within that segment of the An- ological Station,5 Parque Nacional del Manu, dean front between parallels 20 North (south- about 380 m elev., Department of Madre de ern Colombia) and 160 South (northern Bo- Dios, Peru (fig. 2). livia). With a few exceptions, the other species PARATYPES: A total of 17 adults -plus ju- are either more widespread in the Amazonian veniles and larvae -all from Parque Nacion- basin or else occur on the Pacific versant of al del Manu in Depto. Madre de Dios, as Colombia and Ecuador.4 follows: AMNH A-129473, 129474 (juv.), A twelfth species of Epipedobates (fig. 1) 129475 (4 dorsal tadpoles from 129473), endemic to extreme western Amazonia is 133205, 133206 (6 froglets), 133207 (2 free- named here: It came to the attention of the living tadpoles); 134159-134163; MHNSM first author in 1985 during a preliminary fau- 3834 (skinned carcass), 3837, 3838, 3841 nal survey of Manu National Park, in the (skinned carcass), 9093; MHNNP (Paris) upper Madre de Dios drainage of southeast- 1989.3605-1989.3607; USNM 322909, ern Peru. It has since been listed by Rodri- 322910 (juv.) -preceding all collected at the guez (1987; see also Rodriguez and Cadle, type locality across river from Cocha Cashu 1990) as a new species being described by Biological Station, 1986-1989, by Lily Rod- Rodriguez and Myers. Schulte (1987) and riguez et al. AMNH A-134088, near Tay- Jungfer (1989) also mentioned its existence akome (Machiguenga Indian village), 450 m, on the basis of a photograph taken by one of 11053'S, 71025'W, collected on August 9, the Park's early visitors. 1986, by Andre Bairtschi. ETYMOLOGY: The species name macero, a noun in apposition, is the Machiguenga In- dian word for "frog." The Machiguenga are one ofthe indigenous tribes ofthe Rio Manu 3These two concepts of course are not mutually ex- region. clusive, inasmuch as postulated refugia may be theorized DEFINITION AND DIAGNOSIS: A medium- as centers ofspeciation. But virtually every part ofmain- an snout-to-vent land tropical South America has been designated a Pleis- size dendrobatid with adult tocene refuge for one kind of organism or another, usu- length (SVL) of about 24-30 mm. Head and ally without supporting data other than current back bright red and strongly granular; sides distributions. Distribution maps alone cannot differen- black, with a bright yellow stripe extending tiate between recent speciation, impending extinction, obliquely from groin to above shoulder; yel- or a range of other possibilities. low labial stripe not reaching snout; a yellow 4The genus Epipedobates Myers was erected explicitly spot on anterodorsal base of thigh; ventral to accommodate the basal, plesiomorphic assembage of surfaces blue with black reticulum. Lacking lipophilic-alkaloid producing dendrobatids. These spe- axillary and groin spots and lacking a con- cies can be arranged in species groups (although only one cealed calf spot on proximoventral part of is demonstrably monophyletic), but relationships among the groups are unknown. As recommended by Myers (1987: 303), "further splitting is inadvisable unless two 5 Stephens and Traylor (1983: 57) placed Cocha Cashu or more separate lineages can be objectively [i.e., phy- at approximately 115 1'S, 711l9'W, at an elevation of logenetically] determined." For that reason (with further about 400 m above sea level. Terborgh et al. (1984) comments by Myers et al., 1991: 18), we follow Schulte stated that Cocha Cashu is "located about 45 km north- (1989) in considering, for the time being, the 1988 names west (= 80 river km upstream) from the mouth of the Allobates Zimmermann and Zimmermann and Phobob- Rio Manu . 1 1°55'S, 77°[error for 71°]18'W; elev. ap- ates idem as junior synonyms of Epipedobates. prox. 380 m." 1993 RODRIGUEZ AND MYERS: POISON FROG 3 Exiarachi .........E.0mtmaceroorak 0 200 400 600k,..0 Fig. 2. Western Amazonia, showing distributions of the species of Epipedobates discussed in this paper. Distribution of E. petersi-in the Rio Ucayali and part of the Rio Huallaga drainages- is based mainly on the type specimens mapped and listed in Silverstone (1976: 19, 37). The ques- tioned locality is for petersi paratypes from La Union in the Rio Tambobata drainage (Madre de Dios drainage), Depto. Puno, extreme southeastern Peru. See footnote 10. Fig. 1. Epipedobates macero, new species. Lat- eral and ventral views of a living specimen pho- tographed on September 8, 1986, at Tayakome, Parque Nacional del Manu. [Courtesy Andre Bart- E. petersi (fig. 12), but E. macero differs from schi] these in that the oblique lateral stripe and the labial stripe are both incomplete. In E. macero, the anterior terminus ofthe oblique lat- shank. Teeth present on maxillary arch. Ap- eral stripe lies above or slightly anterior to pressed first finger about equal to second; fin- the shoulder (conspicuously short of the eye) ger discs weakly to moderately expanded; and the labial stripe usually does not extend third finger disc ofadults about 1.3-1.7 times rostrad from the eye (conspicuously short of wider than finger. Toes completely without the narial region). Also, there are differences webbing. Advertisement call a train of short, in color in life: The central Peruvian E. pe- harsh notes at a dominant frequency of 3500 tersi has a brown dorsum, a pale (yellowish, Hz; terrestrial. Defensive skin alkaloids in- perhaps with greenish tinge) oblique lateral clude a pyrrolizidine oxime and members of stripe continuous to the eye, and a pale (white the decahydroquinoline and histrionicotoxin or yellowish) labial stripe extending anteri- classes of dendrobatid alkaloids. orly to the end of the snout. Epipedobates Epipedobates macero is a member of the macero is more similar to the geographically E. petersi species group (sensu Schulte, 1989). distant E. cainarachi (NW Peru, fig. 2) in that It is similar in morphology and basic color both have red dorsa and yellow stripes, but pattern (fig. 1) to E. cainarachi (fig. 11) and E. cainarachi has a slightly longer first finger, 4 AMERICAN MUSEUM NOVITATES NO. 3068 Fig. 3. Epipedobates macero, new species. Dorsal and ventral views ofholotype (MHNSM 07262), x 2. its coloration seems less bright and/or more longest (third) finger 7.6; width ofdisc ofthird variable, and its stripes are complete as in E. finger (and width of penultimate phalanx be- petersi. See Notes on Other Species for fur- low disc) 1.0 (0.6); width of discs (and pen- ther comment and apparent differences in vo- ultimate phalanges below discs) of third and calizations. fourth toes 1.0 (0.6) and 1.2 (0.7), respec- MEFASupREmENTS OF HOLOTnPE (in mm): The tively. undissected holotype (fig. 3) is a femalejudged to be sexually mature because of its size; it has toes 2-3 aberrantly fused on the left foot DESCRIPTION OF TYPE SERIES but appears normal in all other respects. External Morphology: A medium-size den- Length from snout to vent 27.3; tibia length drobatid, with adult males attaining a size of between heel and outer surface offlexed knee about 27 mm SVL and adult females about 14.8; greatest width of body 8.4; head width 30 mm SVL (measurements and proportions between angles of jaws, and between outer summarized in table 1). Dorsal skin of head, edges upper eyelids, 8.5, 7.6 respectively; ap- body, and hind limbs coarsely and conspic- proximate width ofinterorbital area 3.0; head uously granular; skin smooth on forelimbs, length (sagittal) from tip of snout to angle of sides ofhead and body, and ventral surfaces.

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