LIBRARY OCCASIONAL PAPERS of the MUSEUM OF NATURAL HISTORY The University of Kansas Lawrence, Kansas number 155, pages 1-33 11 march 1993 New Species of Centrolenid Frogs from Northern Peru William E. Duellman and Rainer Schulte Museum ofNaturalHistory andDepartment ofSystematica andEeologx, The University ofKansas, Lawrence, Kansas 66045-2454, USA (WED); Proyecto de Investigation de laBiologia de las Cordilleras Orientates,Apartado258, Tarapoto, Peru (RS) ABSTRACT Eightnewspeciesofcentrolenidfrogsaredescribedfromtheeastern slopesoftheCordilleraCentral and ridgesextendingeastwardandsouthwardfrom the main cordillera of the Andes in Departamento San Martin in northern Peru. Three ofthese species are described in thegenus Centrolene;they havedark dorsal pigmentation in preservative, and the males have humeral spines. Four speciesareplaced inthegenusCochranella;theyhavedarkdorsalpigmentationin preservative, but males lack humeral spines. One ofthe species ofCochranella is known only from females. One species is white in preservative and is amemberof the genusHyalinobatrachium. Three species recognized in the genus Cochranella by Ruiz-Carranza and Lynch ( 1991) are placed in the genus Centrolene. Key words: Anura, Centrolenidae, New species, Peru. RESUMEN Ocho especies nuevas de ranas centrolenidas se describen de las laderasorientalesdelaCordilleraCentralydelasmontanasqueseextiendenaleste yalsurdelacordilleraprincipaldelosAndesenel DepartamentodeSanMartinen elnortedePeru. TresdeestasespeciessedescribenenelgeneroCentrolene:tienen pigmentacion oscura en el dorso en preservative, y los machos tienen espinas humerales. Tres especies se ponen en el genero Cochranella; tienen pigmentacion oscura en el dorso en preservativo, pero los machos no tienen espinas humerales. SololashembrasseconocendeunadelasespeciesdeCochranella. Unaespeciees blanca en preservativo y es un miembro del genero Hyalinobatrachium. Tres especiesreconcidasenelgeneroCochranellaporRufz-CarranzaandLynch(1991) se ponen en el genero Centrolene. Palabras claves: Anura, Centrolenidae, Especies nuevas, Peru. ©MuseumofNaturalHistory.TheUniversityofKansas.Lawrence. 1SSN:0091-7958 2 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155 In recent years, six species of frogs have been named in the genus Centrolenella from the cloud forests on the eastern slopes ofthe Ande—s or isolated mountain ranges in central and southern Amazonian Peru C. spiculata and C. truebae from DepartamentoAyacucho (Duellman, 1976), C. mariae from Departamento Huanuco (Duellman and Toft, 1979), C. phenax and C. pluvialis from Departamento Cuzco (Cannatella and Duellman, 1982), and C. azulae from Departamento Huanuco (Flores and McDiarmid, 1989). Even with the descriptions of these new species (all except the holotype of C. azulae were collected for the first time in the 1970s), it seemed reasonable that many additional species would be discovered as new areas were explored, especially in northern Peru. This supposition wasverified with thediscovery ofC. euhystrixand C. hesperia on the Pacific versant of the Andes in northern Peru in 1987 (Cadle and McDiarmid, 1990) and by our discovery of eight undescribed species on the easternAndean slopes in Departamento San Martin in northern Peru in 1989. In the Departamento San Martin, a complex series of ridges reaching elevations of more than 1000 m extend eastward and southward from the Cordillera Central of the Andes. These steep, rocky ridges support lower montane rainforest and cloud forest, whereas the intervening valleys, most of which are extensively cultivated, contain remnants of dry tropical forest. Many small streams cascade down these slopes; major drainage systems flow south (e.g., Rio Mayo) oreast (e.g., Rio Cainarache) into the Rio Huallaga. which separates the Cordillera Central from the Cordillera Oriental. In northern Peru, the Cordillera Oriental terminates in a series of ridges with elevations of 1000-1200 m. The Rfo Huallaga cuts a canyon through these front ranges between Shapaja and Chazuta, to the south of which is the northern end ofthe CordilleraAzul (Fig. 1 ). Adultcentrolenidswerecollectedon theeastern slopesofthe Cordillera Central and on two associated ridges. One site is on the west slope of a pass,AbraTangarana, to the west ofthe valley ofthe Rio Mayo. The other sites are on a ridge north ofTarapoto and east ofthe Rio Mayo. METHODS AND MATERIALS Inthe followingdescriptions, species are assignedtogeneraand species groups according to the definitions provided by Rufz-Carranza and Lynch (1991), and the numbered characters in the diagnoses follow the standard established by Lynch and Duellman ( 1973) as modified by Cannatella and Duellman (1982) and Flores ( 1985). None ofthe species ofcentrolenids in Central America, Chocoan South America, Venezuela, and southeastern Brazil is sharedwiththe upperAmazon Basin andtheAmazonian slopes of theAndes. Consequently, comparisons ofthe new taxa are made only with NEWCENTROLENIDFROGS FROM NORTHERN I'ERl Fig. 1. Map of part of Departamento San Martfn, Peru, showing localities mentioned in text; circled numbers are: 1 = Abra Tangarana, 2 = Cataratas Ahuashiyacu. 3 = northern end ofCordilleraAzul. Basedon Mapa Fisico Politico Departamento de San Martin, Ed. 2, 1985, Instituto Geografico Nacional. Lima, Peru. those species from the upperAmazon Basin and the Amazonian slopes of the Andes. mm All measurements were taken to the nearest 0.1 with dial calipers. Abbreviations used in the descriptions are: ED (eye diameter) = greatest length of eye visible externally; EW (eyelid width) = greatest diagonal width of upper eyelid; FL (foot length) = distance from proximal base of inner metatarsal tubercle to tip of fourth toe; HL (head length) = distance betweentipofsnoutandposterioredgeofangleofjaws; HnL(handlength) =distancebetweenproximal baseofpalmartubercle andtipofthird finger; HW (head width) = greatest width ofhead at lateral margins ofjaws; IOD (interorbital distance) = width of top of head between nearest medial margins ofuppereyelids; SL (snout length) = distance from tip ofsnout to anterior corner of orbit; SVL (snout-vent length) = distance from tip of snout to posterior end of body; T-E (tympanum-eye) = distance between anterior border of tympanic annulus and posterior corner of eyelid; TL (tibia length) =distance from knee to heel; TYM (tympanum) = horizontal 4 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155 length of tympanum including annulus. Relative lengths of digits were determined by adpressing adjacent digits. Webbing formulae follow the scheme ofSavage and Heyer ( 1967) as modified by Myers and Duellman (1982). Nuptial pad terminology follows Flores (1985). Museum collections are abbreviated as follows: KU = Museum of Natural History,TheUniversityofKansas; LSUMZ=MuseumofZoology, Louisiana State University; MHNSM = Museo de Historia Natural, Universidad Nacional Mayorde San Marcos, Lima, Peru. Comparisons of the new taxa were made with KU specimens ofthe other species. DESCRIPTIONS OF NEW SPECIES Centrolenefernandoi new species Figure 2 — Holotype. KU 211770, an adult male, from the west slope of Abra Tangarana, 7 km (by road) northeast of San Juan de Pacaysapa (06°12'S, 76°44'W, 1080 m), Provincia Lamas, Departamento San Martin, Peru, one of a series collected on 5 February 1989 by William E. Duellman and Rainer Schulte. — Paratopotypes KU 211771-75 and MHNSM 6176-78; same date and collectors.— Diagnosis. A species in the Centrolene prosoblepon group characterized by: (1) vomerine teeth present; (2) bones green; (3) parietal peritoneum white; visceral peritoneum clear; (4) color in life, lime-green with bluish white flecks dorsally; in p—reservative, pale lavender; (5) modal w—ebbing between outer fingers III 2 2 IV; (6) modal webbing on foot I 1 1+II 1—2 III 1—2 IV 2—1 V; (7) snout rounded in dorsal view, bluntly rounded in profile: (8) dorsal skin shagreened with scattered minute spicules; (9)armsandlegslackingtuberclesanddermal folds; (10) humeral spine present; (11) tympanum distinct, oriented posterolaterally with slight dorsal inclination; (12) prepollex enlarged; unpigmented nuptial excrescences(TypeI)present; prepollical spineabsent; (13)pairofenlarged tubercles below vent; (14) first finger longer than second. Of the other Centrolene in the region with vomerine teeth, C. audax differs from C.fernandoi (characters in parentheses) by having the snout truncate in profile (bluntly rounded), skin on dorsum shagreened without spicules (shagreened with scattered small spicules), and in life, golden (bluish-white) flecks on dorsum, digits pale yellow (pale green), and iris pale bronze (silvery-green) with black reticulations. Centrolene marine differs by having large, pale spots on the dorsum, truncate snout in profile, and only basal webbing between Fingers III and IV; C. azulae and C. puyoensisdifferbyhaving ulnarfolds, snouttruncate inprofile inC.azulae NEWCENTROLENID FROGS FROM NORTHERN PERU — KU Fig. 2. Upper left para—type ofCochranella croceopodes, 211804, female, 24.7mm SVL. Upperright. —holotypeofCentrolenefernandoi,KU211770,male, 24.0 mm SVL. Middle left holot—ype of Centrolene lemniscatum, KU 217300, male. 27.0 mm SVL. Middle right hol—otype of Hyalinobatrachium lemur, KU 211768, male, 20.4mm SVL. Lowerleft holotype o—fCochranella saxiscandens, KU 211779, male, 21.7 mm SVL. Lower right holotype of Cochranella tangarana, KU 211776, male, 23.3 mm SVL. 6 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155 and truncate in dorsal view in C. puyoensis, and less extensive webbing between Fingers—III and IV in C. puyoensis. Description. Nine adult males; SVL 22.5-26.4 mm. Body slender, elongate. Head distinct, about as wide as long, round in outline as viewed from below; HW 31.3-38.1% (3c = 33.2) of SVL; HL 30.0-36.0% (3c = 33.7) of SVL; snout rounded in dorsal view, bluntly rounded with slight anteroventral inclination in profile: SL 34.2^11.0% (x = 37.3) of HL; canthusrostralisstraight, indistinct,round insection; lorealregionconcave; lips barely flared anterior to orbit (Fig. 3C). Nostril ovoid, directed anteriorly on protuberance; internarial area slightly depressed. Eye large, ED 34.3^1.1% (x = 37.3) of HL, directed anterolateral^ at 37^0° (x = 39.2) from midline; greatest diagonal EW 57.1-67.0% (x = 60.0) of IOD. Tympanumdistinct,lowerfour-fifthsvisible,orientedposterolaterally with slight dorsal inclination, separated from eye by distance 1.64-1.90 (x = 1.71) times TYM, which is 31.2-41.4% (x = 35.7) of ED; tympanic annulus slightly elevated anteriorly and ventrally; supratympanic fold moderate, obscuring tympanic annulus dorsally. Skin on dorsal surfaces ofhead, body, and limbs shagreened with small spicules on side of head ventral and posterior to orbit and on dorsolateral surfaces ofbody. Flanks smooth dorsally, granular ventrally. Skin on belly and proximal four fifths of ventral surfaces of thighs coarsely granular; otherventral surfaces smooth. Ventdirectedposteroventrallyat upperlevel of thighs; cloacal sheath short, unmodified; para- and subcloacal folds absent; one pairoflarge, round, flat tubercles on ventral surfaces ofthighs below vent. Breadth of upper arm about one half that of forearm; humeral spine present; ulnarfold and tubercles absent; hand moderately large; HnL88.0- 95.2% (x = 91.0) of HL; digits broad, bearing narrow lateral fringes, including outeredge ofFinger IV but not inneredge ofFinger I; length of digits II < I < IV < III; webbing basal between Fingers I and II; webbing 2—2 2— formula for other fingers II III (l-l1/:) IV (Fig. 5C). Terminal discs elliptical: width ofdisc on Finger III about 1.3 times TYM; width of disc on Finger I 64.3-69.2% (x = 66.7) ofthat on Finger III. Subarticular tubercles as wideasdigits, round, elevated; penultimate tubercle on Finger IV largest; supernumerary tubercles present on proximal segments (absent on Fingers III and IV in some specimens); palmar tubercle large, broadly ovoid with diagonal orientation, elevated, its length 15.5-20.5% (x = 17.7) of HnL. Thenar tubercle elliptical, diffuse proximally, barely elevated, aboutas longaspalmartubercle; prepollex slightlyenlarged;Type I nuptial excrescence present; prepollical spine absent. Hind limbs long, slender; TL 54.3-59.2% (x = 56.7) ofSVL; FL43.0- 48.2% (x = 45.2) of SVL; dermal fringes, tarsal tubercles, and tarsal fold absent; inner metatarsal tubercle elongately elliptical, barely elevated, not NEWCENTROLENIDFROGS FROM NORTHERN PERU 7 visible from above; outer metatarsal tubercle absent: toes moderately slender, bearing lateral fringes, including outer edge of Toe V and inner edge ofToe I; length oftoes I < II < III < V—< IV; toes about three-fourt—hs webbed; webbing formula I 1—2 II (1-1'/:) (2"-2+) III (l-l1/:)—2 IV 2 ( 1-1'/:) V. Terminal discs subtruncate; width ofdigit on Toe I 45.5-55.5% (x = 50.3) of that on Toe IV, which is 69.2-73.3% (x = 71.3) of that on Finger III; subarticular tubercles not as wide as digits, round, elevated. Dentigerous processes of vomers ovoid, between nearly quadrangular choanae, narrowly separated medially, each bearing 2-5 (x = 3.3) teeth. Tongue nearly round, barely free posteriorly. Vocal slit elongate, extending from posterolateral margin oftongue toward angle ofjaw; vocal sac single, median, subgular. Color in life: Dorsum lime-green with small bluish-white flecks; digits pale green; parietal peritoneum white; visceral peritoneum clear; heart not visible; bones green; iris pale silvery-green with fine black reticulations. Colorinpreservative:Generalappearance lavenderwithsmall,elevated, cream flecks dorsally, cream ventrally. Under 25x magnification, dense melanophores in skin of dorsal surfaces of head and body with scattered, small, unpigmented elevations; equally dense melanophores in loreal region; less dense melanophores on upper lip, in tympanic region, and on dorsal surfaces offorelimbs. hind limbs, third and fourth fingers, tarsi, and fourth and fifth toes. Edge ofuppereyelid pigmented. Flanks, anteriorand posterior surfaces ofthighs, inner two fingers and inner three toes, and all ventral surfaces cream with no melanophores. Measurements (mm with means in parentheses): SVL 22.5-26.4 (23.7), TL 12.9-14.3 (13.4), FL 10.2-11.5 (10.7), HW 7.3-8.9 (7.9), HL 7.6-8.4 (8.0), SL 2.8-3.2 (3.0). ED 2.8-3.2 (3.0). IOD 2.8-3.1 (3.0). T-E 1.7-1.9 (1.8), TYM 1.0-1.2 (1.1), Hn—L 7.2-7.4 (7.3). Distribution and ecology. All individuals were on the uppersurfaces m of leaves of trees 1.5-2.0 above a small stream in a narrow ravine at night. Although centrolenids were calling, no call was associated with this species. The type locality is on the road between Moyobamba and Tarapoto. The stream flows southward at a point about 0.5 km above (northeast of) Somos Libres (a road maintenance camp) and about 1 km below (southwe—st of) the crest ofthe ridge. Etymology. The specific name isapatronymforFernandoM. Cuadros V.oftheUniversidadNacional Mayorde San MarcosinLimainrecognition ofhischeerfulcompanionshipanddedicationtocollectingfrogsinnorthern Peru in 1989.— Remarks. The most similar species to C. fernandoi is C. audax, which is known from elevations of 1660-1700in on theAmazonian slopes ofthe Andes in northern Ecuador. UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155 ALathRlM***' Fig. 3. Heads of centrolenids: A. Cochranella chancas, KU 211778. B. Cochranella croceopodes, KU 211804. C. Centrolenefernandoi, KU 211770. D. Centrolene lemniscatum, KU 217300. Scale = 5 mm. NEWCENTROLENIDFROGS FROM NORTHERN PERU J Fig. 4. Heads of centrolenids: A. Hyalinobatrachium lemur, KU 211768. B. Centrolene muelleri, KU 217301. C. Cochranella saxiscandens. KU 211779. D. Cochranella tangarana, KU 211777. Scale = 5 mm. 10 UNIV. KANSAS MUS. NAT. HIST. OCC. PAP. No. 155 -I L 1 | mmm m Ml » m %& ... Mm Fig. 5. Hands of centrolenids: A. Cochranella chancas, KU 211778. B. Cochranella croceopodes, KU 211804. C. Centrolenefernandoi, KU 211770. D. Centrolene lemniscatum, KU 217300. Scale = 5 mm.