Scientific Papers Natural History Museum The University of Kansas 15 November 2004 Number 35:1-49 Frogs of the Genus Colostethus (Anura; Dendrobatidae) in the Andes of Northern Peru By OBP^^^ William E. Duellman 1008 \ Natural Histon/ Mitsciiiii ami Biodi'ocrsiti/ Research Coitcr T?\7\ /^r'R&v'l ' The University of Kansas Lawrence, Kansas 66045-7561 USA CONTENTS ABSTRACT 2 RESUMEN 2 INTRODUCTION 3 Acknowledgments 3 MATERIALS AND METHODS 4 ANDES OF NORTHERN PERU 4 SUMMARY OF TAXONOMIC CHARACTERS 3 IDENTIFICATION OF SPECIES 10 Key TO THE Species 10 Clave de las Especies 11 ACCOUNTS OF THE SPECIES 11 Colostethus aeruginosus new species 11 Colostethus argyrogaster Morales and Schulte 13 Colostethus craspedoceps new species 16 Colostethus elachyhistus Edwards 18 ColostetJius eleutlierodactylus new species 21 Colostethus idioiftelus Rivero 23 Colostethus iiisulatus new species 24 Colostethus leiicophaeus new species 26 Colostethus mittermeieri Rivero 28 Colostethus )iexipus Frost 30 Colostethus ornatus Morales 32 Harvard Universsl^/ © Natural History Museum, The University iit Kansas . r nrnVi^-XZ'*'!^ i-U'J-^iBJ ISSN No. UW4-0782 Scientific Papers Natural History Museum The University of Kansas 15 November 2004 Number 35:1-49 Frogs of the Genus Colostethus (Anura; Dendrobatidae) in the Andes of Northern Peru ^AC■z. By ^^gp,AR^ William E. Duellman Nntuivl H;s/i)n/ MusciDii and Biodiversity Research Ce)iter The Universiti/ of Kansas Laioreiiee, Ka)isas 66045-7561 USA CONTENTS ABSTRACT 2 RESUMEN 2 INTRODUCTION 3 Acknowledgments 3 MATERIALS AND METHODS 4 ANDES OF NORTHERN PERU 4 SUMMARY OF TAXONOMIC CHARACTERS 5 IDENTIFICATION OF SPECIES 10 Key to the Species 10 Clave de las Especies 11 ACCOUNTS OF THE SPECIES 11 Cohistethus aerii^^iiiosiis new species 11 Colostethus argyro^iaster Morales and Schulte 13 Colostethus craspedoeeys new species 16 Colostethus elachyhistus Edwards 18 Colostethus eleutherodacti/his new species 21 Colostethus idioiuelus Rivero 23 Colostethus iiisulatus new species 24 Colostethus leueophaeus new species 26 Colostethus luitteniieieri Rivero 28 Colostethus lu'xipus Frost 30 Colostethus oniatus Morales 32 © Natural History Museum, The UniverMt\ of Kcinsds , rAmn'^.r'-'l V'& i-Uu.v-ieJ ISSN No. 11)94-0782 Harvard Univors.iV Museum Scientific Papers, Natural History Museum, The University of Kansas Colosti'tlnifi pocciloiiotiifi Rivero 33 Colostetlnis piilchcrriinu^ new species 34 Colostethus sordidatu^ new species 35 Colostctlnif i^piloto;^nfti'r new species 37 Colostethus s\/lvaticiis Barbour and Noble 38 Colostethus utcubambensis Morales 40 Status of Phyllobates peruviamis Melin 41 DISCUSSION...! 41 LiPE History and Tadpoles 41 Ecology 43 Bioceography 44 LITERATURE CITED 45 APPENDICES 46 Appendix 1. Specimens and Localities 46 Appendix 2. Gazetteer 48 abstract Seventeen species of the dendrobatid frog genus Colostethus are recognized in the Andes and adjacent lowlands in northern Peru. Nine of these species have been reported previously from the region and eight are described as new. Tadpoles of eight species are described and illustrated, and growth trajectories of six species of tadpoles are compared. The greatest diversity of Colostethus is in humid montane forests in the northern part of the Cordillera Central, where five species occur at eleva- tions above 1 600 m; six other species occur at elevations below 1 000 m. Two of these also range into the valley of the Ri'o Maranon, where two additional species occur. Colostethus elachyhistus is widespread in the Andes in southern Ecuador and in the northern part of the Cordillera Occidental, including the Cordillera de Huancabamba, the higher parts of which are inhabited by C. sylvaticus. Colostethus ela- chyhistus and C. nexipus are the only species that occur in Ecuador and northern Peru. Seven instances of sympatry reveal slight differences in size between co-occurring species at most of these sites. Key Words: Anura, Dendrobatidae, Colostethus, new species, tadpoles, Andes of northern Peru, tax- onomy, biogeography. RESUMEN Se reconocen diecisiete especies de dendrobatidos del genero Colostethus en los Andes y tierras bajas adyacentes del norte del Peru. Nueve de estas especies han sido reportadas previamente para la zona y se describen ocho especies nuevas. Los renacuajos de ocho especies son descritos e ilustrados; se compara los patrones de crecimiento de los renacuajos de seis especies. La mas alta diversidad de Colostethus se encuentra en los bosc]ues humedos montanos de la region norte de la Cordillera Central, en donde habitan cinco especies por sobre los 1 000 m; dos de estas especies tambien se encuentran en el valle del n'o Maranon; en el mencionado valle existen dos especies adicionales. Colostethus elachyhistus habita en los Andes del sur del Ecuador y en la region norte de la Cordillera Occidental del Peru, incluyendo la Cordillera de Huancabamba, en cuyas partes mas altas se encuen- tra C. sylvaticus. Colostethus elachyhistus y C. nexipus son las linicas especies que se encuentran en el Ecuador y en el norte del Peru. Siete ejemplos de simpatn'a revelan pcquenas diferencias en el tamano de especies en la mayorfa de los lugares mencionados. Palabras Clave: Anura, Dendrobatidae, Colostethus, nuevas especies, renacuajos, Andes del norte del Peru, taxonomfa, biogeografi'a. COLOSTETHUS OV NORTHERN PeRU INTRODUCTION Frogs currently allocated to the genus Colo$ti'tliu> are especiallv speciose in the Andes of Colombia (Ruiz- Carranza et al., 1996; Rivero and Serna, 2000; Grant and Ardila-Robayo, 2002) and Ecuador (Coloma, 1995; Santos et al., 2003). Many fewer species ha\'e been reported from the highlands of northern Peru (Rivero, 1991; Duellman and Wild, 1993; Morales and Schulte, 1993; Morales, 1994). Field parties from the Natural History Museum at The University of Kansas worked in the Andes of northern Peru (departamentos de Amazonas, Cajamarca, Piura, and San Martin) in 1970, 1975, 1979, 1989, and 1991. This fieldwork produced more than 400 specimens of Colostc- thus that are deposited in the Natural History Museum at The University of Kansas and in the Museo de Histo- ria Natural at the Universidad Nacional Mayor de San Marcos in Lima, Peru. In addition to these specimens, additional material was collected by: Kenneth E, Camp- bell in 1970 and deposited in the Florida State Museum; Richard Thomas and others in the 1970s and deposited in the Museum of Natural Science at Louisiana State Uni- versity (some specimens transferred to The University of Kansas); Russell A. Mittermeier in 1978 and deposited in the Museum of Comparative Zoology at Harvard Univer- sity; and Rainer Schulte, a resident of Tarapoto, Peru, in the 1980s and deposited in the Museo de Historia Natural in Lima. The first species of Colostetliiif recorded from northern Peru was "Phyllobatcs" sylvatkus collected by G. K. Noble in 1916 and described by Barbour and Noble (i920). From that inauspicious beginning, more than seven decades passed before Rivero (1991) described Colostethus idiomclus and C. mittcrmeieri from Venceremos on the eastern slope of the northern part of the Cordillera Central and C. poccilonotits from the Rio Maranon Valley. The principal Peruvian researcher on Colostctlnts, Victor R. Morales, described C. iitcubamhensis and C. argywgaster from the northern Cordil- lera Central, and middle Ri'o Maranon valley, respectively (Morales 1994; Morales and Schulte, 1993). In a revision of the Colostethus triliueatus Group, Morales (2002) named C. ornntus from the Tarapoto region at the eastern base of the Cordillera Central. The range of one Ecuadorian species, C. clachyliiftiis, was extended into the Cordillera de Huan- cabamba in Peru by Duellman and Wild (1993); another species described from Ecuador, C. ncxipu^, was reported from Peru by Coloma (1995). Thus, to date, nine species of Colostethiis have been reported from the Andean region of northern Peru. The extensive collections from northern Peru include eight new species described herein and the first Peruvian records of another species previously known only from Ecuador. The purposes of this paper are to describe the new species and to review the species of Colostethiis known to occur in the Andes and associated mountain ranges, inter- vening valleys, and the immediately adjacent Amazonian lowlands in northern Peru (departamentos of Amazonas, Cajamarca, Piura, and San Martin). Despite the large number of specimens now known from the region, our knowledge of the anurans in the Andes of northern Peru is far from complete. More thorough collecting certainly will extend the known ranges of many species, and the explora- tion of previously uncollected valleys and mountain slopes surely will reveal additional species, as is evident by the discoveries in other taxa (e.g., Centrolenidae by Duellman and Schulte, 1993, and Eleutherodactylus by Duellman and Pramuk, 1999). Acknowledgments I am indebted to mv field companions — Thomas J. Berger, David C. Cannatella, Fernando M. Cuadros V., Mi- chael E. Morrison, Rainer Schulte, John E. Simmons, Linda Trueb, and John J. Wiens — whose efforts resulted in man\' of the specimens of Colostcthus; to many residents of north- ern Peru who provided shelter, food, and assistance to the field parties; to Jesus H. Cordova and Victor R Morales for courtesies at the Museo de Historia Natural in Lima; to B. Anthony Luscombe and Rainer Schulte for logistic support; to Cesar Aguilar, Christopher C. Austin, Jesus Ccirdova, Alison Jennings, Kenneth L. Krysko, Max A. Nickerson, and Jose P. Rosado for the loan of specimens; to Stefan Lotters for information on, and photos of, a Peruvian species, and to JuajT Manuel Guayasamin for aid with the Resumen and the key in Spanish. This manuscript benefited from critical review by Luis A. Coloma, Victor R. Morales, and espe- cially Taran Grant and thorough editing by Linda Trueb, who also illustrated the tadpoles and provided images of preserved specimens; I thank them profusely for their ef- forts. Collecting and export permits were issued by Luis J. Cuento Aragon, Armando Pimental Bustamento, Gonzalo Bravo Meji'a Munoz, and Jose Purisaca, Direccion General Forestal y de Fauna, Ministerio de Agricultura, Lima, Peru. The research reported herein is part of a study on the pat- terns of speciation and biogeography of Andean anurans supported by a grant (BSR 8805920) from the National Science Foundation (W. E. Duellman, PI). SciENTiiic Papers, Natl'rai, History Museum. The University of Kansas MAIERIALS AND METHODS Specimens in museum collections nre identified by their catalogue numbers preceded by the following codes: BMNH = British Museum (Natural History); KU = Natu- ral History Museum, University of Kansas; LSUMNS = Museum of Natural Sciences, Louisiana State University; MHNSM = Museo de Historia Natural, Universidad Nacio- nal Ma\or de San Marcos; MCZ = Museum of Comparative Zoolog)', Harxard University; NHMG = Naturhistorika Museet Goteborg; UF = Florida Natural History Mu- seum. All specimens from northern Peru and southern Ecuador that ha\'e been studied are listed in Appenciix 1. Localities from which specimens have been examined are listed with their geographic ctiordinates, elevation, and vegetation type in Appendix 2. Geographic coordinates were obtained from maps, principally the Mapa Fisico Politico 1:1,000,000 (1973) but also the Carta Nacional del Peru, 1:100,000 (1986) for those regions so mapped; the Institute Geografico Militar, Lima, Peru, produced both sets of maps. ANDES OF NORTHERN PERU Measurements, definitions of coloration and structural characters, and numbered characteristics in diagnoses follow those of Duellman and Simmons (1988) v\ith the addition of testes color (Coloma, 1995) and the median lingual process (Grant et al., 1997). Measurements were taken with dial calipers to the nearest 0.1 mm. If sex and reproducti\'e condition were not evident externally (swol- len Finger 111 or vocal sacs in males; eggs visible through body wall in females) sex was determined by dissection. Males with granular testes were determined to be adults; females with expanded, convoluted oviducts or large ova were considered to be adults. Throughout, snout-vent length is abbreviated SVL, and larval labial tooth row formula is abbreviated LTRF. Tadpoles were staged accord- ing to Gosner's (1960) table; terminology for tadpoles was based in Altig and McDiarmid (1999). Neuromasts were not evident in any of the tadpoles under 20x magnifica- tion; therefore, lateral line systems are not included in the descriptions of tadpoles. The geological history, physiography, climate, and vegetation of the northern Andes were described by Du- ellman and Pramuk (1999); thus, only a thumbnail sketch is presented here. A major orogenic difference between the northern and southern Andes is the deflection in a region of complex interdigitation of relatively low ranges, basins, and valleys — the Huancabamba Depression en- compassing the Ecuador-Peruvian border (Figs. 1 and 2). In this region, major rivers flow north or south between intervening north-south highlands. The backbone of the Andes in Peru is the Cordillera Occidental, which north of the Nudo de Pasco in central Peru is separated from the Cordillera Central by the valley of the Rfo MarancSn; this Cordillera has its northern terminus at about 5'S latitude and is separated from the Cordillera Oriental b\- the \ al- ley of the Rfo Huallaga, which turns eastward and forms the northern edge of the Cordillera Oriental at about 7 S latitude. Nine types of vegetation can be recognized by simpli- fying the system used by Tosi (1960), as follov\-. Desert scrub. — Dr\' Pacific lov\ lands receiving less than 150 mm of rain annually with legumes and cacti near the western base of the Andes up to as much as 500 m. Thorn forest. — A forest dominated by legimies, cacti, and othiM drought-resistant trees in areas receiving less than 500 mm of rain annually on the western slopes of the Cordillera Occidental to about 1200 m and in the interior valleys of the rfos Chamaya, Chichipe, Huancabamba, Marafion, and Utcubamba. Dry forest. — In areas receiving 1000-2000 mm of rain annualh', this forest develops peripheral to thorn forest and in the rain shadows at elevations of 25t)-600 m of the Ri'o Mavo and middle Rfo Huallaga. Montane dry forest. — Isolated patches of this forest exist at elevations of principally at 2000-2500 m in the up- per Rio Maranon Valle\', in the Cordillera Occidental and Cordillera Central. Humid montane forest. — This is the dominant type of forest on the lower slopes (up to about 2500 m) of the Cordillera de Huancabamba and Cordillera Colan. Very humid montane forest. — This is the so-called cloud forest characteristic of ele\'ations of 2500-3000 m on windward slopes of the Andes that recei\-e 1000-2000 mm of rain annually. I his type of forest occurs at high elevations of the Cordillera de Huancabamba and Cordillera Colan, and the eastern slopes of the Cordillera Central. Humid subtropical forest. — Forest receiving 1000- 3000 mm of rain annually at elevations of 500-1900 m; this is characteristic of the eastern slopes of the Cordillera Central and middle Rio Mayo Valley. Subtropical pluvial forest. — In northern Peru, this type of forest, which receixes rainfall in excess of 4000 mm annually, is known only from the crest (> 3000 m) of the Cordillera Central west of the Ki'o Mavo. Humid tropica! forest. — The Amazonian rainforest at elevations < 300 m receiving rainfall in excess of 2000 mm annualh'; great species diversitx' of trees, some of which form a ciimplele, or nearly so, canopy 30^0 m above the ground. COLOSTETUVS OV NoRTHKRN PeRU SUMMARY OF TAXONOMIC CHARACTERS Compared to their colorful allies (AUobates, Cryp- topln/llobntes, Dendrobaics, Epipcdobatcs, Miuyobates, and Phyllobntes), frogs of the genus Coloftethufi are rather drab in appearance. Living individuals of most species have distinctive colorations, but individuals that were easily distinguished in life commonly are difficult to distinguish once they have been preserved. Consequently, taxonomic studies necessitate detailed and comparable diagnoses and descriptions. Edwards (1971) made the first attempt to standardize diagnostic character states in Colofitctliu> bv recognizing 21 features in adults and larvae. Duellman and Simmons (1988) reduced the nimiber to 15 characters of adults. This scheme was partially followed by Morales (1994, 2002) and modified slightly by Coloma (1995) with the addition of tes- tes color; Grant et al. (1997) added the presence or absence of a median lingual process to the diagnostic characters of Colostethus, and Grant and Castro (1998) and Grant and Ardila-Robavo (2002) added the so-called black arm band, characteristic of the Colostctliu^ nviio^i Group. Lotters et al. (2003) added cloacal tubercles and cloacal sheath to their diagnosis. Herein, I define 17 diagnostic features. 1. Size and proportions. — Snout-vent length (SVL) is the straight-line distance from the tip of the snout to the posterior terminus of the body. In making comparisons, 1 use some subjective terms — small (< 20 mm), moderately small (20-25 mm), and moderately large (> 25 mm). Head length is the chord of head length, i.e., the distance from the tip of the snout to the posterior edge of the articulation of the jaw. Head width is the greatest width of the head. The eye and tympanum sizes are horizontal measurements of the eye and visible part of the tympanum, respectively, and the eye-nostril distance is measured from the ante- rior corner of the orbit to the posterior edge of the nostril opening. Tibia length is the distance from the knee to the distal end of the tibia. Foot length is the distance from the proximal edge of the inner metatarsal tubercle to the tip of Toe IV. Snout-vent lengths vary from less than 18.0 mm in males of Colostetliiis nrgi/wgastcr, claLht/histiis, iiisulntii^, and oniatiis to 30.0 mm or more in C. sordidntiis and C. si/lvnticus. The latter two species and C. ncrugiiiosus and C. utcubniiibeiisis have the greatest sexual dimorphism in SVL (Table 1). Proportions are much the same in all species of Colostetliiis in northern Peru. The ratio of head length to SVL is slightly more than the ratio of head width to SVL in most species, but the ratios are reversed in C. acriigiiiosits and are equal in C. iiisiilatiis and C. sordidntiis. The length of the snout is expressed as the ratio of the eye-nostril distance to the length of the eye. Ratios are less than 60% in three short-snouted species (C. elacln/histiis, Table 1. Body sizes (SVL in mm) of Coloslethu^ in the Andes of northern Peru. Values include range (mean, sample size); SexD = sexual dimor- phism. Species Males (em ales SexD c. twruiiitwsu^ 22.1-25.0 (23.0, S) 29,2 1.27 c. ar^i/w;^ii^tcr 17.5-19.8(18.7,2) 19,0-22.1 (20.4,3) 1 ,09 c. crnspcdoci'p'^ 19.1 18,.5-20.5(19.7, 5) 1,01 c. tiaclii/lustiis 15.2-20.9(17.7,43) 17.2-23.6(20.7,53) 1,17 c. elfiithcwitactyliii 21.0 22.7 1.08 c. iiliomeliis 19.5-24,8(22.0.38) 23,0-27.8(25.1.23) 1.14 c. iiiiulntui 17.2-22.0(19.7,9) 20.2-26.0 (22.8. 22) l,lh c. IcHCophnfu^ 24.7-25.2(25,0, 3) 26.1 1,04 c. miUcniicicri 27.2 28.2 1.04 c. nexipus 18.6-20.5(19.3,10) 20.0-23.1 (21.4, 11) 1,11 c. ornntus 15.7-16.9(16.3,2) 15.5-18.4(17,0,2) 1.04 c. pulchcrrimiii 28.2 26.4-29.7 (28.2, 4) 1 ,00 c. poccilouotu^ 19.7 20.8-25.2 (22.9, 3) 1,16 c. iordidatus 24.2-29.9 (25.7, 8) 31,8-36.1 (33.6,3) 1,31 c s;'i7ofo\>(?sfi'r — 24,0 — oniatiis, and spilotogastcr), more than 70'7< in four long- snouted species (C. aemgiiiosiis, craspedoccps, idionicliis, and sordidntiis), and 60-70' <'. in the other species. The ratio of the length of the tympanum to the length of the eye is less than 50''r'. in five species (C. Iciicoplmeiis, inittcnncieri, nexi- pus, sordidntiis, and iitciibaiiibeusis), more than 60% in two species (C. idionieliis and C. pidclierriiuiis), more than 78'":. in C. spilotognster, and 50-60% in the other species. Tibia length is proportionally slightly greater than foot length in most species, but the ratio of each to SVL is essentiallv the same in C. poeciloiiotus, and foot length is proportionately more than tibia length in C. leucoplineiis, pulcherriinus, and siflvnticiis. In the species accounts, 1 describe the limbs in general, subjective terms that are used in comparison with other species in the region. Thus, the forearm (elbow to wrist) is described as long, slender, or robust. The hind limb is described as short, long, robust, or slender; unless specified otherwise this pertains to the thigh and shank. 2. Disc on Finger IIL — In most species of Colostetliiis in the Andes of northern Peru, the terminal discs on the fingers, exclusive of the thumb (Finger I) are expanded to be 1.5-2.0 times the width of the preceding (antepenultimate) segment of the digit (Fig. 3). The discs are barely wider that the preceding segment in C. elncln/liistiis, piilcherrinnis, and sylvaticiis, and not expanded in C. idiomeliis (Table 2). Standard comparison is made on Finger III. 3. Relative lengths of Fingers I and IL — When the Fin- gers 1 and 11 are adpiessed. Finger I (thumb) is shorter than Finger II in Colostetliiis nexipus and longer than Finger II in ScinN'TiFic Papers. Nati'rai, History Museum, Thh University OF Kansas 78 / /" 7"^. .- ( /" /' ( n 11 \ El O ro Saraguro , San Lucasi I -A' ' o Abra de Zamora o • Cities and towns o- Collecting localities Major roads Rivers International boundary Departmental and provincial boundaries 50 100 r-<;s^' iLoja o / Zamora .^ / Macara^ — -"'^Cahamanga i ^' ■' Ay abaca ^ \ ) <& ,/ \ / 'V /' \ V ,) (' \ Piura \ _ ; \ . . ' ) Tabaconas V Huancabamba . / v 'K- l El Tambo / ] [J K- o -^ {Chihaco ^ / \ \ L r e t o \V \ rAbra Pardbde Miguel ^-^ Venceremos — - , Balzapata ^o^ Alva rT' of'-^ , Nx^ //^ \ x^^^ o/ Pomacocha PorcJilr^K=^-B=^'''^ V- Pedro^^P\ngenio ^. La Riojo v ^■ Porculla J -~~-R^ ^ \ o Ga„o \chachapoya>- \ ^-i,-^ \, ' "^ ^ Santa Cruz \%- ^. T.^^^.X'i^- — ^ ^ '^ Lambayequek s°^ \* <>. ^'"go'Kr ^Moimopampa\ t ' - O; Q Ss ^ .79- Caiabambai^ \ \ \ 11 Fie- 1. Political map ot thu Andes ol northom I'oru and southern Fauidor shins inj; niajor risers and KHalities mentioned in the text. CoiosTFTHis oi- Northern Pkrl! 1 1 < 1000 ni iOO 1 1 1000-2000 m 1 1 2000-3000 ni ' 3000-4000 m > 4000 m 50 Kilometers Fig. 2. Physiographic map ot the Andes of northern Peru and southern Ecuador. Scientific Papkrs, Natural History Museum, The University of Kansas Table 2. States of structural characters and coloration in Cd/iisfcf/ms from the Andes of northern Peru. Finger lengths Disc Width Lateral fringes Inner tarsal fold Toe webbing Lateral stripes Species Dorso- Oblique Ventro- C. aeruginosus I > II 1.5X Yes Distal i/i No No \o C. arf>\rof>asler I > II 1.5X No Weak Basal IIl-lV No Yes Yes C. craspciloceps 1 > II 1.5X No — — No Diffuse No C. clachyhistus I = II I.IX Yes Distal '/2 1/4 No Yes No C. eleulhewdactyhis I = II 2.0X No Distal '/^ Yes Yes No C. idiomelus I = II Not Yes Distal '/i Yes No No C. IMSU/fl/KS I > II 1.5 X Yes Distal '/2 Basal No Yes No C. teiicophaetis I > II 1.1 X Yes Distal >^ 'A No Groin No C. mittermeieri I = II 1.5X Yes Distal '7 Va, No No No C. iiexipus I < II 1.5X Yes Weak Vi Yes Yes No C. onmtiis I.5X No — Basal Yes No Yes C. poecihiiotiii 1 = 11 1.2X No Distal '/^ — No Short No C. pukiwrnmiis I > II 1.1X Yes Distal 1^ — No Yes No C. sordidciius I = 11 2.0X Yes Distal % 3/4 No Yes No C. spilotoga:>tcr 1 = II 1.5X Yes Distal 'A. Basal Yes No No C. si/lvaticus I = 11 I.IX Yes Weak — Yes No No C. iitciihamheiisi:i I >n 1.2X No Distal '/7 Basal Yes Yes No C. aeruginosus, argi/rogastcr, craspcdoccps, insulatus, Icucophacus, omatiis, pidchcniiiius, dnd utciibambciisis. Fingers 1 and II are about equal in length in the other species (Table 2). 4. Fringes on fingers. — The tingers of Colostctliiis never have more than basal webbing, but many species (e.g., C. twriis^iiiosiis, elachyhifitiis, idioiiifliis, iusiilatus, Icucophacus, nexipus, pulclicrrinius, sordidatus, spHotogastcr, and si/lvali- cus) usually have fringes on the inner (preaxial) edges ot Finger IV and both sides of Finger I! and 111. The other spe- cies in northern Peru lack fringes (Table 2). The degree ot development of fringes is variable; therefore, the standard comparison is made on Finger III. The fringes are broad and conspicuous in C. aeruginosus and C. Icucophacus, rather thick and keel-like in C. idiomelus, and narrow in other spe- cies that have fringes. Nearly all of the specimens treated herein were preserved in the same manner, so states of fringes can be compared with relative ease; however, slight desiccation can result in fringes appearing more extensive than in properly prepared specimens. 5. Disc on Toe IV. — The variation in expansion of the terminal discs on the toes approximates that on the fingers (Character 2). Standard comparison is made on Toe IV. In three species (Colostcthus craspcdoccps, clachi/hislus, and uiionu'lus), the toe discs are slighth' more expanded than the finger discs. 6. Fringes on toes. — The presence, absence, and nature of lateral fringes on the toes parallel the condition of the fringes on the fingers (Character 4). Standard comparisiMi is made on Toe IV. 7. Tarsal folds and tubercles. — Of the species of Colo- stctlius in the Andes of northern Peru, only C. clcuthcrodac- ti/lus has a noticeable outer tarsal fold (= metatarsal fold sensu Grant and Rodriguez, 2001) that extends the lull length of the tarsus; a weak outer tarsal fold is present in some individuals of C. clachi/histus. Except for C. craspcdo- ccps and C. or)iatus, all species in the region have an inner tarsal fold. This fold is sigmoid on more or less the distal half of the tarsus; the fold is weak in C. argyrogaster and C. cladn/histus, and it is flaplike in C. aeruginosus and C. pul- chcrriiiuis. The inner tarsal fold is contiguous with the inner metatarsal tubercle and extends to a tubercle at about the midlength of the tarsus in five species; the tarsal tubercle is small in C. argyrogaster, iiisulatiis, and utcubambensis, and it is much larger in C. elcutherodactylus. Two species that lack an inner tarsal fold have tarsal tubercles; the tubercle is small in C. craspcdoccps and elongate and cur\ed in C. ornatus. 8. Toe webbing. — The toes are webbed basal!) in Colo- stcthus argyrogaster, insulafus, ornatus, and utcubambensis. In two species (C. elachyhistus and C. niitteniicicri), the toes are about one-fourth webbed; the\' are about one-half webbed in C. Icucophacus and C. nexipus, and about three-fourths webbed in C. sordidatus. Other species of Colostcthus in the region lack webbing between the toes (Table 2). In the in- dividual accouiits of those species having more than basal webbing, the webbing is described following the protocol of Sa\'age and Heyer (1967) as modified by Myers and Puellman (1982). 9-11. Pale longitudinal stripes. — The color patterns of most species of Colostcthus include one or more kinds of pale stripes; the combinations of these stripes are help- ful in identifying species (Table 2). The stripes usually are pale tan or cream, but the\' are orange-red in C. nexipus. Two species in the northern Andes, C. aeruginosus and C. mittermeieri, lack longitudinal stripes. The dorsolateral stripe (Character 9) extends from at least the occiput to the posterior end of the body above the insertion ot the COLOSTETHUS OF NORTHERN PeRU hind limb; in most species, the stripe is continuous ante- riorh' along the outer ecige ot the upper eyelid and along the canthus rostralis to the tip of the snout. Dorsolateral stripes are present in only six species in the region (C. dciitlii'ivdactylus, luwipiis, oniatiis, fipiloto^y^a^tci-, $iflvnticiis and iitciibiiiiibeii^is). The oblique lateral stripe (Character 10) is present on the flank in man\' species; it originates in the groin (at about midheight of the insertion of the hind limb) and ma\' extend anteriorly to the midflank region or exen to the posterior edge of the upper eyelid. An oblic]ue lateral stripe is absent in C. onintus and C. si/Ivaticiif. The stripe is short and confined to the groin in C. Icucoplmcus and C. pocciliviotiis; it is diffuse in C. cmspcdoccpf and is fragmented in C. clciitlicrodtu'tj/lus and some individuals of C. ;/('.v//'((.s. In some species, a pale ventrolateral stripe (Character 11) is present; this is most noticeable in species ha\'ing a dark x-enter. Among the species in the Andes of northern Peru, a ventrolateral stripe is present only in C. argywgastcr and C. oiiiatiis. 12. Coloration of the gular-chest region. — Uniform coloration of the gular-chest region characterizes adults of seven species of Colostcthus in the Andes of northern Peru — cream in C. argyrogastcr, cleutlieyodncti/his, Icucoplm- t'i/s, lu'xipus, pocciloiiotui': pale gray in C. iitcubaudh'iisis, grayish brown or cream in C. cmspedoceps, and gray or white in C. onintus. The most common markings in the gular-chest region are a pair of dark brown to black spots posterolaterally on the throat; these are present in adults of C. elnchyhistiis, idioiiicliis, iiifidatiis, iiiittenueicri, and present or not in C. sylvnticus. Other color patterns on the gular- chest region consist of cream spots on a brown background (C. nerugijiosiis and C. piilchcmiutis), many small brown spots (C. spilotogastci), or a dark gray intrusion laterally on the throat in C. sordidatiis (may be an incipient pair of dark spots). 13. Coloration of abdomen. — In life, the entire ab- dominal region is pale cream to creamy yellow or white in most species of Coloftctlnis in the Andes of northern Peru, but it is silver in C. nrgi/wgnster and gray or white in C. foididatub. Other species have distinctive patterns on the abdomen — mottled gray and white anteriorly and yellow posteriorly on abdomen anci ventral surfaces of hind limbs (C. idioincliis and C. sordidatiis), mottled throughout with gray (C. aerugiuosis), gray with white spots (males of C. utcubambeiifis and some males of C. elncbyhistiis), mottled with gray anteriorly (C. iiiittcniwicii), or cream with brown mottling (C. piilclicniiiiiis) or spots (C. spdotogastcy). 14. Sexual dimorphism in coloration. — As is evident from the character states in Characters 12 and 13, sexual dimorphism in coloration of preserveci specimens is evi- dent in five species of Colostethub in the Andes of northern Peru. Sexual dimorphism in ventral cokiration occurs in four species — gray throat or abdomen in males and cream or white in females (C. onmtus, fioididatii>, and utcubain- boisis), or gray with cream spots in some females versus cream in males (C. einchyhistiis). The males of Colostethiis ivgyiognstL'v are distinctive in having an orange inguinal region in life. 15. Median lingual process. — The presence of a me- dian lingual process on the dorsal surface of the tongue was reported in nine species of Colostethus from Chocoan Ecuacior and Colombia, Guyanan Venezuela, and Amazo- nian Brazil (Grant et al., 1997; Grant and Rodriguez, 2001; Myers and Donnelly, 1997). No species from Peru is known to possess this structure. 16. Swollen fingers in males. — The preaxial part Finger III is swollen laterally in males of many species of Colostethus, especiallv in members of the Cohstctluis tri- liucntus Group (Morales, 2002). Finger II also is swollen in some males of C. triliiicntus (Grant and Rodriguez, 2001), and in several other species, e.g., C. iiiguinalis (Grant, 2004) and C. iiubicoln (Ibaiiez and Smith, 1995). The only species with a swollen Finger III in the region under consideration is C. onintus, a member of the Colostcthus triliiicntus Group (sensu Morales, 2002). 17. Testis color. — Coloma (1995) noted that the testes are brown in three species of Colostetluis in Ecuador. All male Colostethus examined from Peru have white testes, as do most species in Ecuador. In addition to the foregoing characters that are used in the diagnoses of species of Colostethus, other structural features and aspects of coloration differ interspecifically. Structural features include texture of the skin on the dor- sum, which is smooth in most species, but with minute tubercles posterior to the sacral region in C. neruginosus, nrgywgnster, crnspiedoceps, and idioinelus. The skin on the dorsum is finely shagreen in C. iiiitteniieieri, sordidntus, and sylvntieus, and shagreen with minute tubercles posteriorly in C. iiexipus and C. poeeilouotus. The skin on the flanks usu- ally has more texture (usually shagreen) than that on the dorsum; in most species, it is shagreen, but it is smooth in C. neruginosus, nrgyrognster, elachyhistus, and uteidmuibensis, anci weakly granular in C. crnspedoceps and C. sordidntus. Distinct tubercles are present below the tympanum in C. poeeilouotus. In most species of Colostethus in the Andes of northern Peru, the cloacal opening is directed posteriorly or posteroventrally at the upper level of the thighs; how- ever, in C. ernspedoeeps and C. nexipus, the cloacal sheath is relatively long and the opening is located at the midlevel of the thighs and directed \-entrally. None of the species of Colostethus in the Andes of northern Peru has supernumerary tubercles on the digits. Subarticular tubercles are always present and usually smaller on the toes than on the fingers; these tubercles are 10 Scientific Papers. Natural History Museum, The University of Kansas small and round in most species, but they are distinctly subconical on the fingers of C. clncln/hi^tKs, ek'iitherodacty- lus, insulatus, mittermeieri, oniatiis, and si/hmticm. Nuptial excrescences were not discernable on any males. In addition to the combinations of longitudinal stripes and differences in ventral coloration, other features of the color pattern are noteworthy in some species. In most spe- cies of Colo^tftlniti, the dorsum of the head and body is dull brown with darker brown to black flecks, spots, or irregular marks, but a few species have distinctive colors — dull green in C. acnigiiio^ii^, reddish brown in C. iu^ulatiis, grayish brown in C. leucophaeus, grayish green with coppery brown markings in C. piilcherrimus, and olive brown to coppery brown in C. sylvatkus — or patterns — V-shaped marks in C. idiomelus, X-shaped mark connected to a teardrop-shaped mark posteriorly in C. oniatus, middorsal brown stripe in C. spilotogn^ter, and chevrons in some C. sylvaticus. The dorsal surfaces of the hind limbs are tan or pale brown, usually with two to four dark brov\'n transverse bars on each segment; there is only one bar on each segment in C. oniatiis. Irregular dark brown markings are present on the dorsal surfaces of the hind limbs in C. cmfpcdoccp^ and C. spilotogaster, and dark markings are absent on the dorsal surfaces in C. nr;f\/wgaster. A distinctive longitudinal dark brown stripe is present on the anterior surface of the thigh in C. argyrogaster, craspedoceps, and poecilonotus, and on the posterior surface in C. argyrogaster, craspedoceps, and leuco- phaeus. In contrast, C. poeciloiiotus has a pale cream stripe on the posterior surface; C. insulatus, on the anterior surface; C. eleutherodactylus, diagonal pale stripe on the anterior surface. Seven species (C. elachyhistus, idiomelus, leucopha- eus, or}iatus, poeciloiiotus, sordidatus, and spnlotogaster) have longitudinal brown stripes on the upper arms; the mark is crescent-shaped in C. sordidatus. IDENTIFICATION OF SPECIES Key to the Species of Coeostethus i\ northern Peru 8. 1. Finger I < II; toes about one-half webbed; dorsolateral and oblique lateral stripes present C. iie.xipus 9. Finger I > II or I = II; webbing and stripes variable 2 2. Finger I > II; digital discs expanded 3 1(1. Fingers I and II equal in length; digital discs expanded or not 11 3. Lateral fringes present on fingers 4 Lateral fringes absent on fingers 8 1 ' • 4. Oblique lateral stripe present 5 Oblique lateral stripe absent 7 12. 5. Oblique lateral stripe present only in groin; toes about one-half webbed C. leucophaeus 13. Oblique lateral stripe extending from orbit to groin (may be indistinct anteriorlv); toes with no more than |_| basal webbing 6 6. Toes webbed basally; flanks gray with brown mot- tling; pair of dark spots JalcialK' on throat C. insulatus Toes not webbed; flank black with pale llecks or streaks; throat cream with or without brin\n mot- tling \(x C. pulcherrimus 7. Toes not webbed C. aerugiuosus Toes about one-fourth webbed C. iiiittenneieri Oblique lateral stripe absent C. oniatus Oblique lateral stripe present 9 Dorsolateral stripe present C. utcubambensis Dorsolateral stripe absent 10 Oblique lateral stripe distinct, extending from orbit to groin C. argyrogaster Oblique lateral stripe diffuse, restricted to groin C. craspedoceps Dorsolateral stripe present 12 Dorsolateral stripe absent 15 Digital discs greatly expanded 13 Digital discs not or barely expanded 14 Oblique lateral stripe present C. eteutlierodacti/lus Oblique lateral stripe absent C. spilotogaster Lateral fringes present on fingers C. si/h'aficus Lateral fringes absent on fingers C. idiomelus Oblique lateral stripe only on posterior part of body; toes about two-thirds webbed C. sordidatus C^biique lateral stripe extending from orbit to groin; toes no more than one-fourth webbed 16 Lateral fringes present tin fingers; toes at least webbed basalK- C. elachyhistus lateral fringes absent on fingers; toes not v\'ebbed .. C. poecilouotus COI.OSTETHUS OF NoKTHHRN PkRU Clave para i,as Especies de Coio^tethus dee norte del Peru 1 . Dedo manual I < Dedo manual II; dodos de los pies con mombranas interdigitales que se extienden aproximadamente hasta la mitad de su longitiid; li'neas dorsolaterales v oblicuas presentes C. iifxipii> Dedo manual I > Dedo manual II; membranas inter- digitales y lineas en el doiso variables 2 2. Dedo manual 1 > II; discos de los dedos manuales expandidos 3 Dedo manual I con longituci similar a la del Dedo manual II; discos de los dedos manuales expandidos o no expandidos 11 3. Pliegues laterales presentes a lo largo de los dedos manuales 4 Dedos manuales sin pliegues laterales 8 4. Linea lateral oblicua presente 5 Li'nea lateral oblicua ausente 7 5. Li'nea lateral oblicua presente solo en la ingle; dedos de los pies cc^n membranas interdigitales cjue se extienden aproximadamente hasta la mitad de su longitud C. Iciicopliacus Li'nea lateral se extiencie desde la orbita ocular hasta la ingle (aunque puede ser indistinta anteriormente); dedos cie los pies con membranas interdigitales solo en la base o sin membranas interdigitales 6 6. Dedos de los pies con membranas interdigitales basales; flancos del cuerpo color gris con un patron irregular de manchas color cafe; garganta con un par de puntos oscuros (un punto en cada lado de la garganta) C. iiibulntu> Dedos de los pies sin membranas interdigitales; tlan- cos del cuerpo color negro con pequehas manchas claras; garganta crema con o sin patron irregular de manchas color cafe C. pulclifiriiiui!^ 7. Dedos de los pies sin membranas interdigitales.. ..C. aentgiiiosui Dedos de los pies con membranas interdigitales que se extienden hasta aproximadamente 1 /4 de su lon- gitud C. niiticnncicri 8. Li'nea lateral oblicua ausente C. oruatus Li'nea lateral oblicua presente 9 9. Li'nea dorsolateral presente C. iitaibiwibcnsis Li'nea dorsolateral ausente 10 1 1). Linea lateral oblicua e\'idente \' que se extiende desde la orbita ocular hasta la ingle C. argywgaster Li'nea lateral oblicua difusa v restringuida a la ingle C. cmspcdoccps 11. Linea dorsolateral presente 12 Linea dorsolateral ausente 15 12. Discos de los dedos manuales muy expandidos .. 13 Discos de los dedos manuales no expandidos o ap- enas expandidos 14 13. Linea lateral oblicua present C. clciithi'iviiiutylus Linea lateral oblicua ausente C. spilotogaster 14. Pliegues laterales presentes a lo largo de los dedos manuales C. si/lvaticiis Dedos manuales sin pliegues laterales... C. idiomcliis 1 5. Linea lateral oblicua ex'idente I'lnicamente en la parte posterior del cuerpo; dedos de los pies con membra- nas interdigitales que se extienden hasta aproxima- damente 2/3 de su longitud C. sordiddfiis Li'nea lateral oblicua se extiende desde la orbita ocu- lar hasta la ingle; dedos de los pies con membranas interdigitales que no se extienden mas alia de 1 /4 de su longitud 16 16. Dedos manuales con pliegues laterales; dedos de los pies con membranas interdigitales presentes por lo menos basalmente C. clacln/hii^tiis Dedos manuales sin pliegues laterales; dedos de los pies sin membranas interdigitales C. poccilojiotus ACCOUNTS OF THE SPECIES Colostcthiis aeruginosiis new species Holotype. — KU 211940, an adult female from the east slope of Abra Pardo de Miguel (05"46' S, 77"42' W), 2180 m, Provincia de Rioja, Departamento deSan Martin, Peru; one of a series collected by Fernando Cuadros, Michael E. Morrison, and John ]. Wiens on 31 [anuarv and 01 Februarv 1989. Paratypes.— KU 211941-43 and MHNSM 6225-27 with same data as holotvpe; KU 211945 from the east slope of Abra Pardo de Miguel, 1980 m, Provincia de Rioja, Depar- tamento, San Marti'n, Peru. Diagnosis. — A moderately large species of Colofitcthus with the following characteristics: (1) maximum SVL of males 25.0 mm, of females 29.2 mm; (2) disc on Finger III 12 Scientific Papers. Natural History Museum. The Univhrsit'i of Kansas (7) outer tarsal fold and tarsal tubercle absent; inner tarsal fold curved, distinct on distal half of tarsus; (8) webbing absent between toes; (9) dorsolateral stripe absent; (10) oblique lateral stripe absent; (11) ventrolateral stripe ab- sent; (12) gular-chest region brown with cream spots; (13) posterior part of abdomen mottled or cream; (14) sexual dimorphism in ventral coloration; (15) median lingual process absent; (16) Finger 111 not swollen in males; (17) testes white. The only other species in Ecuador and Peru that has Finger 1 longer than Finger 11 and lacks longitudinal stripes is Colostethus inittenrieieri; this species differs from C. aeni- ginosiis by having the toes about one-fourth webbed and having a cream venter with a pair of brown spots on the throat. Description . — (.V = 8: 5 males, 1 female, 2 juveniles). Body moderately robust; head slightlv wider than long; head length 32.4-37.1% (x = 35.6':;,) of SVL; head width 36.3-39.0',', (x = 37.7'y,) of SVL; snout moderately long, bluntly rounded in dorsal and lateral profiles; loreal region slightly concave; nostrils barely protuberant, anterior bor- der at level slighth' posterior to anterior margin of lower jaw; eye-nostril distance 72.4-78.6':?< (x = 74.5%) of length of eye; supratympanic bulge weak', covering upper edge of tvmpanum; t\'mpanic annulus distincth' elevated; length of tympanum 51.7-62.9% (x = 58.9','(',) length of eye, separated from eye by distance about one-fifth length of eye. Forelimb moderatelv long, slender; Finger 1 noticeably longer than Finger 11; fingers unwebbed, bearing broad lat- eral fringes; Finger 111 not swollen in males; terminal discs expanded, width about half again width of penultimate phalanges; subarticular tubercles rounded; supernumer- ary tubercles absent; palmar tubercle large, round; thenar tubercle small, elliptical; nuptial excrescences absent. Hind limb short, robust; tibia length 42.8-44.7'./, (\ = 44.0';,) SVL; foot length 41.1-42.6% (x = 41.9%) SVL; outer tarsal fold and tarsal tubercle absent; inner tarsal fold tlaplike, form- ing shallow sigmoid on distal half of tarsus; inner meta- tarsal tubercle small, elliptical; outer metatarsal tubercle small, subconical; toes unwebbed, bearing broad lateral fringes; terminal discs expanded, width about half again width of penultimate phalanges; subarticular tubercles small, rounded; supernumerary tubercles absent. Skin on dorsum of head, body, and hind limbs smooth with scattered tubercles posteriorly on dorsum and on hind limbs; skin on flanks and ventral surfaces smooth; cloacal opening directed posteroventrally at midlevel of thighs; cloacal sheath short. Tongue long, widest and shallowly notched posteriorly, free behind for about one-third of its length; median lingual process absent. Color in preservative: Pale longitudinal stripes absent. In females, dorsum of head and bod\' black with faint tan flecks on head; flanks black with cream spots; upper lip cream with faint gray markings; dorsal surfaces of limbs black with indistinct tan spots; anterior and posterior surfaces of thighs brov\'nish black with irregular, small cream spots; throat, chest, anterior two-thirds of belly, and ventral surfaces of limbs black with cream spots (Fig. 3); posterior one-third of bellv and proximal surfaces of thighs cream; palmar and plantar surfaces dark gray. In males, dorsum dull brown with faint irregular darker brown markings; limbs tan with dark brown transverse bars; throat and chest dark brov\'n with cream spots; belly and ventral surface of limbs cream. Color in life Female holotvpe. — Dorsum dull green with faint bluish tint; bluish cream flecks on flanks; dorsal surfaces of digital scutes white, contrasting with bordering gray surfaces; throat chest, and anterior part of belly olive brown with cream spots; posterior part of belly and ventral surfaces of thighs dark yellow; iris brown with cream ring around pupil (Fig. 5A). Measurements of holotype (in mm): S\'L 29.2, tibia length 14.0, foot length 13.5, head width 10.4, head length 9.5, eye-nostril distance 2.6, length of eve 3.5, length of tympanum 2.1. Distribution and ecology. — This species is kncn\n from two localities at elevations of 1980 and 2180 m along the road from Abra Pardo de Miguel to Moyobamba on the steep east slope of the northern part of the Cordillera Central in northern Peru (Fig. 4). The frogs were associated with small streams in cloud forest; all VNere under rocks at the edges of streams by day. 'I use the term "bulge" instead of "fold." According to Grant (2004), the bulge is formed by a slip of the m. dcpri'^Mr mandihulae and is not a dermal modification as is the fold of skin usually referred to as a supra- tympanic fold. Fig. 3. Ci'/i's(i'f/;i(S ncnif;i>w:ius, KU 21 1440, female venter, 29.2 mm S\'l.. Coi-osii-THCs OF Northern Phru 13 4° > 7y' ■' M^l^r" ' A ;' I'" 11' 4' <; l(XMI 111 _ 1000-3000 m _: 3000-1000 m -^ 1000 -JOOO 111 so KHi oc. • c aeruginosus insulatus 5° -X AC ■ c ornatus sordtdatus 5° V '%'" s -AJ. 6° ^ \ 'O) V --. V^.--^, " \ 6' \ >•: _J 4. ■ 7° \ \ 74" 7K' 77° Fig. 4. Localities of occurrence ot four species of Culostctlius. Etymology. — The specific name is Latin meaning full of verdigris and refers to the dull bluish green ciorsal col- oration. Remarks. — At least some of the apparent sexual differ- ences in coloration may be related to size. Three juveniles (KU 211945, MHNSM 6227) with SVLs of 20.5 and 18.8 mm have uniformly cream venters and bars on the limbs that are more distinct than those in males ha\ing SVLs of 22.1-25.0 mm. Colostcthiis argyrognstcr Morales and Schulte Colostt'tluis (ngiiivf;a>lL-r Morales and Schulte, 1993:98. Holotype: MHNSM 14256 from Imaza, Provincia Bagua Grande, Departamento de Amazonas, Peru. Diagnosis. — A small species of Coloiitctliiif^ with the following characteristics: (1) SVL in males 17.5-19.8 mm, in females 19.0-22.1 mm; (2) disc on Finger 111 expanded; (3) Finger I longer than Finger II; (4) lateral fringes absent on fingers; (5) disc on Toe IV expanded; (6) lateral fringes absent on toes; (7) outer tarsal fold absent; (8) webbing basal between Toes III and IV, alisent between other toes; (9) dorsolateral stripe absent; (10) oblique lateral stripe present; (11) ventrolateral stripe present; (12) gular-chest region uniform cream; (13) abdomen bright silver; (14) males with orange inguinal region in life; (15) median lingual process absent; (16) Finger III not swollen in males; (17) testes white. Few species of Colostcthiifi in Ecuador and Peru lack lateral fringes on the digits and ha\'e Finger 1 longer than Finger II. Of these, only C. ar^i/ivgiistcr and C. oiiiatiis have ventrolateral stripes; the latter differs from C. nri^i/rogaster by having dorsolateral stripes and by having a lemon yellow belly instead of a silver-colored venter; likewise, C. utcubambeiisis has dorsolateral stripes. Colostiilnis iiiit- termcieri differs by lacking all longitudinal stripes, and C. cmspcdoccps differs by having net webbing between the toes and a diffuse (instead of distinct) oblique lateral stripe. Description. — (N = 5: 2 males, 3 females) Body moderately slender; head longer than wide; head length 33.9-38.5% (x = 36.6) of SVL; head width 29.0-32.3% (x = 31.1) of SVL; snout long, bluntly rounded in dorsal view, acutely rounded in profile; loreal region flat; nostrils not protuberant, anterior edge slightly posterior to anterior end of lower jaw; eye-nostril distance 63.3-75.9% (x = 69.3%) eye; supratympanic bulge diffuse, obscuring posterodorsal part of tympanum; length of tympanum 44.8-59.1' o (x = 50.1) length of eye, separated from eye by distance about one-tenth length of eye. Forelimb long, slender; Finger I longer than Finger II; fingers unwebbed, lacking lateral fringes; Finger III not swollen in males (Morales and Schulte, 1994); terminal discs expianded, about half again width of penultimate phalanges; subarticular tubercles rounded; supernumerary tubercles absent; palmar tubercle moderately small, round, partially bifid; thenar tubercle large, elliptical; presence of nuptial excrescences absent. Hind limb moderately short, robust; tibia length 41.6^6.0% (x = 44.4"o) of SVL; foot length 40.3-42.1'v, (x = 41.2%) of SVL; outer tarsal fold ab- sent; inner tarsal fold barely discernible, continuous with small tarsal tubercle on distal one-third of tarsus; inner metatarsal tubercle small, ovoid; outer metatarsal tubercle small, subconical; toes unwebbed, lacking lateral fringes; terminal discs slightly expanded, about half again width of digits; subarticular tubercles small, rounded; supernumer- ary tubercles absent. Skin on all surfaces smooth, with a few low, scattered tubercles posteriorly on dorsum of body; cloacal opening directed posteroventrally at upper level of thighs; cloacal sheath short. Tongue cordiform, shallowly notched pos- teriorly, free behind for about one-half its length; median lingual process absent. Color in preservative: Dorsum of head and body brown without darker markings; flanks uniform dark brown, color confinuing anteriorly as brown stripe through upper two thirds of tympanum, loreal region, and around tip of snout; dorsolateral stripe absent; oblique lateral stripe pale tan, extending from posterior corner of orbit to groin; creamy white labial stripe through lower one third 14 SciiiNTihic Papers, Natural History Museum, The University of Kansas ir- tU'f ttsii}io>u>. KV 2]1'-'41), tomali' 2'' 1 mm b\ L B. Colostctluii 211952, female, 19.5 mm SVL. c. L,i;. kl 212 i2(l, li'iiuk', 21, T mm s\ I D. I i';i'-,/r;/;iis m^iiLili,-. KL 21 IS33, temalc, 22.5 mm SVL. G. Coloatetliiit^ idiomelu^, KU 2I18S4, female, 2.3.0 mm SVL. H. Cohsti'lhiis idioii!chi>. KU 21 ISS4, lemdle, 23.1' mm SVL. Fig. 5. Six species of Colo^lclliu> friim llie Ande^ ol norlhern Peru. Color [iholos on .AmphlbiaVVeb (htlp; / /elib. cs.berkeley.edu /aw/ COLOSTETHUS OF NORTHERN PhRU 15 B. Colo>tiiliu< /.-IK, >;''""'"". KU 211879, female, 2(i.l mm S\ L. C. Gi/os(c//;i/s iicxipiiii, KU 211805, female, 22.8 mm SVL. D. Colo^tcUui^ oiu,itii<, KU 211M51, tenuis I " 1 mm SVL E. Co/o.s/cf/dL- fmUiu'niiiiii>. KU 211946, female, 28.0 mm SVL. F. Cotostcthus sylvaticus, KU 181667, female, 28.7 mm SVL / ^**^!r*^ ! : G. Coloslethus sorduiatus, KU 211860, female, 33.0 mm SVL. H. Co/()S(e//!wssorrf;.i.(;i.,. KL 211861, female, 36.1 mm SVL. Fig. 6. Seven species of Colo'^tcthiis from the Andes of northern Ecuador. Color photos on AmphibiaWeb (http:/ ;'elib.cs.berkcley.cdu/aw/ 16 Scientific Papers. Natlrai, History Mushi'm. Thf University of Kansas Fig. 7. Colostethus argyrogaster, MHNSM 14257, temale, 22.1 mm SVL. of tympanum and continiuius with ventrolateral stripe extending to groin; foreiimbs tan; dorsal surfaces of hind limbs tan; anterior surfaces of thighs tan with dark brown longitudinal stripe; posterior surfaces of thighs brown with longitudinal dark brov\'n stripe. Venter uniform creamy white; palmar surfaces tan; plantar surfaces brown (Fig. 8). Color in life: "Dorsum pale brown with granules on posterior part of body dark brown; tlanks dark brown; oblicjue lateral line brilliant silver; dorsal surfaces of thighs cream; posterior surfaces of thighs and inner surfaces of shanks orange; inguinal region diffuse orange; white mark on either side of cloaca extending onto posterior surface of thigh; throat and chest cream; abdomen bright silver; iris bronze" (translation from Spanish in Morales and Schulte, 1994:100). Sexual dimorphism in coloration is the orange inguinal region in living males. Distribution and ecology. — This small species is known from two localities in the Ri'o Cainarachi drain- age east of the Cordillera Central and from two localities in the middle Ri'o Maranon near the northern end of the Cordillera Central (Fig. 8). These localities are at elevations of 500-650 m in partially cutover lowland humid triipical forest and dry forest. Remarks. — The foregoing description is based on three paratypes (MHNSM 1808-09 and 14257) and two additional specimens (KU 196717 and LSUMNS 32523). Colostethus craspedoceps new species Holotype.— KU 211952, an adult female, from 30 km southwest of Zapatero (about 10 km NE San Jose de Sisa [ca. 06°45' S, 76"33' W], 500 m, Provincia de Lamas, Depar- tamento de San Marti'n, Peru; one of a series collected by William F. Ducllman and Rainer Schulte on 13 February 1989. Paratypes.— KU 211953-59 collected with the ln>lo- type. 3f, 79° / -J.^ - 77' 3° ,; < 1000 m 1000-2000 m .-S i \ CZZ 2000-3000 m \ ■ ma 3000-4000 m '~i. ^m >«X»m / ' 50 lOO a" r 1 Kdometef^ A C. argyrogaster 4° / ▼ C. craspedoceps ! / ^. ■ C, eleutherodaclylus * C sylvatlcus V 1 v4 \ ■'• n A \ 'r\ y- h- { \ -^ 6° \ ^ r^ ^ --•' ' ^ \ 7" .-,/" \, r \ V \ - 79 78° 11° Fig. 8. Localities of occurrence of four species of Cokvtcthuf. Diagnosis. — A small species of Colostethus with the following characteristics: (1) maximum S\'L of males 19.1 mm, oi females 20.5 mm; (2) disc on Finger 111 expanded, about half again width of penultimate phalange; (3) Finger I longer than Finger II, (4) lateral fringes absent on fingers; (5) disc on Toe IV expanded, about twice width of penultimate phalange; (6) lateral fringes absent on toes; (7) outer and inner tarsal folds absent; low, indistinct outer tarsal tubercle present; (8) webbing absent between toes; (9) dorsolateral stripe present; ( 1 0) oblique lateral stripe diffuse, present in groin only; (11) \'entrolateral stripe absent; (12) gular-chest region cream; (13) abdomen cream; (14) sexual dimorphism in \entral coloration; (15) median lingual process absent; (lb) Finger 111 not swollen in males; (17) testes white. TweKe other species in Fcuador, Peru, and Amazonian Brazil resemble Ci>lo>ti'tlnis cnisi'i'doicps in having Finger I longer than Finger II and in having dorsolateral stripes, but no obli(.|ue lateral stripe.' One of these, C talatuaticne, is slightK- larger (maxinuim S\'L 24 mm), has moderate webbing between the toes and a \entrolateral stripe, and is restricted to the Chocoan Region. 1 he other 11 species are ■Most specimens of Colostethus triUneatus have a pale streak anterior to the groin (Grant and Rodriguez, 2001 ); C. mindu'sianus has a diffuse pale line or series of diffuse pale spots in the groin or extending anteriorly to midflank or ne.ir to the iiisi'rtion of the arm (Caldwell el al., 2002). COLOSTETHUS OF NORTHERN PeRU 17 Amazonian and belong to the Colo^tcthus triliiicntiiti Group as defined by Morales (2t)tl2). All of these have ventrolateral stripes; seven of the species (C. fii^ccllu^, ^'^ascoiii, iiiairhcsia- iiiis, oruntus, fiUDiptiiofiU'^, tiHincntiis, imiizoliiiiiis) also differ from C. crnspetioccps by having Finger 111 swollen in males. Six species (C. coii^piciiiif, ^asconi, iiuirclic^iaiiiis, innsuiger, sh/h;'^;/os(/s, I'tvizoliniin^) also differ by having lateral fringes on the toes (also on fingers in C. vivizoliiuiifi). Five species (C. in^pcmtiis, onmtus, siiiuptiiosus, hiUucatii^, I'lvizoliiiius) are distinguished by having webbing between Toes ill and IV. Description. — (N = 8: 1 male, 5 females, 2 juveniles). Body robust; head slightly longer than wide; head length 35.6-38.5% (x = 37.8%) of SVL; head width 33.2-35.9% (x = 34.4%) of SVL; snout moderately long, bluntly rounded in dorsal view, rounded above and inclined posteroventrally in profile; loreal region flat; nostrils noticeably protuberant laterally, anterior margins at level of anterior margin of lower jaw; eye-nostril distance 68.0-76.9%) (x = 72.2'y;) eye; supratympanic bulge diffuse, covering posterodorsal edge of tympanum; otherwise, tympanic annulus distinct; length of tympanum 48.0-63.0% (x = 55.3%) length of eye, separated from eye by distance about one-eighth length of eye. Forelimb short, moderately robust; Finger 1 distinctly longer than Finger II; fingers not webbed, lacking lateral fringes; Finger III not swollen in male; terminal discs ex- panded, about half again width of penultimate phalanges; subarticular tubercles rounded; supernumerary tubercles absent; palmar tubercle large, round; thenar tubercle small, elliptical; nuptial excrescences absent. Flind limb short, robust; tibia length 43.5-45.3% (x = 44.5%) SVL; foot length 41 .4-42.9% (x = 42.2%,) SVL; tarsal folds absent; outer tarsal tubercle low, indistinct; inner metatarsal tubercle elliptical- outer metatarsal ttibercle rounded; toes unwebbed; lateral fringes absent on toes; terminal discs expanded, about twice width of penultimate phalanges; subarticular tubercles small, subconical; supernumerary tubercles absent. Skin on dorsum of body and hind limbs, and venter smooth; skin on flanks weakly granular; cloacal opening directed ventrally at midlevel of thighs; cloacal sheath long. Tongue long, widest and shallowly notched posteriorly, free behind for about one-half of its length; median lingual process absent; vomerine odontophores absent. Color in preservative: Dorsum of head and body brown with or without darker brown dashes middorsally; flanks brown; dorsolateral stripe pale tan; stripe narrow and distinct across tip of snout, along canthus rostralis and outer edge of eyelid, becoming wider and somewhat diffuse on body and continuous to supracloacal region, bordered below for its entire length by wider dark brown stripe; oblique lateral stripe short, diffuse, restricted to groin; ventrolateral stripes absent; upper lip creamy tan. Forelimbs tan with irregular brown marks on forearm and longitudinal brown stripe on anterior edge of upper arm; dorsal surfaces of hind limbs pale brown with irregular darker brown markings; anterior and posterior surfaces of thighs pale brown with diffuse darker brown longitudi- nal stripes, fragmented on posterior surfaces of thighs in some indiviciuals. Venter cream, with male having gray- ish-brown throat and chest; palmar and plantar surfaces brown. Coloration in life: Dorsum brown with yellowish-tan dorsolateral stripes, distinct on head, diffuse on body, bordered below by dark brown; side of head below brown stripe ancl dorsal surfaces of upper arms dull yellow; dor- sal surfaces of digital scutes gray; flanks creamy tan with irregular brown spots (Fig. 5B); bellv creamy yellow with greenish tint; ventral surfaces of limbs dull yellow; throat and chest of male brown; iris coppery bronze with median horizontal dark brown streak. Measurements of holotype (in mm): SVL 19.5, tibia length 8.8, foot length 8.3, head width 7.0, head length 7.7, eye-nostril distance 2.0, length of eye 2.7, length of tympanum L9. Tadpoles. — Tadpoles associated with this species are from the type locality and include seven back-riding tad- poles and eight free-swimming tadpoles from a pool in a stream in a rocky ravine. One male with a SVL of 19.1 mm was transporting seven tadpoles on its back. Back-riding tadpoles have body lengths of 3.7-4.8 mm (x = 4.36) and total lengths of 11.0-12.9 mm (x = 11.91). These tadpoles have jaw sheaths and a LTRFof 2(l)/2. In life, the tadpoles were dark brown. In preservative, the body is dark brown dorsally, becoming cream ventrally; the caudal muscu- lature is cream with brown reticulations, and scattered guanophores are present on the body, caudal musculature, anci translucent caudal fins. Five of the free-swimming tadpoles are in Stage 25 and have bocly lengths of 6.5-9.8 mm (x = 8.72) and total lengths of 15.9-23.7 mm (x = 20.68). Three tadpoles in Stages 31, 34, and 35, have body lengths of 10.0, 10.0, 11.5 mm, and total lengths of 24.5, 25.3, and 27.0 mm, respectively. A typical tadpole (KU 215610) in Stage 34 has a body length of 10.0 mm and a total length of 25.3 mm; body ovoid, wider (5.5 mm) than high (4.5 mm); snout rounded in dorsal view and in profile; nares directed dorsolaterally at point midway between tip of snout and orbits; eyes moderately large (1.2 mm), situated dorsally, directed dorsolaterally, not visible from below; interorbital dis- tance 2.1 mm; spiracle sinistral; tube short, attached to body throughout its length; spiracular opening directed posteriorly just below midline at about midlength of body; cloacal tube dextral, short, cone-shaped, attached to ventral fin. Caudal musculature robust, equal height throughout proximal third of tail, gradually diminishing to pointed tip; dorsal fin originating on caudal musculature, highest at about two-thirds length of tail, gradually diminishing 18 Scientific Papers, Natural History Museum. Thii University oi- Kansas f C ^ D »* ?::i;"!»»^ ■^ //- H Fig. 9. Tadpoles of eight species of Colostlictliiis from the Andes of northern Peru. A. C. cmspedoccps, KU 215610, Stage 34, 23.3 mm total length. B. C. clachyhi^tiiii, KU 219754, Stage .34, 32.5 mm total length. C. C. hiiomcla^. KU 215607, Stage 34, .34.7 mm total length. D. C. in^uktus, KU 215600, Stage 34, 38.3 mm total length. E. C. Icucoplumi^, KU 215603, St. 28, 42.5 mm total length. F. C. iwxipuf, KU 215594, Stage 37, 30.5 mm 'total length. G. C. wrdidatns, KU 215611, Stage 25, 30 mm total length. H. C. t.ylvatinis, KU 181869, Stage 33, 29.4 nim total length. Scale bars = 5 mm. to acutely rounded tip; ventral fin originating on body, highest at about three-fifths length of tail; at midlength of tail; heights of dorsal and ventral fins about equal, less than height of caudal musculature (Fig. 9A). Width of oral disc 2.8 mm, directed anteroventrally; median half of anterior labium bare; elsewhere, labia bearing single row of moderately long, pointed marginal papillae; labia having shallow lateral folds; submarginal papillae absent. Jaw sheaths slender, finely serrate; anterior sheath in form of broad arch; posterior sheath broadly V- shaped; LTRF 2(l)/3; A, slightly longer than others. In life, body and tail gray. In preservative, dorsum and sides of body brown; belly cream; caudal musculature pale creamy tan with small, irregular brown spots; caudal musculature translucent with brov\n spots; guanophores scattered on body and tail (Fig. 9A). Distribution and ecology. — This species is know n only from the type locality at the eastern base of the northern part of Cortiillera Central (Fig. 8); this is a stream in a shal- dorsolateral stripe absent; (10) oblique lateral stripe pres- low ravine in cutovcr humid tropical forest. The frogs were ent; (11) ventrolateral stripe absent; ( 1 2) gular-chest region active on wet boulders in the stream bv dav. Tadpoles of with pair of dark marks; (1,3) abdomen with white spots; Colostftlnis soriiidiitii> were found in the same stream as tadpoles of C. cra^pcdoccps. Etymology. — The specific name is an adjective derived from the Creek kinspciloii meaning edge or border and the CJreek kcphnlc meaning head. The name refers to the pale stripe borclering the anterior and lateral margins of the head. Colostetlins elachyhistiis Edwards Colo^telhu^ elndn/hi^itti^ Edwards, 1971:149. Holotvpe: KU 12().S4() from Loja, 21. "SO m, Provincia de Loja, Ecuador. Diagnosis. — A moderately small species of Colostetlnifi with the following characteristics: (1) maximum SVL of males 24.1 mm, of females 24.8 mm; (2) disc on Finger 111 not, or only slightly, expanded; (3) Fingers 1 and II equal in lengtii; (4) lateral fringes present on fingers; (5) disc on Toe IV slightly expanded; (6) lateral fringes present on toes; (7) outer tarsal fold weak or absent; inner tarsal fold curved on distal third of tarsus; (8) toes about one-third webbed; (9) COWSTETHVS OF NORTHERN PkRU 19 (14) no conspicuous sexual dimorphism in coloration; (15) median lingual process absent; (Id) f'inger 111 not swollen in males; (17) testes white. Colosk'thus cliuin/histiis resembles C. awn, iiifrn^uttatiis, and z'ciichmlis in having an oblique lateral stripe and pair oi dark marks on the gular-chest region. It ditters from C. vcitcbialis in ventral coloration and bv lacking a middorsal stripe, from C. inoa bv having a well-detined, continuous oblique lateral stripe (diffuse and fragmented in C. irwii), and from C. i}ifn\':iiittntii^ by having more extensive web- bing (toes webbed basally in C. iiifn\^iittntiis). Colostctlius eleutlwivdnctiflit^, C. ^oniidatiit^, and C. sylzmticiis also ha\'e Fingers I and II equal in length, but these species lack webbing between the toes, and C. clciithcrodactyliis and C. si/lvaticiis have dorsolateral stripes. Description . — (N = 36: 20 males, 16 females). Body moderately robust; SVL in males 16.7-19.8 (x = 18.4) mm, in females 18.6-23.4 (x = 20.4) mm; head slightly longer than wide; head length 32.6-40.r'o (x = 36.4%)'of SVL; head width 30.9-37.6','- (x = 34.6'';<) of SVL; snout short, bluntly rounded in dorsal view, rounded in lateral profile; loreal region flat; nostrils slightly protuberant laterally, anterior border; eve-nostril distance 50.0-68.0',7- (x = 57.7%) eye length; supratympanic bulge low, diffuse, covering posterodorsal part of tvmpanum; length of tympanum 48.3-62.1' ,'i (x = 54.5"n) length of eye, separated from eye by distance about one-tenth length of eye. Forelimb moderately short, slender; Fingers 1 and II equal in length; fingers unwebbed, bearing narrow lateral fringes; Finger III not swollen in males; terminal discs not or slightly expanded; subarticular tubercles subconical; supernumerary tubercles absent; palmar tubercle large, round; thenar tubercle elliptical; nuptial excrescences ab- sent. Hind limb short, robust; foot longer than tibia; tibia length 42.9-50.6';;, (x = 47.1'7.) SVL; foot length 44.4-52.2"v, (x = 48.9'o) SVL; outer tarsal fold weak or absent; inner tarsal fold conspicuous, curved on distal third of tarsus; inner metatarsal tubercle ovoid; outer metatarsal tubercle subconical; toes at least partially webbed, usually about one-third webbed; webbing formula I(r-2) — (2-2')II(2 -2')— (3-3-)III(2'-3-)— (3 4)IV(3 -4)— (2*-3-)V; toes with narrow lateral fringes; terminal discs slightly expanded, no more than half again width of penultimate phalanges; subarticular tubercles small, round; supernumerary tu- bercles absent. Skin on all surfaces smooth; small, scattered tubercles present postsacrally in some specimens; cloacal opening directed posteriorly at upper level of thighs; cloacal sheath short. Tongue half again as long as wide, widest and dis- tinctly notched posteriorly, free behind for about two-thirds of its length; median lingual process absent. Color in preservative: Dorsum of head and body brown with darker brown irregular spots; flanks brown with small white spots ventrallv in some specimens; dark brown stripe continuous trom nostrils, below canthus, along supratympanic area to flanks; dorsolateral stripe absent; oblique lateral stripe creamy tan, extending from grciin to edge of upper evelid or just posterior to eye, con- tinuous with irregular edges on anterior surface of thigh; ventrolateral stripe absent (except in one female). Upper arm creamv tan with brown stripe on anterior surface; fore- arm creamy tan with cir without irregular brown cross bars; dorsal surfaces of hind limbs tan with two or three dark brown transverse bars extending onto posterior surfaces of thighs; upper lip dull tan. Venter tan with or without pair of brown spots posterolateralK' on throat; some females with gray mcittling enclosing dull \\ bite spots on abdomen; one female with small brown spots on throat and abdomen; palmar and plantar surfaces tan. Color in life: Dorsum variable from pale olive to brown; oblique lateral stripe pale yellow to tan, usu- ally narrowly bordered bv dark brov\'n or black (Fig. 5C). Specimens from Huancabamba. — Dorsum brown with dark brown or black markings; flanks dull brown; oblique lateral stripe dull creamy tan; abdomen creamv yellow with gray mottling; throat creamy wliite with gray suffusion; iris grayish brown. Specimens from Canchaque. — Dorsum yellowish olive with black markings; oblique lateral stripe silvery white; venter white; iris reddish brown. Tadpoles. — Six adults were transporting tadpoles in February and March. A female (KU 212514) having a SVL of 24.5 mm was carrying five tadpoles (KU 212488) at Aya- baca, Departamento de Piura, Peru, on 21 February 1989; other adults transporting tadpoles are males from Abra de Porculla, Departamento de Piura, and 1 km S Cajabama, Departamento de Cajamarca. Size and development of keratinized structures in the oral disc vary (Table 3); the smallest individuals lack differentiated intestines, and the body cavity is filled with yolk. Table 3. Moasurements (in mm) .ind laliuil tooth row formul.i (LTRF) of transporting adults and hack-riding tadpoles of Colosliihii^ cliicln/liistus. Range of variation followed on next line hy mean. Adult Bodv Total S\L Se\ ;V length length LTRF 24.5 Male S 3.7-3.8 3.75 10.4-10.6 10.53 2/1-2/2 21.8 Female 12 3.4-3.7 3.58 10.0-10.4 10.14 1/0-1/1 18.7 Female 11 3.3-3.5 3.38 9.8-10.1 9.97 0/0-1/0 18.3 1 emale '' 6.5-7.4 15.4-17.4 16.68 2/2 18.3 lemale h 3.6-3.7 3.68 9.9-10.3 10.1 1 / 1 17.8 Female 14 3.3-3.5 3.40 10.2-10.4 10.26 1/0-1/2 20 Scientific Paphrs. N.vn rai. History Museum. Tin Univi-rsity oe Kansas Table 4. Measurements (in mm) of free-swimming tadpoles of Co/cish'/Ziiis elachyhistus from the vicinit)- of Canchaque, Oepartamento de I'iura, Peru. Range of variation followed bv mean in parentheses. Body Total Stage ,\' length length 2? 2 7.8-90 (X = 8.4) 20.2-22.4 (x = 21.20) 26 3 5.1-7.0 (x = 6.26) 15.0-18.5 (x = 16.62) 27 7 7.4-8.5 (X = 7.98) 19.2-27.2 (x = 22.29) 28 3 8.1-8.9 (x = 8.63) 22.1-26.3 (x = 24.43) 29 1 9.3 22.6 30 2 8.9 20.6-25.9 (x = 23.25) 32 1 9.6 29.0 33 3 10.3-13.3 (x= 11. ,30) 30.5-38.2 (x = 33.41) 34 5 10.1-12.2 (x = 11.28) 31 .3-33.8 (x = 33.74) 35 4 10.9-12.3 (x = 11.53) 32.7-36.6 (x = 34.58) 36 3 10.6-12.5 (x= 11.27) 31 .8-38.2 (x = 35.87) 39 2 10.5-11.7 (x= 11.10) 36.2-36.5 (x = 36.35) 40 1 12.6 38.2 43 I n.h 35.1 Free-swimming tadpoles have been found in pools in streams in January-March. Series of tadpoles from the vi- cinity of Canchaque, Departamento de Piura, are in Stages 25-43 (Table 4). A typical tadpole in Stage 34 (KU 219754) has a body length of 10.8 mm and a total length of 32.5 mm; body ovoid, much wider (7.2 mm) than high (4.7 mm); snout bluntly rounded in dorsal view and in profile; nares directed anterolaterally at point about midway between tip of snout and orbits; eyes large (1.5 mm), situated dorsally, directed dorsolaterally, not visible from below; interorbital distance 1 .9 mm; spiracle sinistral; tube short, attached to body throughout its length; spiracular opening directed posterodorsally well below midline at about midlength of body; cloacal tube dextral, short, attached to ventral fin. Caudal musculature robust, equal height throughout proximal third of tail, gradually diminishing to pointed tip; dorsal fin originating on caudal musculature, highest at about three-fourths length of tail, gradually diminish- ing to acutely rounded tip; ventral fin originating on body, highest at about two thirds length of tail; at midlength of tail, height of dorsal fin greater than that of ventral fin, much less than height of caudal musculature (Fig. 9B). Width of oral disc 3.0 mm, directed anteroxentrally; median half of anterior labium bare; elsewhere labia bear single row of moderately long, pointed marginal papillae; labia having shallow lateral folds; submarginal papillae absent. Jaw sheaths moderately robust, coarsely serrate; anterior sheath in form of broad arch; posterior sheath broadly V-shaped; ITRF 2(l)/3; A, slightlv longer than others. in life, body olive-gray; tail yellow proximally, becom- ing more orange distallv, flecked with gray. In preservative, dorsum and sides of body brown; belly translucent gray; caudal musculature pale creamy tan with brown flecks; cau- dal musculature translucent with brown flecks (Fig. 9B). Distribution and ecology. — As presenth' recognized, this species has the largest geographic and ele\ational distribution of all Cc/hs/c/Z/hs in northern Peru (Fig. 10). It ranges from the vicinity of San Lucas, Ecuador, to Caja- bamba, Peru, an airline distance of about 425 km. The elevational range in northern Peru is 710-276t) m; the spe- cies occurs in thorn forest, drv forest, montane drv forest, and hmnid montane forest. On the western slopes of the Cordillera de Huancabamba, it occurs sympatrically with C. si/lvaticiis at elevations of 1920-2590 m. At Cutervo, Departamento de Cajamarca, it occurs sympatrically with C. pulclh'nisiiini>. In cultivated areas, such as the \icinit\' of Huancabam- ba, Peru, and Macara Ecuador, the frogs inhabit irrigation ditches. At otlier places in thorn forest or dr\- forest, the frogs seem to be restricted to the immediate vicinitv of small streams, a habitat in which thev also are abundant in humid montane forest. Remarks. — The foregoing description is based on spec- imens from the vicinities of Huancabamba and Canchacjue, Departamento de Piura, Peru. Duellman and Wild (1993) noted minor differences in coloration and the amount of 79 7S° .-^'' 77- ' A o " ' r \ ": 7' ,^/' ../ f ^ V ,7S < IflOO Til ;000- .111(10 m .loawooom ; >4<]00m 50 IIKI 4 • C elachyhistus A C. idiomelus ▼ C leucophaeus ■ C poealonotus C- spilotogaster J r f Fig. 10. Localities of occurrence of fi\e species of Co/i'sM/ii/s. COLOSTETHUS OF NORTHERN PeRU 21 webbing behveen specimens from those localities and those trom the Loja Basin in Ecuador. Comparison ot specimens from nine localities in northwestern Peru reveals minor differences in SVL and different proportions of individuals that have a pair of brown spots on the gular-chest region or of females that have a mottled abdomen (Table 5). How- ever, specimens from all of these localities are consistent in having a cream longitudinal stripe on the anterior surface of the thigh and a brown longitudinal on the upper arm. In some individuals from Huancabamba, the toe-webbing formula for Toe IV is 3'; in others it is 4" or 4. At all other Peruvian localities the formula for Toe IV is 4 to 4*. Topotvpic specimens from the vicinity of Loja, Pro\in- cia de Loja, Ecuador, and from other localities in southern Ecuador resemble those from northern Peru in having a cream longitudinal stripe on the anterior surface of the thigh and a brown longitudinal stripe on the upper arm, as well as a toe webbing for Toe IV as 4" to 4*. All have a pair of brown spots on the gular-chest region, and 52-9T'(. of the females in four samples have a mottled abdomen (Table 5). On the average, individuals of both sexes from Peru are smaller than those from Ecuador. Males from Ecuador have SVLs of 16.3-2L5 mm (x = 19.1, N = 28), in contrast to 15.2-20.9 mm (x = 17.7, N = 44) from Peru, whereas females from Ecuador have SVLs of 19.6-25.8 mm (x = 22.6, N = 47), in contrast to 17.2-23.6 (x = 21.1, N = 52) from Peru. Mean snout-vent lengths of females of 13 samples were regressed against elevations of their respective localities; the analysis revealed no significant correlation (P = 0.9452). The variation in this species is further confounded bv differences among tadpoles from separate sites. In contrast to the coloration of tadpoles described here, Duellman and Wild (1993) noted that living tadpoles from Huan- cabamba, Departamento de Piura, Peru, had a dull brov\'n body and a cream tail with a reddish-brown mid-lateral line proximally. Edwards (1971:153) described the color of preser\ed tadpoles from Loja, Provincia de Loja, Ecuador, as "black above, dark gray to black below, becoming pale gray anterior to eyes; papillae and lips impigmented; caudal musculature creamy white with black reticulations concentrated into small blotches anteriorly on dorsal half of tail." Taran Grant (pers. comm., 16 July 2004) found that sequences of Cytochrome b are 13% divergent between the sample from 4 km W of Llama, Departamento de Caja- marca, and those from 8.5 km E Canchaque, Departamento de Piura, and Ayabaca, Departamento de Piura, whereas the divergence is only 2''(. between the last two samples. Careful examination of the specimens from 4 km W Llama (KU 212518-24) reveals no consistent morphological dif- ferences from other populations of Cokvtethus eladn/histiis, a nominal species that may include two or more cryptic biological species. Colostcthtis eleiithcroiiactyliis new species Holotype.— KU 211814, an adult female, from 16 km BSE of Shapaja (ca. 06"37' S, 76"'15' W), 360 m, Provincia de San Martin, Departamento de San Martfn, Peru; obtained on 19 February 1989 by John J. Wiens. Paratype. — KU 211813, an adult male, collected with the holotype by Rainer Schulte. Diagnosis. — A moderately small species of C(i/osft'f/;»s with the folkiwing characteristics: (1) SVL in male 21.0 mm, 22.7 mm in female; (2) disc on Finger III greatly expanded; (3) Fingers I and II equal in length; (4) lateral fringes absent on fingers; (5) disc on Toe IV greatly expanded; (6) lateral fringes absent on toes; (7) outer tarsal fold present; curved Ttilile ?. Geographic variation in Colostctliiif clacln/histii^. Localities arranged from north to south. Locality N Male SVL (mm) Ecuador: San Lucas 7 17.4-21.4 (x = 19.4, ;; = 2) Loja 45 17.2-21.5 (x = 19.4, )/ = 20) Cariamanga 12 16.3-19.5 (x = 18.L" = 6) Macara 11 — Peru; Avabaca 4 20.2, n = \) Huancabamba 28 16.7-19.8 (x = 18.4, (/ = 18) Canchaque 8 16.8-19.3, (x = 18.1,;; = 2) Abra de Porculla 32 15.2-18.7 (x = 16.3, » = 17) Santa Cruz 7 — Cutervo 3 — Lamas 7 16.0, }i =1 Cajabamba g 19.3-20.^) (x = 20.1, )/ = 3) Female SVL (mm) Females with mottled abdomen Pair of brown spots on throat 23.5-25.1 (x = 24.2, ii = 5) 19.6-25.8 (x = 22.8, ii = 25) 20.4-23.0 (x = 21.4, )i = 6) 20.5-23.7 (x = 2L9,;! = 11) 20.5-2L7(x = 21.2, 11 = 3) 20.2-23.4 (x = 21.8, )/ = :0) 18.6-22.5 (x = 20.7,11 = 6) 17.2-21.5 (x = 19.7, (1 = 9) 21.4-22.7 (x = 22.0, n = 7) 18.5-19.2 (x = 18.9, /; = 2) 20.2-22.8 (x = 21.5, )? = 6) 20.9-23.6 (\ = 21.9, (, =6) 60% 52% 67% 91% 33% 0% 67% 67% 33"; 100% 100% 100% 100% 75% 100% 86% 100% 71% 89'; 22 Scientific Papers, Natural History Museum. Tin UMMRsn v of Kansas inner tarsal fold on distal half of tarsus; (8) webbing absent between toes; (9) dorsolateral stripe present; (10) shi)rt oblique lateral stripe present; (11) ventrolateral stripe absent; (12) gular-chest region uniformly creamv white; (13) abdomen creamy white; 14) no sexual dimorphism in coloration; (15) median lingual process absent; (16) Finger ill not swollen in males; (17) testes white. In northern Peru and Ecuador, Colostcthufi cxafiperatu^ is the only other member of the genus that has Fingers I and 11 equal in length and possesses dorsolateral and oblique lateral stripes; it differs from C ck'iithcivdacti/liis by having a swollen black gland on the inner surface of the elbow, having dark marbling laterallv on the belly, and by lacking an outer tarsal fold. Colostcthii^ ^oniidatu::^, C. idioiiwlus, and C. si/lvaticus also have the Fingers I and 11 equal in length; C. >ordidatii> lacks a dorsolateral stripe, and the other two lack oblique lateral stripes. Furthermore, C. idioiiicliis and C. sylvaticiis ha\e only slightly expanded discs on the dig- its, and both have lateral fringes on the fingers and toes. The coloration of C. clciitlicivdncti/lii^ is much like that of C. nexipiis, a species that has Finger 1 shorter than Finger 11, distinct lateral fringes on the digits, and toes about half webbed. The coloration also resembles that of C. patitac in the Serranfa de Sira in central Peru; that species differs from C. I'leutlicivdacti/liis by having Finger 1 shorter than Finger 11, a bright yellow dorsolateral stripe, and dark brown upper lip in contrast to the yellowish tan upper lip in C. elciitheivdncti/liis (Letters et al., 2003). Description. — (N = 2: 1 male, 1 female). Body moder- ately robust; head slightly longer than wide; head length 33.9% and 35.2% (x = 34.6%) of SVL; head width 32.2% and 34.3% (x = 33.3'/o) of SVL; snout long, nearly truncate in dorsal view, bluntly rounded in profile; loreal region barely concave; nostrils protuberant laterally, anterior edges at level of anterior margin of lower jaw; eye-nostril distance 67.7% of eye length; supratympanic bulge diffuse, obscur- ing upper edge of tympanum; length of tympanum 51.6% length of eye in male, 64.5' i/ in female, separated from eye by distance about one-fourth length of eye. Forelimb moderately long, slender; Fingers 1 and II equal in length; fingers unwebbed, lacking lateral fringes; Finger III not swollen in male; terminal discs expanded, nearly truncate, about twice width of penultimate pha- langes; subarticular tubercles prominent, subconical; supernumerary tubercles absent; palmar tubercle large, elevated, nearly round; thenar tubercle elevated, bluntly ovoid; nuptial excrescences absent. Hind limb moderately robust, short; tibia length 45.4% and 47.6% (x = 46.5%) SVL; foot length 43.6% and 45.7% (x = 44.7%) SVL; outer tarsal fold low, thin, extending full length of tarsus; inner tarsal foki low, a shallow sigmoid curve from prominent tarsal tubercle to elliptical inner metatarsal tubercle; outer metatarsal tubercle subconical; toes imwebbed, lacking lateral fringes; terminal discs expanded, about twice width of penultimate phalanges; subarticular tu- bercles prominent, subconical; supernumerar\' tubercles absent. Skin on dorsum of body and hind limbs smooth; skin on flanks finely shagreen; skin of ventral surfaces smooth; cloacal opening directed posteroventrally near upper level of thighs; cloacal sheath short. Tongue elongate, widest posteriorly, not notched behind, free posteriorly for about two-thirds of its length; median lingual process absent. Color in preservative: Dorsiun of head and body brown; flanks dark brown; dark brown stripe across snout, through loreal and tympanic regions, confluent with brown flanks; dorsolateral stripe tan, continuous across snout, canthal region, outer edge of upper eyelid to point above insertion of hind limb; oblique lateral stripe white, extend- ing from groin to midtlank, fragmented; \entrolateral stripe absent; dorsal surfaces of upper arm creamy tan; anterior and posterior surfaces of upper arm brown; forearm tan with brovsn transverse bars; dorsal surfaces of hind limbs brown with dark brtnvn transverse bars; anterior surfaces of thighs brown with diagonal cream mark proximallv; posterior surfaces of thighs brown with tan flecks; upper lip creamy tan. Venter creamy tan with minute brown flecks on throat and chest; palmar surfaces cream\' tan; plantar surfaces brown. Color in lite: Dorsum brown with dark brown mark- ings; dorsolateral stripe tan with orange tint on head; flanks black with broken creamy v\hite oblique lateral stripe; thighs and upper arms pale dull yellow w ith brow n markings; dorsal siufaces of digital scutes white, contrast- ing with adjacent brown surfaces; venter dull yellow; iris pale copper (Fig. 5E). Measurements of holotvpe (in mm): SVL 22.7; tibia length 10.3; foot length 9,9; head width 7.3; head length 7.7; eye-nostril distance 2.1; length of eve 3.1; length ot tympanum 2.0. Tadpoles. — An adult male v\ith a SVL of 21.0 mm was carrying five tadpoles. Five back-riding tadpoles (KU 21 5593) have body lengths of 3.8-4.3 mm (x = 4.04) and total lengths of 10.8-11.2 mm (x = 10.9). The tadpoles are in Stage 25; the intestines do not seem to be fully differentiated, and the body cavity is filled with yolk. All individuals have thin jaw sheaths and variable development of labial tooth rows; fyv'o have LTRF of 1/0, two have 2/1, and one has 2/2. In preservative, the dorsal and lateral surfaces of the body are brown; the caudal musculature is creamy white with faint brown spots above the midline, and the fins are translucent. Distribution and ecology. — This species is known onl\- from the t\ pe localitx at M'i elevation of 360 m in disturbed humid tropical forest just north ot [he Rio Huallaga in the upperniost reaches of the .Xma/iin Basin (Fig. S). Both in- COLOSTETHI'S OF NORTHERN PeRU 23 dividuals were active in leaf litter near a small stream in a rocky ravine bv day. Etymology. — The specific name is a compound adjec- tive derived from the Greek elcutheivs meaning free and the Greek daktylos meaning toe. The name refers to the unwebbed toes of this species. Colostcthiis idioiiiclus Rivero Cohstethus idiomelin Rivero, 1991a:I0. Holotv'pe: MCZ A-100260 troni Venceremos, Departamento de San Martin, Peru, 162(1 m. Diagnosis. — A moderate-sized species of Colostctlius with the following characteristics: (1) ma,ximum SVL of males 24.8 mm, of females 27.8 mm; (2) disc on Finger III not expanded; (3) Fingers I and II equal in length; (4) narrow lateral fringes present on fingers; (5) disc on Toe IV slightly expanded; (6) narrow lateral fringes present on toes; (7) curved inner tarsal fold on distal half of tarsus; (8) webbing absent between toes; (9) dorso- lateral stripe present; (10) oblique lateral stripe absent; (11) ventrolateral stripe absent; (12) gular-chest region with pair of brown spots; (13) abdomen mottled gray and white anteriorly, yellow posteriorly; 14) no sexual dimorphism in ventral coloration; (15) median lingual process absent; (16) Finger 111 not swollen in males; (17) testes white. Three other species in Ecuador and Peru have dorso- lateral stripes and Fingers 1 and II equal in length. Of these, Colostcthiis eleiitheroLiacti/liis and C. exnspi'mtiis also have oblique lateral stripes and expanded discs on the digits (oblique stripe absent and discs on fingers not expanded in C. idioiiifhis). Colostethiis sylvaticiis is most like C. idioiii- clus in having the first and second fingers equal in length, weakly expanded discs on the digits, no oblique lateral stripe, and a pair of dark spots on the throat; however, C. sylimticiis differs from C. idioiiiclus by having more exten- sive lateral fringes on the fingers and toes, much broader dorsolateral stripe, more dark bars on the hind limbs (3 or 4 bars vs. 2), and a yellow throat in life. Also, C. s\/lvaticus is slightly larger (Table 1). Description.— (N = 61: 38 males, 23 females). SVL 19,5-24.8 mm (x = 22.0) in males, 23.0-27.8 mm (x = 25.1) in females; bodv robust; head slightlv longer than wide; head length 33.9-39.9";- (x = 36.6";,) of SVL; head width 32.3-41.4';, (x = 34.4'/;,) of SVL; snout moderately long, rounded in dorsal view, bluntlv rounded in profile; loreal region flat; nostrils barelv protuberant laterally, anterior edge just pcisterior to level of antericir margin of lower jaw; eye-nostril distance 60.0-83.3';^, (x = 70.4'T,) of length of eye; supratympanic bulge distinct, angular posterodorsal to tympanum, obscuring dorsal edge and upper postericir edge of tympanum; length of tympanum 57.1-72.4% (x = 63.3";,) length of eye, separated from eye bv distance about one-fifth length of eye. Forelimb moderately short, robust; Fingers 1 and II equal in length; fingers unwebbed, bearing narrow, keel- like lateral fringes; terminal disc not expanded; subarticular tubercles small, rounded; supernumerary tubercles absent; palmar tubercle large, round; thenar tubercle small, ellip- tical. Hind limb short, robust; foot longer than tibia; tibia length 39.9-52.5% (x = 48.1%) SVL; foot length 40.4-52.0% (x = 47.2%) SVL; outer tarsal fold absent; inner tarsal fold elevated, shallow sigmoid on distal third of tarsus; in- ner metatarsal tubercle small, elliptical; outer metatarsal tubercle subconical, nearly as large as inner tubercle; toes unwebbed, bearing narrow keel-like lateral fringes; termi- nal discs slightly expanded; subarticular tubercles small, rounded; supernumerary tubercles absent. Skin on dorsum smooth to shagreen, usually with scat- tered small tubercles postsacrallv on dorsum; skin on flanks shagreen to finely granular; skin on venter smooth; cloa- ca! opening directed posteriorly at upper level of thighs; cloacal sheath short. Tongue elongate, slightly widest and shallowly indented posteriorly, free behind for about half of its length; median lingual process absent. Color in preservative: Dorsum of head and body dull tan with dark brown markings consisting of (1) irregular interorbital mark, V-shaped marks in occipital and scapular regions, and pair of longitudinal postsacral marks (68.3";,); (2) small interorbital spot or longitudinal mark, pair of diagonal marks in scapular region with small spots posteri- orly, pair of longitudinal postsacral marks (15.0%,); (3) pair of diagonal marks or V-shaped mark in occipital region; rest of dorsum with many small spots (11.7' ;.); dorsum of head and body plain or only with a pair of longitudinal dark postsacral marks (5.0"<,) flanks brown, gray below in some individuals; narrow, dark stripe across snout, through loreal and supratvmpanic regions, to midflank; dorsolat- eral stripe tan, narrow across snout and along outer edge of upper eyelid, usually widest and commonly bordered by black on body; oblique lateral stripe and ventrolateral stripes absent. Forelimb tan with one or two narrow brown transverse bars on dorsal surface of forearm; brown streak proximally on lateral surface of upper arm; dorsal surfaces of hind limbs dull tan with faint transverse brown bars — one or two on thigh, three on shank, one or tw'o on foot; anterior surfaces of thighs dark brown; posterior surfaces of thighs pale brown with \'ertical dark brown extensions of bars on dorsal surfaces. Side of face and upper lip tan; cream stripe on posterior part of upper jaw; venter creamy tan with faint brown mottling anterior to midlength of abdomen; pair of diffuse brown spots posterolaterally on throat; palmar and plantar surfaces brown or tan. Color in life: Dorsum dull tan to rich orange-brown, to grayish tan with green tint in some places; dorsal mark- ings olive-brown to black (Figs. 5F and G); dorsolateral stripe pinkish tan, yellowish orange or cream, usually 24 Scientific Papers. Natural History Muselm. Tin University of Kansas Tdble 6. Measurements (in mm) of transporting males and bacl<-riding tadpoles of Co/os/i'Hiiis idiomflim. Range of variation followed by mean in parentheses). Table 7. Measurements (in mm) of free-swimming tadpoles of Ci'/i)s/('f/n(S ulioiiii'lu^. Range of variation follow ed b\' mean in parentheses. Male SVL Bod\ length total length 20.5 20.7 23.8 H) 4.3-4.4 (x = 4.39) 4.1-1.5 (x = 4.29) 3.7-3.9 (x = 3.7(1) 9.8-10.5 (x = 10.31) I0.2-Il.8(x = 11.04) 10.2-10.4 (\= 10.29) bordered by dark brown; flanks tan to gray; labial stripe pinkish cream; anterior and posterior surfaces of thighs yellow to dull orange; digital scutes gray; iris dull bronze. Throat, chest, and anterior part of abdomen white with dark brown or gray mottling, yellow suffusion on anterior part of throat; posterior part of belly and ventral surfaces of hind limbs yellow; ventral surfaces of forclimbs dull yellow (Fig. 5H). Tadpoles. — Males transporting 7-12 tadpoles each were found in Januarv and February (Table b). The intes- tines of back-riding tadpoles contain yolk. The tadpoles probably grow and begin developing keratinized jaw sheaths and labial teeth while being transported. Thin jaw sheaths are present in some of the smallest tadpoles (body lengths of 3.7-3.8 mm); two tadpoles with body lengths of 4.4 mm have LTRFs of 0/1, one of the same size has an LTRF of 0/2, and one with a body length of 4.5 mm has an LTRF of 0/3. Anterior tooth rows ap- parently develop after the tadpoles are free-swimming, the smallest individual of which as a body length of 6.7 mm and an LTRF of 2/3. Free-swimming tadpoles in various stages of develop- ment were found in January and February in slow-moving, even marshy, streams (Table 7). These are associated with Colostcthus idiomclus because adults of that species were found at three of the four localities where tadpoles were collected, and C. iiliomelus is the only species of Coloi^tetluis known to occur at two of those localities. Furthermore, a tadpole in Stage 43 has a well-defined dorsolateral stripe as in adults, anci recently metamorphosed young with SVLs of 10.8-14.1 mm ha\e the incipient color pattern of the adults. A typical tadpole (KU 215607) in Stage 34 has a hod\ length of 14.2 mm and a total length o\ 34.7 mm; body wider (9.3 mm) than high (7.2 mm); snout bluntly rounded in dorsal view, sloping anteroventralh' from le\el of or- bits to rounded tip in profile; external nares small, about midway between snout and orbits; eye small (1.4 mm) situated dorsally, directed dorsolaterally, not visible from below; interorbital distance 3.5 mm; spiracle sinistral: tube short, attached to body for its entire length; spiracu- lar opening directed posterodorsally just below midline at about midlength of body; cloacal tube short, dextral, attached to ventral fin. Caudal musculature moderateh' Stage Body L'ngth Total length 25 23 28 5 29 5 34 2 35 2 36 4 40 3 4-! 1 8.4-13.5 (x = 10.64) 12.2-13.1 (x = 12.50) 13.0-14.2 (x = 13.66) 14.2-14.3 (x = 14.25) 14.0-14.1 (x = 14.05) 14.6-15.5 {x = 14.98) 15.(V16.2(\ = 15.53) 16.3 16.9-35.6 (x = 29.10) 32.2-35.3 (x = 33.48) 33.2-39.0 (x = 35.10) 34.7-35.5 (x = 35.10) 36.8-37.0 (x = 36.90) 36.0-38.5 (x = .37.93) 36.9-12.6 (x = 39.90) 36.6 robust, approximately uniform in depth on anterior third of tail, gradually diminishing distally to pointed tip; dorsal fin originating on base of caudal musculature, gradually increasing in height on proximal two thirds of tail, and declining posteriorly to an acutely rounded tip; ventral fin originating on body wall, highest at midlength of tail, where equal in height to caudal musculature, noticeably less than height of dorsal fin (Fig. 9C). Oral disc 3.9 mm wide, directed anteroventrally; deep lateral folds in labia; median half of anterior labium bare; elsewhere labia with single irregular row of small, sub- conical marginal papillae; submarginal papillae absent; few rounded papillae in lateral folds. Jaw sheaths thin, finely serrate; anterior sheath in form of broad arch; posterior sheath broadly V-shaped; LTRF 2/3; all rows of teeth about equal in length. In life, body olive-brown; belly creamy gray; tail tan with olive flecks and brown spots or reticulations; it is tan to bronze, in preservative, dorsum and sides of body dark brown; belly gray; caudal musculature tan with midlateral brown streak on proximal third of musculature; caudal fins translucent with brown reticulations, most numerous and minute white flecks on fins and musculature on distal third of tail (Fig.9C). Distribution and ecology. — Colo^tcthiis idionieliifi is kninvn from several localities at elevations of 1620-2200 m and two localities at 22S0 m and 2840 m in humid montane forest in the northern part of the Cordillera Central in Peru (Fig. 10). Individuals were found in spring seepages and along small streams. At a site at 2180 m on the east slope of Abra Pardo Miguel, C. idi- oiticliis occurs sympatricalU w ith the slightly larger C. lU'iiisiiiiosii^, and at a site at 1620 m, it occurs s\-mpatri- cally with C. iiiittrnticicri. Remarks. — Ihe original description ot Colo>tcthiifi itlioiiiclu> was based on a single subadult female, not an adult temale as stated by Rivero (1991a). Colostcthiis insiilatiis new species Holotype. — KU 211857, an adult temale, trom 17 km FNL (by road) of Balsas (ca. 06 30' S, 77'56' VV), 1810 m, Proxincia de Chachapoyas, Departamento de Amazonas, COLOSTETHUS OF NORTHERN PeRU 25 Peru; one of a series collected by Fernando M. Cuadros and John J. Wiens on 21 January 1989. Paratopotypes.— KU 211858-70, 2 adult males, 11 adult females; same collectors and date. Referred specimens. — See Appendix 1. Diagnosis. — A moderately small species of Colostethiis with the following characteristics: (1) maximum SVL of males 22.2 mm, of females 26.0 mm; (2) disc on Finger III expanded, half again width of penultimate phalange; (3) Finger I longer than Finger 11, (4) lateral fringes present on fingers; (5) disc on Toe IV expanded, half again wicith of penultimate phalange; (6) lateral fringes present on toes; (7) outer tarsal fold and tarsal tubercle absent; inner tarsal fold curved, distinct on distal half of tarsus; (8) toes webbed basally; (9) dorsolateral stripe absent; (10) oblique lateral stripe present; (11) ventrolateral stripe absent; (12) gular- chest region cream with pair of brown spots laterally on throat; (13) abdomen cream; (14) no sexual dimorphism in coloration; (15) median lingual process absent; (16) Finger III not swollen in males; (17) testes white. Few species of Colostctliiis in Ecuador and Peru have Finger I longer than Finger II and an oblique lateral stripe. Of these, C. argyrogaster differs from C. insulatus by having a pale ventrolateral stripe, orange inguinal region in life, no fringes on the digits, and basal webbing only between Toes III and IV. Colostethus leitcophacus differs by having the toes about one-half webbed and the oblique lateral stripe present only in the groin, whereas C. pulclwrrimus differs by having black flanks with pale blue flecks. Colostctliits fiigax and C. maclialilln differ by having Finger 111 swollen in males and by lacking fringes on the digits and discrete dark marks in the gular-chest region. The larger C. tonchi also lacks fringes on the digits and has a pale ventrolat- eral stripe, as does C. iiiclaiiolneiuus, in which the oblique lateral stripe is diffuse. Other species that have Finger I longer than Finger II in the region include C. conspicuous, kingsburyi, oriiatus, and tiilaiiuvicae, all of which have dor- solateral stripes, and all except C. talamancae lack fringes on the digits. Colostethus acruginosus and C. mittermcicri also have Finger 1 longer than Finger II, but they lack all longitudinal stripes. Description . — (N = 32: 9 males, 22 females, 1 juvenile). SVL 17.2-22.2 (x = 19.7) mm in males, 20.2-26.0 (x = 22.8) mm in females; body robust; head about as wide as long; head length 33.3-43.6% (x = 34.9%) of SVL; head width 32.3-40.2"v, (x = 34.9%) of SVL; snout moderately long, bluntly rounded in dorsal view, rounded in profile; loreal region slightly concave; nostrils barely protuberant later- ally, anterior margins slightly posterior to level of anterior margin of lower jaw; eve-nostril distance 58.6-72.7% (x = 64.6%)) of length of eve; supratvmpanic bulge weak, barelv covering upper and posterodorsal edge of tympanum; otherwise, tympanic annulus distinct; length of tympanum 46.4-58.6% (x = 50.6','J.) of length of eye, separated from eye by distance less than one-fourth length of eye. Forelimb moderately long, robust; Finger 1 slightly longer than Finger 11; fingers not webbed, bearing narrow lateral fringes; Finger 111 in males not swollen; terminal discs expanded, about half again width of penultimate phalanges; subarticular tubercles moderately large, sub- conical; supernumerary tubercles absent; palmar tubercle large, round; thenar tubercle small, elongately elliptical; nuptial excrescences absent. Hind limb short, robust; tibia length 42.9-18.2% (x = 45.6%) SVL; foot length 41.0-t8.0% (x = 44.6%) SVL; outer tarsal fold absent; inner tarsal fold curved from low tubercle at midlength of tarsus to elliptical inner metatarsal tubercle; outer metatarsal tu- bercle subconical; toes webbed basally; webbing formula I2*-2*II2-3III3-4 IV4 -3V; lateral fringes present on toes; terminal discs expanded, about half again width of pen- ultimate phalanges; subarticular tubercles small, round; supernumerary tubercles absent. Skin on dorsum of body and hind limbs, flanks, and venter smooth; cloacal opening directed posteriorly at upper level of thighs; cloacal sheath long. Tongue long, widest and shallowly notched posteriorly, free behind for about half of its length; median lingual process absent. Color in preservative: Dorsum of head and body brown with small, irregular, dark brown spots, principally on body; dark brown stripe across tip of snout through loreal and tympanic regions to anterior part of flanks; midflank gray; groin tan. Dorsolateral and ventrolateral stripes absent; oblique lateral stripe, creamy white, distinct from groin to posterior part of upper eyelid. Forelimb pale tan, with or without irregular brown marks on forearm; dorsal surfaces of hind limbs brown, usually with darker brown transverse bars — two each cin thigh, shank, and foot; anterior surfaces of thighs brown with cream longitudinal stripe on at least proximal half of thigh, continuous with oblique lateral stripe, diffuse or indistinct in some speci- mens; posterior surfaces of thighs brown with or without diffuse darker brown mottling; upper lip pale tan, usu- ally with faint brown mottling. Venter cream with pair of diffuse brown or grayish brown spots posterolaterally on throat; palmar and plantar surfaces pale brown; webbing translucent. Color in life Dorsum reddish brown with dark brown middorsal marks, anterior two or three of which broadly chevron-shaped, posterior two usually round, quadran- gular, or elongate; upper lip and proximal dorsal surfaces of upper arm white; rest of forearm brown with diffuse grayish-white mottling; oblique lateral stripe creamy white; flanks gray mottled with brown; dorsal surfaces of thighs dull brown with darker brown markings ranging from irregular spots to transverse bars; anterior surfaces 26 Scientific Papers, Natural History Museum, The University of Kansas of thighs brown with narrow creamy-white longitudinal stripe connected to oblicjue lateral stripe in groin; throat gray with pair of black spots; belly creamy white; ventral surfaces of hind limbs pale gray with diffuse, dull orange spot on proximal ventral surface of shank; digital scutes white to pale gray in contrast to brown digits; iris bronze with black flecks (Figs. 5D and 6A). Measurements of holotype (in mm): SVL 22.0; tibia length 9.9; foot length 9.6; head width 7.8; head length 7.9; eye-nostril distance 2.0; length of eye 2.9; length of tympanum 1.7. Tadpoles. — Tadpoles were collected at three of the lo- calities where adults were found; because only one species of Colostetlius was found at each of these sites and because metamorphosing young are like the adults, these tadpoles are assigned to Colpftcthiifi iiisulntus. Tadpoles in various stages of development were found in a rocky pool at the base of a waterfall, in a small pool in a seepage area, and in a small spring-fed pool (Table 8). A typical tadpole (KU 215600) in Stage 34 has a body length of 13.0 mm and a total length of 38.8 mm; body glob- ular, wider (9.6 mm) than high (7.0 mm); snout rounded in dorsal view and in profile; nares directed dorsolater- ally about midway between tip of snout and orbits; eyes moderately large (1.7 mm), situated and directed dorso- laterally, not visible from below; interorbital distance 3.2 mm; spiracle sinistral, short, attached for its entire length to body wall; spiracular opening directed posterodorsally well below midline at about midlength of body; cloacal tube short, dextral, attached to ventral fin. Caudal muscula- ture robust, being equal in height throughout proximal half of tail, diminishing gradually to pointed terminus; dorsal fin originating on base of caudal musculature, reaching greatest height at about midlength of tail, diminishing to bluntly rounded tip; ventral fin originating on body wall, reaching greatest height just posterior to midlength of tail, where dorsal and ventral fins equal in height, and slightly more than height of caudal musculature (Fig. 9D). Oral disc 3.8 mm wide, directed anterolaterally; me- dian half of anterior labium bare; elsewhere labia bearing single row of slender, pointed marginal papillae; submar- ginal papillae absent; labia infolded laterally; few small, round papillae in folds. Jaw sheaths moderately slender, coarsely serrate; anterior sheath forming broad arch; lower sheath broadly V-shaped; LTRF 2(1)/ 3; A, longest; P, short- est; other rows equal in length. In preservative, dorsum and sides of body dark brown; belly translucent; caudal musculature tan with fine brown reticulations; caudal fins translucent with brown flecks on dorsal fin (Fig. 9D). In life, body olive-brown; tail brown with olive flecks; iris pale bronze. Recently metamorphosed young colored like adults, except slightly paler and lacking dark spots on throat. Table 8. Measurements (in mm) of free-swimming tadpoles of Colostethus iiisniatus. Range of variation followed by mean in parentheses. Body Total Stage N length length 25 12 5.4-9.7 (x = 7.83) 11.6-25.1 (x = 19.36) 29 10.2 23.3 31 10.4 23.6 32 11.3 28.4 34 12.5-13.5 (x = 13.00) 35.5-38.8 (x = 37.30) 37 15.0 41.5 41 14.2 39.2 43 14.3 34.3 44 14.3 24.2 46 3 14.2-14.8 (x = 14.43) — Distribution and ecology. — Colostethus insulatiis in- habits the middle Rio Marafion Vallev in northern Peru, where it is known from elevations of 1260-2600 m (Fig. 4). This region supports dry scrub forest, which in some areas is dominated by cacti. A few small streams cascade down the steep slopes from the adjacent highlands of the Cordillera Occidental anci Cordillera Central. Colostethus were found by day on and under rocks adjacent to streams and especially in spray zones of waterfalls; others were under rocks in seepage areas or adjacent to small spring- fed streams. Tadpoles were found in quiet pools. Etymology. — The specific name, iiisulntiis, is Latin meaning isolated and refers to the disparate mesic environments in- habited by this species in otherwise inhospitable terrain. Remarks. — One male (SVL 18.3 m) and nine females (SVLs 20.2-24.9 mm; x = 22.8) from 2 km NW of San Juan, 2290 m, Departamento de Cajamarca, resemble typical Co- lostethus iusulatus in size and structure, but differ somewhat in coloration. The dorsum is dull olive-gray with brown markings; the oblique lateral stripe is pale orange-tan (Fig. 5D). The flanks are gray with cream spots, and the venter is dull creamy white with a pair of diffuse brown spots on the throat. This sample is from the headwaters of the drainage of the Rio Cajamarca, which tlows into the Ri'o Marafion. Colostethus leucophaeiis new species Holotype.— KU 211879, an adult female, froni Mo- linapampa (06"11' S, 77"38' W, 2400 m), Provincia de Chachapoyas, Departamento de Amazonas, Peru; one of a series collected by William E. Duellman and Michael E. Morrison on 26 January 1989. Paratypes.— KU 211880-83and MHNSM 6215-17col- lecled with the holotype. Diagnosis. — A moderate-sized Colostethus with the following characteristics: (1) maximum SVL of males 25.2 mm, of /emales 26.1 mm; (2) disc on Finger 111 expanded. COLOSTETHUS OF NORTHERN PeRU 27 slightly wider than penultimate phalange; (3) Finger 1 lon- ger than Finger 11; (4) lateral fringes present on fingers; (5) disc on Toe IV expanded, slightly wider than penultimate phalange; (6) lateral fringes present on toes; (7) outer tarsal fold and tarsal tubercle absent; inner tarsal fold curved, dis- tinct on distal two-thirds of tarsus; (8) toes about one-half webbed; (9) dorsolateral stripe absent; (10) oblique lateral stripe present posteriorly; (11) ventrolateral stripe absent; (12) gular-chest region cream; (13) abdomen cream; (14) ncT sexual dimorphism in coloration; (15) median lingual process absent; (16) Finger III not swollen in males; (17) testes white. Only a few species in Ecuador and nortliern Peru have Finger I longer than Finger II, an oblique lateral stripe, and no dorsolateral stripe. Of these, C. argifwgaster differs from C. Iciicophnciis by having a pale ventrolateral stripe, orange inguinal region in life, no fringes on the digits, and basal webbing only between Toes III and IV. Colostctlnib piilclicr- rimus differs by having black flanks with pale blue flecks. Colostcthuf fiignx and C. niacJialilla differ by having Finger III swollen in males and by lacking fringes on the digits. The equally large C. tonchi also lacks fringes on the digits and has a pale ventrolateral stripe, as does C. iiielaiiolncmiis, in which the oblique lateral stripe is diffuse. Four other species have the toes at least one-half webbed; these are C. fidiginofits, iiexipiis, pcculinris, and soniidatiis. Fingers I and II are equal in length in C. soniidntus, Finger I is shorter than Finger II in C. fiiliginosuf, nexipus, and peciilinris. Description. — (N = 8: 3 males, 2 females, 3 juveniles). SVL 24.7-25.2 mm (x = 25.0) in males, 23.7-26.1 mm (x = 24.9) mm in females; body robust; head slightly longer than wide; head length 34.9-36.0% (x = 35.3%) of SVL; head width 34.1-35.2% (x = 34.7%) of SVL; snout moderately short, bluntly rounded in dorsal view and in profile; loreal region slightly concave; nostrils barely protuberant later- ally, anterior margins slightly posterior to level of anterior margin of lower jaw; eye-nostril distance 62.5-63.3% (x = 62.7%) of length of eye; supratympanic bulge weak, barely covering upper and posterodorsal edge of tympanum; otherwise, tympanic annulus distinct; length of tympanum 53.1-56.2% (x = 54.4%) of length of eye, separated from eye by distance about one-third length of eye. Forelimb moderately long, slender; Finger I much longer than Finger II; fingers not webbed, bearing broad lateral fringes; Finger III not swollen in males; terminal discs expanded, slightly wider than penultimate phalan- ges; subarticular tubercles moderately small, low, rounded; supernumerary tubercles absent; palmar tubercle large, round; thenar tubercle small, ovoid; nuptial excrescences absent. Hind limb short, robust; tibia length 40.9^4. l"-. (x = 42.4%) of SVL; foot length 42.9-49.0% (x = 45.0%) of SVL; outer tarsal fold absent; inner tarsal fold elevated, curved on distal two thirds of tarsus to small, elliptical inner metatarsal tubercle; outer metatarsal tubercle sub- conical; toes slightly more than one-half webbed; webbing formula ir-2Iir-3-III2-3'IV3-l ^V; lateral fringes present on toes; terminal discs expanded, wider than penultimate phalanges; subarticular tubercles small, low, round; super- numerary tubercles absent. Skin on dorsum of body and hind limbs, and venter smooth; skin on flanks finely shagreen; cloacal opening directed posteriorly at upper level of thighs; cloacal sheath short. Tongue long, widest and shallowly notched posteri- orly, free behind for about half of its length; median lingual process absent. Color in preservafive Dorsum of head, body, and limbs dull brown with small, faint, irregular dark brown spots on body; diffuse dark brown stripe across tip of snout, through loreal and supratympanic regions, present or not on anterior part of flanks; otherwise flanks grayish brown. Dorsolateral and ventrolateral stripes absent; oblique lat- eral stripe creamy white, present only in groin. Forelimb brown with darker brown irregular marks on forearm; anterior surface of upper arm tan with brown longitudinal stripe; dorsal surfaces of hind limbs brown with darker brown transverse bars — three or four each on thigh, shank, and foot; anterior surfaces of thighs creamy tan with con- tinuation of transverse brown bars; posterior surfaces of thighs creamy tan with dark brown longitudinal stripe on posteroventral surfaces; upper lip and tympanic region pale tan. Venter cream with fine melanophores on throat; palmar and plantar surfaces brown; webbing cream. Three juveniles having SVLs of 18.5-19.8 mm (x = 19.3) are colored like the adults. However, the pattern on the dorsal surfaces of the hind limbs is more striking; the interspaces between the dark brown transverse bars are creamy tan. Color in life: Dorsum dull grayish brown; flanks pale grayish brown; limbs, including digital scutes, dull brown with dark brown transverse bars; canthal and supratympan- ic stripes brownish black; shtirt, dull cream oblique lateral stripe in groin; faint, pale gray line along posterior surfaces of thighs; belly dirty white; throat and ventral surfaces of limbs tinged with dull yellow; iris dull brown (Fig. 6B). Measurements of holotype (in mm) SVL 26.1, tibia length 11.5, foot length 12.8, head width 9.0, head length 9.0, eye-nostril distance 2.0, length of eye 3.2, length of tympanum 1.7. Tadpoles. — A small series of tadpoles, mostly in later stages of development, was obtained from a slow-moving, marshy stream in a pasture at the type locality. These tad- poles are associated with Colostcthus Icucopliaciis because it is the only species of the genus known from the locality and because the metamorphosing tadpoles closely resemble the adults in coloration. The tadpoles are much larger than those of other species in northern Peru (Table 9). 28 Scientific Papers. Natural History Museum, The University of Kansas Table 9. Measurements (in mm) of free-swimming tadpoles of Colostcthus U'ucophai'Ui. Range of variation follovved line by mean in parentheses. Stage N Body length Total length 28 15.6 32 20.0 40 20.5 43 18.7 44 2 18.8 31.15) 37.5 45.0 50.2 48.5 .2-34.1 (x A typical tadpole (KU 215603) in Stage 28 has a body length of 15.6 mm and a total length of 37.5 mm; body ovoid, slightly wider (9.5 mm) than high (8.6 mm); snout in dorsal view bluntly rounded, nearly truncate, rounded in profile; nares directed dorsolaterally about midway between tip of snout and orbits; eyes moderately small (1.6 mm), situated and directed dorsolaterally, not visible from below; interorbital distance 4.4 mm; spiracle sinistral, tube short, attached to body wall; sinistral opening directed posterodorsally well below midline at about midlength of body; cloacal tube short, dextral, attached to ventral fin. Caudal musculature, gradually diminishing from body to acutely rounded terminus; dorsal fin originating on anterior margin of caudal musculature, highest at about two-thirds of tail length, gradually diminishing to bluntly rounded tip; ventral fin originating on body wall, highest at about three-fourths of tail length; at midlength of tail, dorsal fin lower than ventral fin, neither as high as caudal musculature (Fig. 9E). Oral disc 3.9 mm wide, directed anteroventrally; median two thirds of anterior labium bare; rest of labium bearing short, blunt marginal papillae; two rows of papillae in lateral folds; elsewhere, one row of papillae; submarginal papillae absent. Jaw sheaths moderately robust, serrated; anterior sheath in form of broad arch; posterior sheath widely V-shaped; LTRF 2(l)/3; all rows about equal in length. In preservative, dorsum and sides of body dark brown; belly dark grayish brown; caudal musculature tan, becom- ing gray on distal third; fins brown (Fig. 9F). In life, body and tail dark gray. Distribution and ecology. — This species is known only from the type locality at an elevation of 2400 m in the northern part of the Cordillera Central in Departa- mento de Amazonas, Peru (Fig. 10). The type series and the tadpoles were found amid water cress-like plants in a slow-moving, marshy stream in a pasture bv dav. In the same marshy stream, adults, metamorphosing young, and tadpoles of Gastrotliecn monticola and Scinax ovciic^ v\erc found. Etymology. — The specific name is a Latin adjective meaning ash-colored and refers to the generally dull gray appearance of this nondescript frog. Colostcthus mittcrmeieri Rivero Cohstelhm mittermeieri Rivero, 1991a:.3. Holotype: MCZ A-100217 from Venceremos, Departamento de San Martin, Peru, 1620 m. Diagnosis. — A moderately large species of Colostethus with the following characteristics: (1) SVL in males to 27.2 mm, in females 28.2 mm; (2) disc on Finger III expanded, about half again width of penultimate phalange; (3) Finger I longer than Finger II; (4) lateral fringes present on fingers; (5) disc on Toe IV expanded, about half again width of the penultimate phalange; (6) lateral fringes present on toes; (7) curved inner tarsal fold on distal half of tarsus; (8) toes about one-fourth webbed; (9) dorsolateral stripe absent; (10) oblique lateral stripe absent; (11) ventrolateral stripe absent; (12) gular-chest area with paired brown spots; (13) abdomen mottled anteriorly; (14) no sexual dimorphism in ventral coloration; (15) median lingual process absent; (16) Finger III not swollen in males; (17) testes white. The only other species in Ecuador and Peru with Finger I longer than Finger II and lacking longitudinal stripes is Colostethus aeniginosus; this species differs from C. mitter- meieri by having unwebbed toes and a dark \'cnter with pale spots. Description . — (N = 2: 1 male, 1 female). Body mod- erately robust; head longer than wide; head length 35.1 and 37.7';;, (x = 36.4) of SVL; head v\'idth 32.6 and 33.3'^ (x = 33.0) of SVL; snout moderately short, bluntly rounded in dorsal view, rounded abo\'e and truncate in profile; loreal region flat; nostrils barely protuberant laterally, anterior edge at level of anterior border of lower jaw; eve-nostril distance 71.0 and 72.2'V', (x = 71.6) length of eye; supratympanic bulge rather massive, diffuse, cover- ing posterodorsal part of tympanum; length of tympanum 47.2-52.6''(; (x = 49.9) length of eye, separated from eye by distance about one-tenth length of eye. Forelimb long, moderately robust. Finger I longer than Finger II; fingers unwebbed, bearing lateral fringes; Finger 111 not s\yollen in males; terminal discs expanded, about half again width of penultimate phalanges; subarticular tubercles moderateh' large, subconical; supernumerary tubercles absent; palmar tubercle large, round, elevated; thenar tubercle small, ovoid; nuptial excrescences absent. 1 lind limb short, robust; tibia length 45.4-47.6% (x = 46.5) SVL; foot length 44.7-46.5';, (x = 45.6) SVL; outer tarsal fold absent; inner tarsal fold low, shallow sigmoid curve on distal one-half of tarsus; inner metatarsal tubercle small, nearly round; outer metatarsal tubercle larger than inner, round, separated from inner tubercle by large, ovoid medi- an metatarsal tubercle; toes about one-fourth webbed; web- bing tornuila U— (2-2 )I1(2 -2)— 3111(3 -3)— 41V4— 2' V; COLOSTETHUS OF NORTHHRN PkRL' 29 Fig. W. Ci)/i)s/i'//;i(s iiiittcniicicn. KU 211444, male, 27.2 mm. lateral fringes present on toes; terminal discs expanded, about halt again width of penultimate phalanges; subartic- ular tubercles small, subconical; supernumerar\' tubercles absent. Skin on clorsum of bod\' and hind limbs finely shagreen with scattered, minute tubercles in supracloacal region; skin on flanks shagreen; skin of ventral surfaces smooth; cloacal opening directed posteriorly at upper level of thighs; cloacal sheath short, wrinkled. Tongue elongately cordiform, widest and barely indented posteriorly, free behind for about half its length; median lingual process absent. Color in preservative: Dorsum of head and body dull brown with many small, irregular, faintly darker brown markings (Fig. 11); flanks dull brown, sliglitly darker than dorsum; dorsolateral stripe absent; oblique lateral stripe absent, but white dash (one side) or two elongate white spots (other side) in inguinal region of holotype and faint tan stripe in inguinal region in KU 211944; ventrolateral stripe absent; forelimbs pale brown; dorsal surfaces of hind limbs pale brown with barely discernible darker brown transverse marking; anterior surfaces of thighs pale brown; posterior surfaces of thighs brown with diffuse cream mottling; upper lip tan, continuous cross snout. Venter cream with brown spots anterolaterally on throat and dif- fuse brown pigment on throat, chest, and anterior part of abdomen; palmar and plantar surfaces tan; brownish gray transverse marks on dorsal and ventral surfaces digits. Color in life: Unknown. Unfortunately, the single new specimen (KU 211944) was confused with individuals of ColostetJnif aenigiiiosus in the field and no color notes or photographs were taken. Distribution and ecology. — This species is known from onh' two localities at ele\'ations of 1620 and 2030 m in iiumid cloud forest on the eastern slope of the northern 3°-/ 7'/ .' 78°^--^ 77 ;i Kll.iHR-kTv Am: 1 -, 1 ■ C. rnttermaen • C. nexipus O C. pulchernrws D C- ulcubambenas 4 ' i \ -»- \ ( . 7- .-v/' L.,^ ;/ f \ '\ ' -79 78 1 V=4 \ 7' 77" Fig. 12. Loaililios of occurrence of four species of Colo^tctlms. part of the Cordillera Central in Departamento San Mar- tin, Peru (Fig. 12). Both localities are on the road from Balzapata to La Rioja. The specimen from 2050 m was in a small, mossy stream by day. Remarks. — 1 compared KU 21 1944 with the holotype (MCZ A-100217); the two specimens are extremely similar and differ onlv in KU 211^44 ha\ing slightly less webbing and less tuberculate skin in the supracloacal region. How- ever, there are several discrepancies between the holotvpe and its description bv Ri\'ero (1991a). Rivero stated that Finger I was shorter than Finger II (longer, and shown as longer in his Figure 1 ) and that lateral fringes were absent on the fingers (definitely present). Rivero (1991a) men- tioned lateral white stripes on the lower flanks and black longitudinal streaks on the forelimbs; although I did not observe these, more than a decade has elapsed since Rivero examined the specimens, and some of the pattern mav have been lost. Hovve\'er, 1 did not obser\e those features on KU 211944. Rivero (1991a:6) referred to the dorsum of the holotype as "solid dark grayish brown"; there are distinct, darker brown marks on the dorsum. Furthermore, Rivero (1991 a:(S) provided measurements and proportions of three "males." These specimens have been examined bv Taran Grant, who stateci that all three are ju\'enile females (Grant, in lift.). 30 Scientific Papers, Natural History Museum, The University of Kansas Colostethiis nexipus Frost Colostetluis iiexipuf Frost, 1986:214. Holotype: BMNH 1983.1061 from Los Tayos, Provincia de Morona-Santiago, Ecuador. Coloma, 1995:44. Colostethuf citrcicola Rivero, 1991b:ll. Holotype: USNM 282687 from the immediate vicinity of LimOn (General Plaza), 1097 m, Provincia Morona-Santiago, Ecuador [Synonymy fide Coloma, 1995:44]. Diagnosis. — A large species of Colostetlui$ with the following characteristics: (1) maximum SVL in males 30.0 mm, in females 33.0 mm; (2) disc on Finger 111 expanded, half again width of penultimate phalange; (3) Finger 1 shorter than Finger II; (4) lateral fringes present as narrow ridges on fingers; (5) disc on Toe IV expanded, half again width of penultimate phalange; (6) lateral fringes present on toes; (7) inner tarsal fold weak or absent; tarsal tubercle absent; (8) toes about half webbed; (9) dorsolateral stripe present; (10) oblique lateral stripe present; (11) ventrolateral stripe absent; (12) gular-chest region cream; (13) abdomen cream; (14) no sexual dimorphism in ventral coloration; (15) median lingual process absent; (16) Finger 111 not swollen in males; (17) testes black. Relatively few species of Colot^tctlms Ecuador and Peru have Finger 1 shorter than Finger II; with the exception of C. nexipus and C. patitae, all of these species (C. aiva, chocoensis, fuli^^iiwsus, idiomelus, maquipucuna, peculiaris, pukhellus, and puniihis) lack a dorsolateral stripe, which is prominent in C. iicxipus. According to L'tterset al. (2003), C. patitae differs from C. iicxipiis by lacking webbing between Relatively few species of Colostcthits Ecuador and Peru have Finger I shorter than Finger II; with the exception of C. nexipus and C. patitae, all of these species (C. aioa, chocoensis, fuli^inosus, idiomelus, maquipucuna, peculiaris, pulchelhis, and pumilus) lack a dorsolateral stripe, which is prominent in C. nexipus. According to L'tters ot al. (2003), C. patitae differs from C. nexipus by lacking webbing between the toes and in having a bright yellow (instead of orange) dorsolateral stripe. Four other species have the toes at least half webbed; these are C. fuliginosus, leucophaeus, peculiaris, and sonlidatus, all of which lack a dorsolateral stripe. Fingers 1 and II are equal in length in C. sordidatus and Finger I is longer than Finger 11 in C. leucoplineus. Four species (C. lwcagei\ chocoensis, mittenneieri, and sauli) are smaller (males < 25 mm, females < 30 mm) and ha\ e less extensive webbing than C. nexipus. Furthermore, all four species lack dorsolateral stripes; Fingers I and II are equal in length in C. bocai^ei and C. sauli, and Finger I is longer than Finger 11 in C. mittermeieri. The toes are only webbed basally in C. aim, elacln/histus, fugax, iiifraguttatus, and sylvaticus, all of which also differ from C nexipus h\ 'In a phylogenetic tree of some dendrobatid frogs based on molecular data, Santos et al. (2003) included CdoMcthus bocn^ci and C. ninculo^ii^. Tlie latter is a junior subjective synonym of C. l^ocn^fi (Coloma, 1995). One of the authors of Santos etal. (Luis A. Coloma) informed me (in litt., 22iulv 2004) that "Our recognition ... |of C. "W(ii/ii.s»s] was not justified." lacking dorsolateral stripes. The smaller C. axva also has less webbing and Finger 1 shorter than Finger 11. All other Colostethus in the region either lack webbing or if basal webbing is present (usually only between Toes 111 and IV), they are smaller in size and have Finger I longer than Finger II; these include C. alessandroi, brunneus, conspicuus, fratisenescus,fuscellus, gascoiii, insperntus, inachalilla, marche- sianus, mcdiarmidi, luelanolaennis, stepheni, toachi, trilineatus, and imnzolinius (Coloma, 1995; Grant and Rodriguez, 2001; Morales, 2002). Description. — Frost's (1986) original description that was supplemented by Coloma (1995) can be expanded considerably by the series of specimens from northern Peru (N = 10 males, 11 females, 5 juveniles). Body robust; SVL in males 18.6-20.5 (x = 19.3) mm, in females 20.0-23.1 (x = 21.4) mm; head about as long as wide; head length 31.2- 38.1% (x = 35.6) of SVL; head width 31.2-37.6";- (x = 35.4) of SVL; snout moderately short, bluntly rounded indor.sal view, bluntly rounded above and inclined postero\'entrally in profile; loreal region flat; nostrils not protuberant later- ally, with posterior border at level of anterior margin of lower jaw; eye-nostril distance 58.1-75.0% (x = 63.9) of length of eye; supratympanic bulge diffuse, barely obscur- ing posterodorsal edge of tympanum; length of tympanum 40.7-46.9% (x = 44.2) of length of eye, separated from eye by distance about one-fifth length of eye; tvmpanic annulus distinct. Forelimb long, moderately slender; Finger 1 shorter than Finger II; fingers unwebbed, bearing narrow lateral fringes; Finger III not swollen in males; terminal discs ex- panded, about half again width of penultimate phalanges; subarticular tutiercles small, rounded; supernumerary tu- bercles absent; palmar tubercle large, nearly o\oid; thenar tubercle slightly smaller, elliptical; nuptial excrescences absent. Flind limb short, moderatelv robust; tibia length 44.7-54.8% (x = 50.1) of SVL; foot length 43.0-53.1% (x = 47.8) of SVL; outer tarsal fold and tarsal tubercle absent; inner tarsal fold low, shallowly sigmoid on distal half of tarsus; inner metatarsal tubercle low, ovoid; outer meta- tarsal tubercle subconical; toes about half webbed; modal webbing formula I1-2II1-2III1-3IV3-1 V; lateral fringes present distally to webbing; terminal discs expanded, about half again width of penultimate phalanges; subar- ticular tubercles small, rounded; supernumerar\' tubercles absent. Skin on dorsum of body and hind limbs finely shagreen with minute tubercles postsacrally on body; skin on flanks shagreen; skin of ventral surfaces smooth, except for weak granules on posterior part of abdomen; cloacal opening directed ventrallv at midlevel of thighs; cloacal sheath short. Tongue long, widest and shallowh' notched posteri- orly, free behind for about [\\o thirds of its length; median lingual process absent. Coi-OSTETHVS OF NORTHERN PeRU 31 Color in preservati\e: Dorsum of head, body, and limbs brown, v\'ith or without irregular dark brown mid- dorsal marks on body; tan dorsolateral stripe extending from posterior margin of upper evelid to paracloacal region, becoming diffuse or fragmented postsacrally, bordered abo\e and below by dark brown stripes; lower brown stripe continuation of facial stripe across snout and through e\'e and t\-mpanum; cream\' tan oblique lateral stripe extending from groin to point above insertion of upper, fragmented posteriorh' in four indi\iduals. Flanks below stripes brown; limbs brown with dark brown trans- verse bars, extending onto anterior and posterior surfaces of thighs; upper lip cream\' tan. Ventral surfaces of body anci limbs creamy tan; minute brown flecks on throat and chest in some specimens; palmar and plantar surfaces pale brown. Color in life: Dorsum of head and body dull brown to nearly black, usualh' v\ith darker brown or black irregular marks; flanks black; dorsolateral stripes originating on snout and extending to paracloacal area, cin-\ed slightly medialh' at midlength of body, variable in color from nearh' red to most commonly orange or to tan; in some individuals, changing from orange anteriorly to bluish tan posteriorly; oblique lateral stripe cream, pale orange, or usually bluish \\ bite (Fig. 6C). Upper lip and proximal dorsal surface of upper arm creamy tan to pale yellow; dorsal surfaces of hind limbs tan to pale bluish gra\' with dark brown trans\'erse bars or less commonK' irregular marks; anterior and posterior surfaces of thighs and ventral surfaces of limbs bluish gray; digital scutes \\ hite to pale gray; throat and belh' wliite to pale X'ellow; iris reddish copper. Tadpoles. — A male having a SVL of 19.7 mm was transporting 12 tadpoles on 15 February 1989. The tad- poles have body lengths of 3.6-3.8 mm (x = 3.66) and total lengths of 8.7-8.9 mm (x = 8.80). These small tadpoles are in Stage 24; the oral disc and associated structures are not developed, and the body cavity is filled with yolk. Free-swimming tadpoles were found in a pool in a stream and in a pool adjacent to anther stream. Tadpoles are in various stages from Stage 25 to metamorphosis (Table 10); these tadpoles are associated with Colostetlnis ncxipus by the toes being about half webbed in metamorphosing young and by the dorsolateral stripe in large individuals. Furthermore, Colostetlnis iie.xipiif was the only member of the genus found at localities where the tadpoles were col- lected. A typical tadpole (KU 215594) in Stage 37 has a body length of 9.2 mm and a total length of 30.5 mm; body ovoid, wider (5.4 mm) than high (3.5 mm); snout bluntly rounded in dorsal view, rounded in profile; nares directed anteriorly about midway between tip of snout and orbits; eyes large (1.3 mm), situated and directed dorsolaterallv, not visible Table 10. Measurements (in mm) of free-swimming tadpoles of Colostetlnis iicxipus. Range of variation followed b\' mean in parentheses. Stage N Body length Total length 23 -> 14.25) 28 2 29 1 31 3 21.17) 32 T 21.20) 34 -) 37 3 29.3) 39 1 5.3-6.2 (x = 3.75) 6.5 6.6 7.3-8.7 (x = 8.00) 7.9-8.3 (x = 8.20) 8.9-9.0 (x = 8.95) 8.8-9.2 (x = 8.97) q.8 12.7-15.8 (x 16.0 18.5 20.5-22.4 (x 19.1-23.3 (x 23.3 28.4-30.5 (x 24.5 from below; interorbital distance 1 .7 mm; spiracle sinistral; tube short, attached to bod\' for its entire length; spiracu- lar opening directed posterodorsally well below midline at about two thirds of length of body; cloacal tube short, dextral, attached to ventral fin. Caudal musculature robust, about equal in height cm proximal third of tail, gradually diminishing to pointed terminus; dorsal fin originating on base of caudal musculature, greatest height at mid- length of tail, gradualh' diminishing to acutely rounded tip; ventral fin originating cin body wall, about equal in height throughout its length; at midlength of tail, dorsal fin slighth' higher than \entral fin and about equal to height of caudal musculature (Fig. 9F). Oral disc 2 mm wide, directed antero\'entrall\'; median two thirds of anterior labium bare; elsewhere, labia with single row of subconical marginal papillae; submarginal papillae absent; labia with shallow lateral folds bearing few small, rounded papillae. Jaw sheaths moderateh' slender, coarsely serrate; anterior sheath forming broad arch; posterior sheath broadh' V-shaped; LTRF 2(l)/3 ; anterior rov\'s slightly longer than posterior rows. In preservative, dorsum and sides of bod\' dark red- dish brown with minute white flecks; caudal musculature tan, reddish brown dorsallv; fins translucent, essentialh' unmarked (Fig. 9F); indixiduals in Stage 41 with larger, bluish-white lichenous flecks on all body surfaces. In life, body dark brown with tan dorsolateral stripe in largest individuals; tail tan with brown flecks. Distribution and ecology. — This species is widely distributed in departamentos of Amazonas and San Martin in northern Peru, where it has been taken at an elevation of 360 m in the floodplain of the Rio Huallaga, elevations up to 810 m in a spur of the Andes northeast of Tarapoto, and at an elevations of 325 and 520 m on the western slope 32 Scientific PapiiRs. N.vi l km. HisroK'i Museum, The University of Kansas of the northern part of the Cordillera Central (Fig. 12). Most of these localities are at lower elevations than those in southern Ecuador, where the species has been found at elevations of 500-1500 m (Coloma, 1995). All Peruvian specimens were associated with rocky streams in February; most were active by day, but a few were on boulders in streams at night. One juvenile and one adult male carrying tadpoles v\'ere on leaves in the spray zone of a waterfall at night; another male carrying tadpoles was on a rock in a stream by day. Free-sw imming tadpoles were in pools in streams. At Ponga de Shilcayo, a narrow ravine at an elevation of 470 m, 4 km NNVV of Tarapoto, Colo^tetlius ih'xip>us is sympatric with se\ en other dendrobatid frogs — Colosethus ornaliis, Cryptopln/llobates azueriiroentris, Deiiciwbatesfantasti- ciif, D. imitator, Epipcdolmtcs /wss/cr/, £. hnhin-ii, and £. trivit- tatui. Only C. uexipus was found along the rocky stream in the ravine; the other dendrobatids were on moss-covered boulders, on the forest floor, or in terrestrial bromeliads {D. fanta>ticu^). Remarks. — Colofti'thii^ iicxipiis is one of the most brightl V colored members of the genus; the presence of skin toxins in C. iicxipus has not been determined. In the most extensive molecular phylogeny of dendrobatids (Santos et al., 2003), C. nexiput^ is the sister taxon of an aposematic species, Cryptophyllolmtcs nziiiriz'ciitri^. Colostcthus ornatus Morales Cohstclhuii ornatus Morales, 2002:37. Hololype: MHNSM 17713 from Tarapoto, 350 m, Provincia San Martin, Departamcnto de San Martin, Peru. Diagnosis. — A small species of Colostctiiiis with the following characteristics: (1) maximum SVL in males 16.9 mm, in females 18.4 mm; (2) disc on Finger 111 expanded, half again width of penultimate phalange; (3) Finger 1 lon- ger than Finger II; (4) lateral fringes absent on fingers; (5) disc on Toe IV expanded, half again v\'idth of penultimate phalange; (6) lateral fringes absent on toes; (7) inner and outer tarsal folds absent; outer tarsal tubercle present; (8) basal webbing present between Ti>es II and 1\'; elsewhere webbing absent; (9) dorsolateral stripe present; (10) i>blic]ue lateral stripe absent; (11) ventrolateral stripe present; (12) gular-chest region gray in male, white in female; (13) abdo- men creamy white; (14) sexual dimorphism in coloration; (15) median lingual process absent; (16) I'inger 111 swollen in males; (17) color of testes unknown. The only other species of Coiostetlnis in Ecuador and Peru with Finger I longer than Finger 11, dorsolateral and ventrolateral stripes, and no oblique lateral stripe is C. talamancae, a larger frog (SVL to 24 mm) that is confined to the Chocoan Region and that also differs from C. oniatiis by having lateral fringes on the fingers and toes. Fur- thermore, the middorsum of C. ialamaiicae is essentially uniform dark brown, whereas a rhomboidal dark brown mark is present on the tan dorsum of C. onintu^, in which the posterior surfaces of the thighs lack the hook-shaped dark mark characteristic of C. talamancae (Savage, 1968). Description. — The description of this species by Mo- rales (2002) is minimally adequate. Here, it is augmented by a description of a paratype (adult female, KU 211950) from Ponga de Shilcayo, Departamento de San Martin. Body slender; head longer than wide; head length 32.9% of SVL; head width 30.6% of SVL; snout long, broadly truncate in dorsal view, bluntly rounded in profile; loreal region flat; nares not protuberant, posterior margins at level of anterior margin of lower jaw; eye-nostril distance 58.3% of length of eye; supratympanic bulge diffuse, barely obscuring upper edge of t\mpanum; otherwise, tympanic annulus distinct; length of t\mpanum 50'r. of length of eye, separated from eye by about one-fourth length of eye. Forelimb long, slender; Finger I much longer than Fin- ger II; fingers not webbed, lacking lateral fringes; Finger III not swollen in males (fide Morales, 2002); terminal discs ex- panded, about half again width of penultimate phalanges; subarticular tubercles large, subconical; supernumerary tubercles absent; palmar tubercle large, nearly round; the- nar tubercle large, elliptical. Hind limb moderately short, slender; tibia leiigth 50.3' ;, of SVL; foot length 46.8'; of SVL; tarsal folds absent; inner tarsal tubercle small, elongate; in- ner metatarsal tubercle elliptical; outer metatarsal tubercle small, subconical; toes long, slender, with basal webbing between Toes II and III and between Toes III and IV; lateral fringes absent; terminal discs expanded, about half again width of penultimate phalanges; subarticular tubercles small, subconical; supernumerary tubercles absent. Skin on all siu-faces smooth; cloacal opening directed posteroventrallv at upper le\el of thighs; cloacal sheath short. Tongue long, widest posteriorly, not notched pos- teriorlv, free behind for about half of its length. Color in preserxative: Dorsum tan v\ith brown markings consisting of a broad, diffuse interorbital bar, X-shaped mark in scapular region with anterior arms of X extending to c\elids and posterior arms confluent with diagonal marks connected to teardrop-shaped mark on posterior part of dorsum; broad, dark brown stripe across snout, through loreal and suprat\inpanic regions, to groin, bordered abo\'e b\- tan dorsolateral stripe. Hind limbs tan with single, broad transverse bar each on thigh, shank, and foot; anterior sinlaces of thighs tan; posterior surfaces brown; forelimbs cream with dark bi'ow n longitudinal stripe on posteriiir surface ot upper arm; upper lip, lower half of tNinpanum, and lower part o\ tlank cream; \enter and palmar siuiaces cream; plantar sinlaces brown. Color in life Dorsum tan with bro\sn markings, con- sisting of an interorbital bar X-shaped mark in scapular region; posterior parts of X-shaped mark connected to a roughR- triangular mark in sacral region: dorsolateral COLOSTETHLS Ul- NuRTHtRN PeRU 33 stripe pale vellow, narrow on head, diffuse on biKiv; broad, dark brown stripe across tip of snout, tlirougln loreai and tympanic regions becoming broader on tlank and extend- ing to groin, bordered below on flanks b\ narrow pale vellow \entrolateral stripe; oblique lateral stripe absent; flanks below stripe cream\' tan with brown flecks; dorsal surfaces of hind limbs gra\ish brown with man\' small dark brown flecks and one broad transxerse brown bar on thigh, shank, and foot; posterior surfaces of thighs brown; labial region creamy white with brown flecks; throat and bellv pale lemon vellow; \'entral surfaces of limbs flesh colored; digital scutes pale gray in contrast to adjacent dark brown surfaces; iris greenish l^ronze (Fig. 6D). Measurements (in mm): SVL 17.3, tibia length S.7, foot length 8.1, head \\ idth 5.3, head length 5.7, eye-nostril distance 1.4, length of eye, 2.4, length oi tympanum 1.2. Distribution and ecology. — This small species is known only from the vicinity of Tarapoto in northeastern Peru, where it has been found at elevations of 350-680 m in disturbed lowland rainforest and lower humid montane forest (Fig. 4). Morales (2002) provided no ecological data on the species; an indixidual was acti\e on the forest flcior by day in February. Remarks.— Morales (2002) designated KU 211950 as a paratype of Colostctlm^ onuUii> from 12 km northeast of Tarapoto, 720 m, Departamento de San Martin, Peru. The locality given is erroneous; the specimen is from Ponga de Shilcayo, about 4 km NNW of Tarapoto, 470 m. The color pattern of KU 211950 resembles that of the holot\'pe illustrated bv Morales (2002), except that the an- terior part of the X-shaped mark on the head is separated from that part in the scapular region. The single female examined contained six large, unpig- mented ovarian eggs about 2 mm in diameter. The sizes of the ovarian complement and of the oxarian eggs are like those of Co/os^'f/;((s ^tcpliciii (June-, 1998). The eggs of C. stcphciii hatch into non-feeding nidicolous tadpoles that complete their development in the terrestrial nest (June- et al., 1994). Thus, C. ornatu^ mav be another member of the genus \\ ith nidicolous tadpoles. Colo^tctlni::^ poeciloiiotiis Rivero Caloitetlnif {'oecilonotuf^ Rivero, 1991a: 13. Holotvpo: 1V1CZA-S9108 from Alva, Departamento Amazonas, rem. Diagnosis. — A moderate-sized species of Colostelluis with the following characteristics: (1) maximum SVL in males 19.7 mm, in females 25.2 mm; (2) disc on Finger III slightly expanded; (3) Finger I equal in length to Finger II; (4) lateral fringes absent on fingers; (5) disc on Toe IV slightly expanded; (6) lateral fringes absent on toes; (7) outer tarsal fold and tarsal tubercle absent; inner tarsal fold cur\'ed on distal half of tarsus; (8) toes unwebhed; (9) dorsolateral stripe absent; (10) oblique lateral stripe present from groin to midflank; (11) ventrolateral stripe absent; (12) gular-chest region uniform creamv tan; (13) abdomen creamv tan; (14) no sexual dimorphism in ventral coloration; (15) median lingual process absent; (16) Finger III not swollen in males; (17) testes white. Many species of Colostctliu^ in the region lack a dor- solateral stripe and ha\e an oblique lateral stripe and Fingers I and II equal in length. Of these, only C. hoca^ci has a short ol">lique lateral stripe; in the others (C. aiithra- ciims, hrcviqitartiis, ccvalkvi, dclntoiirnc, linchi/hi^tus, fii^nx. idioDicIiis, iiifrn^iittntus, Ichmanni, saiili, and vcrtebmli^), the olilique lateral stripe extends anteriorly to, or nearly to, the eye, instead of only to the midflank as in C. poccilo- notitt^. Of these species, C. cez'ullo^i and C. Ichmmmi also differ from C. poi'cikviotii> bv having black arm bands, as defined by Grant and Ardila-Robayo (2002). Furthermore, C. clachi/histus, fii^ax, >auli, and ^ortiidafus also differ from C. pocciloiiotu> bv having webbing between the toes, and C. Liacln/histiis, iiifiagiittatub, idionwlus. and sauli haw lat- eral fringes on the fingers and toes. Other species with an oblique lateral stripe but no dorsolateral stripe (C. idioimius and C. insidatuf-) have Finger I longer than Finger II, and in C. in:>ulatus, the oblicjue lateral stripe extends anteriorly to the eve. The only other species in the region ha\ ing Fingers 1 and II equal in length are C. fylvaticut^, wliich has lateral fringes on the digits, a dorsolateral stripe, and no oblic]ue lateral stripe, and C. cxn^pcrntus, which has dorsolateral and olilique lateral stripes, and lateral part of the abdomen marbled. Description. — {N = 4: 1 male, 3 females). Bod\' mod- erately robust; head slighth' longer than wide; head length 32.9-36.5% (x = 34.1) of SVL; head width 29.4-32.4'; (x = 30.8) of SVL; snout moderately long, bluntly rounded, nearly truncate in dorsal view, truncate in profile; loreai region flat; nostrils slightly protuberant laterally, with pos- terior border at level well behind anterior margin of lower jaw; eye-nostril distance 65.3-69.0''v( (x = 67.0) length of eye; supratympanic bulge diffuse, covering posterodorsal part of tympanum; length of tympanum 51.7-60.0% (x = 55.2) length of eye, separated from eye by distance about one- sixth length of eye; tympanic annulus evident externally only anteriorly and \'entrally. Forelimb long, moderately slender; Fingers I and II equal in length; fingers unwebbed, lacking lateral fringes; Finger III not swollen in males; terminal discs slightly ex- panded, barely wider than penultimate phalanges; subar- ticular tubercles large, rounded; supernumerary tubercles absent; palmar tubercle large, rounded; thenar tubercle small, elliptical; nuptial excrescences absent. Hind limb short, robust; tibia length 44.0-48.2% (x = 46.2) SVL; foot length 42.8-50.8'v. (x = 46.3) SVL; outer tarsal fold absent; inner tarsal fold shallowly sigmoid on distal half of tarsus; inner metatarsal tubercle small, elliptical; outer metatarsal 34 SciHN niic Papuks, Natural Hiskjry Museum, Thh Univhrsity ot- Kansas tubercle equal in size, subconical; toes unwebbed, lacking lateral fringes; terminal discs expanded barely expanded; subarticLilar tubercles small, rcuind; supernumerary tu- bercles absent. Skin on dorsum of body and liind limbs shagreen with small tubercles postsacrallv and larger tubercles in t}'mpanic region; skin on flanks shagreen; skin of posterior part of abdomen finely granular; other ventral surfaces smooth; cloacal opening directed posteroventrally at mid- level of thighs; cloacal sheath short. Tongue ovoid, widest posteriorly, not notched, free behind for about half of its length; median lingual process absent. Color in preservative: Dorsum of head and body brown with many small, nearly round, dark brown spots; dark brown stripe across tip of snout extending through loreal and supratympanic regions, continuous with dark brown flanks with irregular creamy white spots ven- trally; pale dorsolateral stripe absent; pale tan to dull white oblit]ue lateral stripe extending from groin to point between axilla and midflank; ventrolateral stripe absent; upper lip and tympanic region tan with faint grayish brown irregular markings. Forelimb pale brown with dark brown transverse bars on forearm and dark brown longitudinal stripes on anterior ancH posterior surfaces of upper arm; dorsal surfaces of hind limbs brown with narrow dark brown transverse bars; anterior surface of thigh tan \\'ith longitudinal dark brown stripe; posterior surface of thigh brown v\'ith diffuse cream\' white longitudinal stripe. All ventral surfaces creamy tan, except plantar surfaces brown. Color in life: Unknown. Distribution and ecology. — Colostcthii^ poccilonotus is known only from the type locality at an elevation of 1000 m on the slopes of the Cordillera Central above the Rio Ut- cubamba Valley, which is mostly cultivated but otherwise mainly supports thorn forest (Fig. 10). Remarks. — Examination of the type series revealed several discrepancies between the specimens and Ri vero's (1991a) description; he correctly stated that Fingers 1 and 11 are equal in length, but his illustration (Fig. 5) clearly shows the first finger much longer than the second. Rivero (1991a) reported all four specimens to be adult females; dissection revealed that one of the paratypes ( MCZ A-S'-) 1 (W) to be .w adult male. Measurements and proportions ri'ported here differ slightly from those given by Ri\'ero. Colostethus pulcherrimus new species Holotype. — KU 21194h, an adult female, trom the im- mediate vicinity of Cutervo (06"22' S, 78"49' W, 2620 m), Pro\'incia de Cutervo, Departamento de Cajamarca, Peru; oneof a series collected by Fernando M. Cuadros and John J. Wiens on 26 February 1989. Paratypes.— KU 211947-49 and MHNSM 6251 col- lected with the holotype. Diagnosis. — A moderately large species of Colostethus v\ith the following characteristics: (1) maximum SVL in males 28.2 mm, in females 29.7 mm; (2) disc on Finger 111 expanded, slightly wider than penultimate phalange; (3) Finger I longer than Finger 11; (4) lateral fringes present on fingers; (5) disc on Toes IV expanded, slightly wider than penultimate phalange; (6) lateral fringes present on toes; (7) outer tarsal fold and tarsal tubercle absent; inner tarsal fold curved on distal half of tarsus; (8) webbing absent between toes; (9) dorsolateral stripe absent; (10) oblique lateral stripe present; (11) ventrolateral stripe absent; (12) gular-chest area cream with or without brown mottling; (13) abdomen cream with or v^'ithout brown mottling; (14) no sexual dimorphism in ventral coloration; (15) median lingual process absent; (16) Finger 111 not swollen in males; (17) testes white. Of the other species in Ecuador and northern Peru that have the Finger I longer than Finger II, an oblique lateral stripe, and no dorsolateral stripe, Colo>tctliii$ argywgafiter differs from C. /»///( //crr/'/Hf/s by having a pale ventrolateral stripe, orange inguinal region in life, no fringes on the digits, and basal webbing only between the Toes III and IV. Colofitcthus Icucophacii^ differs by having the toes about half webbed. Colostethus fiij^nx and C. iiinclinlilla differ by having the Finger III swollen in males and b\' lacking fringes on the digits. The equalh' large C. toaclii also lacks fringes on the digits and has a pale xentrolateral stripe, as does C. incliviolncunis, in which the oblique lateral stripe is diffuse. Description. — (N = 5: 1 male, 4 females). Body robust; head slightly longer than wide; head length 30.9-33.7% (x = 33.1) of SVL; head width 29.1-31.8'; i\ = 29.9) of SVL; snout short, roimded in dorsal view, blunth' rounded in profile; loreal region flat; nostrils not protuberant, anterior border at le\el slighfh' posterior to anterior margin of lower jaw; eye-nostril distance 62.5-70.0% (x = 66.9) of length of e\e; supratxmpanic l^ulge diffuse, co\ering upper and pos- ferodiM'sal edges of t\mpanum; t\mpanic annulus distinct; length of t\mpanum 59.4-66.7% (x = 63.7) of length of eye, separated from e\e b\' distance abciut ime-fmn'th length of t\nipanum. Forelimb inoderafeK short, robust; Finger I longer than Finger II; fingers unwebbed, bearing narrov\- lateral fringes; Finger III not swollen in males; teiminal discs expanded, slightK' wider than penultimate phalanges: subarticular tubercles low, rounded: supernumerar\ tubercles absent; COLOSTETHVS OF NoRTHfiRN PeRU 35 palmar tubercle large, nearly round; thenar tubercle large, elliptical; nuptial excrescences absent. Hind limb short, robust; tibia length 44.2-47.0% (x = 45.2) of SVL; foot length 48.9-51.1':;, (x = 49.9) of SVL; outer tarsal fold and tarsal tubercle absent; inner tarsal fold flaplike, forming sigmoid cur\e on distal half of tarsus; inner metatarsal tubercle small, elliptical; outer metatarsal tubercle small, subconical; toes unwebbed, bearing narrow lateral fringes; terminal discs expanded, slightly wider than penultimate phalanges; subarticular tubercles small, rounded; super- numerary tubercles absent. Skin on dorsum of head, body, and liind limbs, and on venter smooth; skin on flanks finely shagreen; cloacal opening directed posteroventrally at upper level of thighs; cloacal sheath short. Tongue long, widest and shallowlv notched posteriorly, free behind for about half of its length; median lingual process absent. Color in preservative: Dorsum of head and body dull brown with faint, darker brown, irregular marks; flanks dark brown with cream flecks; upper lip tan with brown flecks; dorsolateral and ventrolateral stripes absent; oblit]ue lateral stripe narrow, cream, extending from groin to posterior edge of orbit or midflank. Forelimbs brown; dorsal surfaces of hind limbs dull brown with darker brown transverse bars — three each on thigh, shank, and foot; anterior and posterior surfaces of thighs brown with cream mottline. Throat, chest, abdomen, and ventral surfaces of limbs cream with varying amounts of brown mottling — extensive in holotvpe, slight in cine male and two females, negligible in one female; palmar and plantar surfaces brown. Color in life: Dorsum of head and bod v pale grayish green with irregular coppery-brt^wn irregular markings; oblique lateral stripe dull tan anteriorly, bluish white posteriorly; flanks black with blush-white flecks and streaks; upper lip pinkish tan, becoming white poste- riorly; digital scutes pale gray bordered h\ black; iris bronze-brown (Fig. 6E). Measurements of holotype (in mm): SVL 28.0, tibia length 12.6, foot length 14.3, head width 8.2. head length 8.8, eye-nostril distance 2.1, length of eve 3.2, length of tympanum l.*-^. Distribution and ecology. — This species is known only from two small streams in the immediate vicinitv of Cutervo, a village at an elevation of 2620 m in a culti\'ated valley in the northern part of the Cordillera Occidental (Fig. 12). The frogs were active in the streams bv day; adult Colostcthus chwhxfhistiis were found in the same streams. Etymology. — The specific name is a Latin adjecti\e meaning "prettiest," in reference to the rather striking coloration in contrast to the rather drab appearance of many species of Colostcthus in the Peruvian Andes. Colostctlnis sordidatiis new species Holotype.— KU 21 1960, adult female, from 30 km SVV of Zapatero (about 10 km NE San Jose de Sisa [ca. 06"45' S, 76"33' W]), 500 m, Provincia de Lamas, Departamento de San Martin, Peru; one of a series collected by William E. Duellman and Rainer Schulte on 13 February 1989. Paratopotypes. — KU 211961-72; same collectors and date. Diagnosis. — A relatively large species of Colostctlms with the following characteristics: (1) maximum SVL in males 29.9 mm, in females 36.1 mm; (2) disc on Finger 111 expanded, about twice width of penultimate phalange, nearly truncate; (3) Finger 1 ec]ual in length to Finger II; (4) lateral fringes present on Angers; (5) disc on Toe IV expanded, about twice width of penultimate phalange; (6) lateral fringes present on toes; (7) outer tarsal fold and tarsal tubercle absent; inner tarsal fold curved on distal two-thirds of tarsus; (8) toes about two-thirds webbed; (9) dorsolateral stripe absent; (10) oblic]ue lateral stripe pres- ent from groin to midflank; (11 ) ventrolateral stripe absent; (12) gular-chest region pale gray with dark gray intrusion laterally on throat; (13) abdomen creamv tan; (14) sexual dimorphism in ventral coloration; (15) median lingual process absent; (16) Finger 111 not swollen in males; (17) testes white. Colostctlms sordidatiis is compared only with species that have webbing between the toes from Ecuador, Peru, Bolivia, and western Brazil. Three other species have the toes at least half webbed; these are C. fuligiuosus, iiexipnis, and pccidiaris. Both C. iicxipiis and C. pt'ciilinris differ from C. soididatus by having Finger I shorter than Finger II (equal in length in C. sordidatiis); C. iwxipiis also differs by having a distinct pale dorscilateral stripe, whereas C. pccidiaris also differs by lacking a pale oblique lateral stripe. Four species (C. hocagei, cliocoeusis, mittermcicri, and sauli) are smaller (males < 25 mm, females < 30 mm) and have less extensive webbing than C. sordidatiis. Furthermore, in C. cliocoeusis Finger I is shorter than Finger II, whereas the opposite is the case in C. mittermcicri; Fingers I and II are equal in length in C bocagci, sauli, and sordidatiis. The only other species with moderately webbed toes, no dorsolateral and ventral stripes, and Fingers I and II equal in length are C. clacliyhistus and C. iufragitttatiis; in both of these species, the oblique lateral stripe extends anteriorly to the orbit (only to midflank in C. sordidatiis). The smaller C. aiim also has less webbing and Finger I shorter than Finger II. All other Colostcthus in the region either lack webbing or if basal webbing is present (usuallv onlv between Toes Topotypic Coloitethn^ iniirclifiianui' were described by Caldwell et al. (2002), who noted that most references to this species in Colombia, Ecuador and Peru applied to other species, many of which were named bv Morales (2(1U2). 36 SCIKNTIFIC PaPIRS. NaIIRAI, HiSTORV MuSKUM, ThH Um\ FRSIT^ OF Kansas in and IV, but also between Toes II and III at least in C. maniiesimuif*), they are smaller in size and have Finger I longer than Finger II; these include C. alessaiidivi, bniii- nciis, coiispiciius, fratisoicsciis, fiisccllns, i^ascoiii, iiispcmtus, machalilla, marcheiianiis, mcdiarniidi, nielaiiolaeuius, stcphciii, toachi, trilinealus, and vanzoUnius (Coloma, 1995; Grant and Rodriguez, 2001; Morales, 2002). Description. — (N = ll 8 males, 3 temalos). Body ro- bust; head about as long as wide; head length 32.1-40.0% (x = 36.2) of SVL; head width 31 .9-38.8% (x = 36.2) of SVL; snout moderately long, bluntly rounded in dorsal view, nearly truncate with slight posterolateral inclination in profile; loreal region flat; nostrils slightly protuberant lat- erally, with posterior border at level of anterior margin of lower jaw; eye-nostril distance 67.6-75.6% (x = 71.4) length of eye; supratympanic bulge diffuse, covering upper edge of tympanum; length of tympanum 41.9-55.9%, (x = 49.3) length of eye, separated from eye by distance about one- fourth length of eye; tympanic annulus evident externally only anteriorly and ventrally. Forelimb moderately long, robust; Fingers I and 11 equal in length; fingers unwebbed, bearing narrow lateral fringes; Finger III not swollen in males; terminal discs expanded, about twice width of penultimate phalanges; subarticular tubercles large, rounded; supernumerary tubercles absent; palmar tubercle large, broadly ovoid; thenar tubercle small, narrowly elliptical; nuptial excrescences absent. Hind limb short, robust; tibia length 42.9-50.4% (x = 47.5) SVL; foot length 43.4-49.6' ;, (x = 46.3) SVL; outer tarsal fold absent; inner tarsal fold shallowly sigmoid on distal two-thirds of tarsus; inner metatarsal tubercle low, elongately elliptical; outer metatarsal tubercle round; toes about two-thirds webbed; modal webbing formula I1-2II1-2III2 -3IV3-1 V; lateral fringes present distally to webbing; terminal discs expanded, about twice width of penultimate phalanges; subarticular tubercles small, subconical; supernumerary tubercles absent. Skin on dorsum of body and hind limbs finely shagreen; skin on flanks finely granular; skin of ventral surfaces smooth; cloacal opening directed posteroventrally at upper level of thighs; cloacal sheath short. Tongue about three times as long as wide, widest and shallowly notched posteriorly; free behind for about one-third of its length; median lingual process absent. Color in preservative: Dorsum of head and bod\ dull brown with scattered small brown spots; small, irregular white spots present on two males and two females; flanks dark brown; loreal and supratympanic stripe dark brown; pale dorsolateral and ventrolateral stripes absent; oblique lateral stripe dull white extending from groin to midflank; dorsal surfaces of upper arms and hind limbs brovsn with faint to distinct transverse darker brown bands — tv\'o on forearm and three each on thigh, shank, and foot; widtli of transverse bars about equal to width of pale brov\'n in- terspaces; anterior and posterior surfaces of brown with small, pale cream spots on the latter; margin of upper lip tan. Venter creamy tan \Nith gravish-tan suffusion pos- terolaterally on throat and margin of lower jaw; anterior part of chin grav in one male; throat and chest gray in two males; palmar and plantar surfaces gravish brown; web- bing translucent. Color in life: L^orsum pale brown with irregular dark brown spots and transverse bars on limbs; posterior sur- faces of thighs tan. Dark brown stripe across tip of snout, through nostril, to eye and continuous across t\'mpanum, above insertion of forelimb, onto flank to groin; pale dorso- lateral and ventrolateral stripes absent; dull white oblique lateral stripe extending from midflank to groin; dark brown crescent-shaped mark on anterior surface of upper arm not continuous with dark brown stripe on bod}-; digital scutes grayish tan; upper lip pale creamy tan (Fig. 6G). Venter in males gray; venter in females white, with throat and chest bright yellow; white extending into axillar\- region (Fig. 6H). Iris reddish bronze with small black flecks and median horizontal brown streak. Measurements of holot\'pe (in mm): S\'L 33.0; tibia length 15.5; foot length 14.5; head width 11.2; head length 11.7; eye-nostril distance 3.0; length of eye 4.3; length of tympanum 1.8. Tadpoles. — Two kinds of Colo:>ti'tliu^ tadpoles were found at the type locality of C. craspedoceps and C. sordida- tiis. One kind resembles back-riding tadpoles associated with C. craspedoccps. The other presumably is C. fordidntus and is described below. Tadpoles were in a pool in a stream in a rocky ravine on 13 February 1989. One individual in Stage 34 has a body length of 16.2 mm and a total length of 42+ mm (tail incomplete). Eleven individuals in Stage 25 have body lengths of 8.1-13.1 mm (x = 10.87) and total lengths of 19.4-33.5 mm (x = 26.44). A typical tadpole (KU 215611) in Stage 25 has a body length of 12.5 mm and a total length of 30.0; bodv globu- lar, wider (8.8 mm) than height (6.7 mm); snout bluntly rounded in dorsal \iew, rounded in profile; nares directed dorsolaterallv about midwav between tip of snout and orbits; eyes small (1.2 mm), situated dorsally, directed dorsolaterally, not visible from below; interorbital distance 3.1 mm; spiracle sinistral, spiracular tube short, attached to bodv throughout its length; spiracular opening directed nearly posteriorly just below midline at about midlength of bt)dy; cloacal tube short, cone-shaped, dextral, attached to \entral fin. Caudal musculature robust, about equal height throughout proximal two-fifths of tail, gradually diminish- ing to pointed tip; dorsal fin originating on proximal part of caudal musculature, reaching greatest height at about two-thirds length of tail, declining to acutely rounded tip; \entral fin originating on body, highest at midlength; dor- COLUSTETHUS OF NORTHhRN PeRU 37 sal fin higher than ventral fin and hoth higher than caudal musculature at midlength oi tail (I-ig. 9G). Oral disc 3.1 mm wide, directed anteroxentrally; me- dian half of anterior labium bare; elsewhere, labia bearing single row of small, subclinical marginal papillae; labia with lateral folds with small, rounded papillae; submar- ginal papillae absent, jaw sheaths narrow, finely serrate; antericir sheath forming broad arch; posterior sheath broadly V-shaped; LTRF 2(1 )/3; A, longest, P, shortest. In life, body brown; tail pale orange. In preser\'ative, dorsum and sides of body brown; belly translucent; caudal musculature tan proximallv, gray distallv; fins translucent; scattered guanophores on body, caudal musculatinv and fins (Fig. 9G). Distribution and Ecology. — This species is known only from the type locality at an elevation of 500 m near the eastern base of the Cordillera Central and from an el- evation of 520 m southwest of Chiriaco near the northern end of the Cordillera Central (Fig. 4). At the type locality, individuals were under rocks in a streambed by day; at night, they were perched on wet boulders in the streambed. The call is a series of abc^ut 10 whistle-like ncites. Etymology. — The specific name, ^ordidntus, is Latin meaning "dressed shabbily" and refers to the drab dorsal coloration of this species. Remarks. — The specimens from 20 km SW of Chiriaco (KU l%718-26 ) were collected on 17 and 18 December 1974 by Richard Thomas This series contains two subadult females with S VLs of 21 .3 and 26.5 mm and seven juveniles, two of which (SVLs of 10.3 and 12.0 mm) seem to be recent metamorphs. One of the subadult females has a few white spots on the dorsum. Colostetluis spilotogaster new species Holotype.— LSUMNS 39341, adult female, from the west slope of the Cordillera Coin, SE of La Peca, 8500 ft ( = 2326 m) (ca. 05_37' S, 78_2r W), Provincia de Utcubamba, Departamento de Amazonas, Peru; obtained on 1 7 October 1978 by Thomas S. Schulenberg. Diagnosis. — A moderate-sized species of Coloste- tluis with the following characteristics: (1) SVL of males unknown; maximum SVL in female 24.0 mm; (2) clisc on Finger III expanded, half again width of penultimate pha- lange; (3) Fingers I and II ec]ual in length; (4) lateral fringes present on fingers; (5) disc on Toe IV expanded, half again width of penultimate phalange; (6) lateral fringes present on toes; (7) outer tarsal fold absent; tarsal tubercle present; inner tarsal fold slightly curved on distal half of tarsus; (8) webbing basal between toes; (9) dorsolateral stripe pres- ent; (10) oblique lateral stripe absent; (11) ventrolateral stripe absent; (12) gular-chest region cream with brov\n spots; (13) abciomen cream with brown spots; (14) sexual dimorphism in ventral coloration not known; (15) median Imgual process absent; (16) nature of Finger III in males unknown; (17) color of testes unknown. Six cither species {Colostetluis, clciitlu'iodnct\/liis, exas- poiitiis. iicxipus, oiimtus, si/l~onticus, and utculnvidH'iisis) in the region iia\e pale dorsolateral stripes, but none has man\' small brown spots on the throat, chest, and abdo- men as does C. spilotogaster. Four tif thcise species (C. clcutlu'iodacti/liis, cxasperatiis, iicxipus, and utciibniiibciisis) also differ from C. spilotogaster by ha\'ing oblique lateral stripes. Colostetluis orimtiis also differs from C. spilotogaster by ha\'ing a ventrolateral stripe and Finger I longer than Finger II; C. sylvaticiis also differs by having a pair of large brown spots on the throat. Description. — {N = 1 female). Body moderately ro- bust; head longer than wide; head length 37.1' r- of SVL; head width 34.2"o of SVL; snout moderately short, broad, nearly truncate in dorsal view, truncate in profile; loreal region slightly concave; nostrils distinctly protuberant lat- erally, anterior border at level slightly posterior to anterior margin of lower jaw; eye-nostril distance 50% of length of eye; supratympanic bulge weak, obscuring posterodorsal edge of tympanum; tympanic annulus distinct, elevated; length of tympanum 78% of length of eye, separated from eye by distance about one-fifth length of eye. Forelimb moderately long, slender; Fingers I and 11 equal in length; fingers unwebbed, bearing narrow lateral fringes; terminal discs expanded, wicith about half again with of penultimate phalanges; subarticular tubercles rounded; supernumerary tubercles absent; palmar tubercle large, round; thenar tubercle small, elliptical. Hind limb short, robust; tibia length 44.2' J, of SVL; foot length 41.2' ^ of SVL; outer tarsal fold absent; inner tarsal fold originat- ing at small tarsal tubercle, slightly curved on distal half of tarsus; inner metatarsal tubercle small, elliptical; outer metatarsal tubercle small, subconical; toes webbed basally, bearing narrow lateral fringes; terminal discs expanded, width about half again width of penultimate phalanges. Skin on dorsum of head, body, and hind limbs smooth with scattered tubercles posteriorly on dorsum; skin on flanks finely granular; ventral surfaces smooth; cloacal opening directed posteroventrally at upper level of thighs; cloacal sheath short. Tongue long, widest and shallowly ncitched posteriorly, free behind for about one-half of its length; median lingual process absent. Color in preservative: Dorsum of head and body dull brown with discontinuous middorsal longitudinal dark brown stripe; upper lip cream; flanks brown with cream tips on granules; upper lip cream; dorsolateral stripe pale tan, originating on snout, passing along canthus rostralis and outer edge of upper eyelid to upper insertion of hind limb, bordered below on head by broad dark brov^'n stripe, bordered above on body by narrow dark brown stripe. 38 Scientific Papf.rs, Natural History Museum. Thi. University of Kansas Fig. 13. ColoilelliKS .•;f)i;(if(ix"s/i'r, LSUMNS 39341, fcnidlo, 24.0 mm S\L. Dorsal surfaces of hind limbs tan with indistinct brown spots; anterior and posterior surfaces of thighs brown with irregular small cream marks; forelimbs tan with dark brown longitudinal stripe on proximal part of anterior sur- face of upper arm and on entire length of posterior surfaces of upper arm. Venter cream with many small brown spots on throat, chest, and abdomen (Fig 13); palmar and plantar surfaces dark brown. Color in life: Unknown. Measurements of holotype (in mm): SVL 24.0, tibia length 10.6, foot length 10.0, head width 8.2, head length 8.9, eye-nostril distance 1.6, length of eye 3.2, length of tympanum 2.5. Distribution and ecology. — This species is known only from the type locality at i\n elevation of 2326 m in humid montane forest on the western slopes of the Cordillera Coin (Fig. 10). The only field notes accompan\ing the holotype are "leaf litter, cloud forest." Etymology. — The specific name is derived from the Greek spiloto^i, meaning spotted, and the Greek gnster, meaning belly; the name refers to the spotteci venter in this species. Remarks. — The distinctive combination oi a spot- ted venter, dorsolateral stripe, and large txmpanum sets Co/osf('//!ws ^pilotoj^d^tcr apart from all other species in the Andes of Ecuador and Peru; thus, the recognition of the species based on a single specimen is warranted. The ornithological expedition by the Museum of Zo- ology at Louisiana State University in 1978 revealed the existence of an apparently endemic suite of anurans. In addition to the Colo^tctliii^ named here, the endemics in- clude Gastrotlu'ca abdita (Duellman, 1987), Elcutlicrodacti/lii^ atrabrachtts, E. ai'iciiponini, and E. cuiiciw^tri^ (Duellman and Pramuk, 1999), and Tclniafobiiis colaneiisis (Wiens, 1993). Colostcthus iii/lvnticiis Barbour and Noble Pln/llohatefsylvatica Barbour and Noble, 1920:396. Holotype: MCZ 5344 from Tabaconas (near Huancabamba), Departamento de Cajamarca, Peru. Coloskihiif ^\/lvaticii — Edwards, 1971:148. Diagnosis. — A relatively large species of Colostetlms with the following characteristics: (1) maximum SVL in males 25.7 mm, in females 30.0 mm; (2) disc on Finger III slightly wider than penultimate phalange; (3) Finger I equal in length to Finger 11; (4) lateral fringes present on fingers; (5) disc on Toe IV barely wider than penultimate phalange; (6) lateral fringes present on toes; (7) inner tarsal fold usually present; (8) toes unwebbed; (9) dorsolateral stripe present; (10) oblique lateral stripe absent; (11) ventrolateral stripe absent; (12) gular-chest region pale with or without pair of dark spots on throat; (13) abdomen creamy wite; (14) usually sexual dimorphism in xentral coloration; (15) median lingual process absent; (16) Finger III not swollen in males; (17) Testis white. Colostetlms si/lvaticus can be distinguished from most other species in the region bv having the combination of Fingers 1 and II equal in length; dorsolateral stripe pres- ent, and oblique lateral stripe absent. Colostethus idiomelus shares this combination of characters, but it differs from C. s}/li'aticiis by ha\ing narrower fringes on the fingers and toes, fewer transverse brown marks on the hind limbs (2 each on thigh and shank, instead of 3 or 4 as in C. sylvati- oiis), a narrower and less distinct dorsolateral stripe, and a mottled black and white throat (instead of yellow as in C. si/lzHiticus). Some individuals of C. cxnspcrntiis have the first and second fingers equal in length, but they have a short oblique lateral stripe (Coloma, 1995). Colostetlms ek'iithcivdactylus has Fingers 1 and II equal in length, but it differs from C. s\/lvnticiis by having an oblique lateral stripe and greatly expanded discs on the digits. Likewise, C. sordidntiis has Fingers I and II equal in length, but it dif- fers from C. sylvaticiis by having greath' expanded digits, an oblique lateral stripe, and no dorsolateral stripe. Description. — (.V = 21: 8 males, 13 females). Body robust; SVL 20.8-25.7 mm (x = 23.8) in males, 27.1-30.0 mm (x = 28.5) in females; head longer than wide; head length 39.9-36.5% (x = 33.8) of SVL; head width 39.8-33.7% (x = 32.1) of SVL; snout moderately long, bluntly rounded in dorsal view and in profile; loreal region flat; nostrils not protuberant, anterior border slightly posterior to level of anterior margin of lower jaw; eye-nostril distance 62.5-77.4% (x = 68.6) length of eye; supratympanic bulge moderateh' massi\'e, covering posterodorsal half of tympa- num; length of tympanum 51.4-66.7"o (x = 59.5) length of eve, separated from eye by distance about one-fifth length ot eve. COLOSTETHUS OF NORTHHRN PeRU 39 Forelimb moderately long, slender; Fingers I and II equal in length; fingers unwebbed, with narrow lateral fringes; Finger III not swollen in males; terminal discs slightly wider than penultimate phalanges; subarticu- lar tubercles low, suliconical; supernimierarN' tubercles absent; palmar tubercle large, rinmd; thenar tubercle small, l")roadl\' o\oid; nuptial excrescences absent. Hind limb short, robust; tibia shorter than foot; tibia length 41.7-45.7% (x = 43.9) SVL; foot length 45.7-52.3% (x = 48.2) SVL; outer tarsal fold absent; inner tarsal fold, if evident, lovs-, curved on distal half of tarsus; inner metatarsal tubercle elliptical; outer metatarsal tubercle subconical; toes unv\ebbed or with basal webbing be- tween Toes IV and V, bearing narrow lateral fringes; terminal discs bareh' wider than penultimate phalan- ges; subarticular tubercles low, riumd, inccinspicuous; supernumerary tubercles absent. Skin on dorsum of boci\' and hind limbs weakly shaereen; skin on flanks shacreen; skin of \entral surfaces smooth; cloacal opening directed postero\'entrallv at micl- level of thighs; cloacal sheath short. Tongue long, widest and notched posteriorh', free behind for about half of its length; median lingual process absent. Color in preser\'ati\'e Dorsum of head and bod\' and flanks dull brown with irregular, darker brown markings ranging from flecks to dashes and che\'rons; dark brown stripe in loreal and tympanic regions; upper lip dull brown; dorsolateral stripe dull tan, narrow on snout and edges of upper eyelids, broad and usualh- distinct on bod\', extend- ing to groin, continuing as narrow stripe along upper inser- tion of hind limb and as diffuse stripe on anterior surface of thigh; oblique lateral stripe and xentrolateral stripe absent; arms pale brown with dark brown flecks; dorsal surfaces of hind limbs brown with dark brown transverse bars; ante- rior surfaces of thighs brown; posterior surfaces of thighs brown with cream flecks. Venter, including palmar and plantar surfaces pale tan; dusk\' suffusion cin throat and chest forming diffuse paired brown spots posterolaterally on throat in some males. Color in life: Dorsum olive-brown to copperv brown with dark brown to black flecks; digital scutes pale grav, contrasting to adjacent dark gravish-brown surfaces; up- per lip and dorsolateral stripe creamv tan to pale bronze; throat, posterior part of belh' and \entral surfaces of hind limbs dark vellow to orange; chest and anterior part of belly gray with black flecks; iris dull bronze with black flecks (Fig. 6F). Tadpoles.— In February 1979, a male {KU 181673) having a SVL of 25.7 mm was transporting 15 tadpoles (KU 181868) 4.2-4.9 mm (x = 4.50, N = 9) in body length and 12.1-13.2 mm (x = 12.78, ,V = 9) in total length. In the same month, free-swimming tadpoles (KU 181869-70) were found in muddy pools at elevations of 2560 and 3010 m Table 11. Measurements (in mm) of free-swimming tadpoles of Colostcthus si/lvniicus. Range of variation followed bv mean in parentheses. Body Total Stage N length length 30 13 7.0-8.1 (x = 7.42) 16.8-18.1 (x = 17.46) 31 S 6.9-11.0 (x = 8.32) 16.4-25.9 (x= 19.42) 32 1 11.3 25.1 33 10 12.0-13.0 (x= 12.47) 27.8-32.8 (x = 29.05) 34 4 12.5-14.1 (x = 13.27) 31.2-32.5 (x = 31.80) 33 ") 12.9-14.7 (x = 13.80) 38.1-39.3 (x = 38.50) on the eastern slope of the Cordillera de Huancabamba, Departamentci de Piura, Peru. The description of these tadpoles bv Duellman and Wild (1993) is expanded herein (Table 11).' A typical tadpole (KU 181869) in Stage 33 has a body length of 13.0 mm and a total length of 29.4 mm; body ovoid, wider (7.2 mm) than high (5.5 mm); snout depressed, acutely rounded in dorsal view, rounded in profile; nares directed anterolaterally about midway between tip of snout and orbits; eyes small (1.1 mm), situated dorsally, directed dorsolaterally, not visible from below; interorbital distance 1 .7 mm; spiracle sinistral; tube short, attached to body wall; spiracular opening directed posteriorly well below mid- line at about two-fifths length of body; cloacal tube short, dextral, attached to ventral fin. Caudal musculature mod- erately robust, about equal in height throughout proximal half of tail, then diminishing gradually to pointed terminus short of tip of tail; dorsal fin originating on proximal base of caudal musculattire, highest at about two-thirds length of tail, diminishing to blunth' rounded tip; ventral fin originat- ing on body wall, highest at midlength of tail, where each fin is slightly higher than caudal musculature (Fig. 9H) Oral disc 3.0 mm wide, directed anteroventrallv; me- dian half of upper labium bare; elsewhere, labia bear single row of moderately large subconical marginal papillae; submarginal papillae absent; some individuals with two rows laterally; shallow lateral fold. Jaw sheaths moderately robust, finely serrate; anterior sheath in form of broad arch; posterior sheath widely V-shaped; LTRF 2/3(1); anterior rows slightly longer than posterior rows; P^ shortest. In preservative, dorsum of anterior part of body tan with dark brown interorbital blotch; posterior half of body dark brown; unpigmented transverse bands ventrally in some specimens; anterior part of venter transparent; caudal musculature pale tan with brown streak medially on each side of anterior fourth of tail; caudal fins translucent (Fig. 9H). In life, body and tail olive-tan with green lichenous markings dorsallv on caudal musculature; belly greenish white; edge of oral disc yellow; iris pale bronze. 40 Scientific Paphrs, Natlirai. History Museum, The UNivhRsri'* oi- Kansas Distribution and ecology. — Colo^tctliu^ sylzmticiif is known hum clcwitions of 2450-3100 m in the Cordillera de Cuchaili, Cordillera de Huancabamba, and Cordillera de Tabaconas in northern Peru (Fig. 8). On the west slope of the Cordillera de Huancabamba, the species descends to 1920 m; its distribution there overlaps that of C. clachyhistus at elevations of 1920-2590 m. These frogs have been found under rocks in, and along, streams and active along the streams in cloud forest by day. The call is a short trill. C()/('s/c//;//5 iitculmDihcnsis Morales Cohslethiis ulaibamhensif Morales, 1994:103. Holotype: MHNSM 13986 from near Tingo (approximately 06*21' S, 77°49' W, 2391 m; but see remarks), Provincia de Luya, Dcpartamento Amazonas, Peru. Diagnosis. — A moderate-sized species of Colostcthus with the following characteristics: (1) maximimi SVL in males 19.8 mm, in females 26 mm; (2) disc on Finger 111 slightly expanded; (3) Finger 1 et]ual in length to, or longer than, Finger II; (4) lateral fringes absent on fingers; (5) disc on Toe IV slightly expanded; (6) lateral fringes absent on toes; (7) inner tarsal fold weak; (8) toes unwebbed; (9) dor- solateral stripe present; (10) oblique lateral stripe present; (11) ventrolateral stripe absent; (12) gular-chest area pale gray; (13) abdomen white or gray v\'ith diffuse white spots; (14) sexual dimorphism in ventral coloration; (15) median lingual process absent; (16) Finger 111 not swollen in males; (17) testis white. Onl\' three other species of Colostetlnis in Ecuador and northern Peru have dorsolateral and oblique lateral stripes; C. iicxipiib differs from C. iitciibaiiibeiisis by having Finger 1 shorter than Finger 11 and b\- having lateral fringes on the digits and toes about half webbed. Colostftluis clcittlwiv- dacti/liis differs b\ ha\ing Fingers 1 and 11 equal in lengtii, greatly expanded terminal discs on the digits, and a well- developed, curved inner tarsal fold. Colostcthus kiiii^^biiryi is like C. iitciibanihcii^i^ in having Finger 1 longer than Finger II, but it differs bv ha\'ing a ventrolateral stripe. Description. — (N = 3: 2 males, I female). Bod\' mod- erately stout; head slightly longei- than wide; head length 32.4-35.2% (x = 33.9) of SVL; 'head width 29.2-31.2';! (x = 30.9) of SVL; snout truncate in dorsal view, slightly inclined posteroventrally in profile; loreal region flat; nos- trils slightlv protuberant laterally, with posterior edge at level of anterior margin of lower jaw; eye-nt>stril distance 66.7-75.0% (x = 70.9) length of eye; suprat\mpanu bulge diffuse, covering dorsal part of t\'mpanum; lengtii oi t\'m- panum 41.9-53.1'V'- (x - 46.5) length of e\e, separated from eye by distance al"'out one-fourth length of e\e; txmpanic annulus evident externally onK' anteriorly and \entiall\'. Forelimb long, moderately slender; Fingers I equal to, or slightly longer than. Finger II; fingers unwebbed, lack- ing lateral fringes; Finger III not swollen in males; terminal Fig. 14. Colo^lfthui^ iilcid'iimh'iifii, MHNSM 13990, female, 253 mm S\'L. discs slightly expanded, half again as wide as penultimate phalanges; subarticular tubercles large, rounded; supernu- merary tubercles absent; palmar tubercle large, rounded; thenar tubercle smaller, o\'oid; nuptial excrescences absent. Hind limb short, robust; tibia length 43.9-46.1% (x = 45.0) SVL; foot length 41.9-44.0";, (x = 42.7) SVL; outer tarsal fold absent; inner tarsal fold shailowlv sigmoid on distal half of tarsus, originating at elliptical inner tarsal tubercle; inner metatarsal tubercle large, ovoid; outer metatarsal tubercle smaller, rounded; toes webbed basally, lacking lateral fringes; terminal discs expanded, about half again width of penultimate segments; subarticular tubercles small, round; supernumerar\' tubercles absent. Skin on dorsum and tlanks fineh' shagreen; skin on \entral surfaces smooth; cloacal opening directed posteri- orh' at uppi'r level of thighs; cloacal sheath short. Tongue, widest posteriorK', not notched, free behind for about half oi its length; median lingual process absent. Color in preser\ ative Dorsum of head and body brown with nearly round, dark brown spots (Fig. 13); dark brow n stripe across tip of snout extending through loreal and supralvmpanic regions and continuous \\ ith dark brow n flanks; scattered, small, creanw white spots on lower parts ol tlanks; di>rsolateral stripe broad, tan; oblique lateral sliipe pall' tan, exli'nding h\im groin lo anterior part of tlank; ventrolateral stripe absent; upper lip and tNinpanic region cream. Forelimb pale tan w ith tw i> dark brow n, trans- verse bars on forearm and dark brown longitudinal stripe on anterioi' surface of upper arm; dorsal surfaces of hind limbs tan with daik brown transverse bars — tour t>r five on thigh, four on sliank, twii on toot: anterior surtace of thigh tan with longitudinal daik bi'own stripe: posterior surtace ot thigh COLOSTETHUS OF NORTHKRN PeRU 41 brown with small creamy white spots. All ventral surfaces creamy white, except plantar surfaces brown (Fig. 12). Morales (1994) stated that the belly is gray in males and white in females; I noted no differences between the sexes 10 years after the description. Color in life: Unknown. Distribution and ecology. — This species is known onl\' from the tvpe localit\' in the valle\' of the Rio Ut- cubamba, a tributary of the Rio MaraOUn in northern Peru (Fig. 10) Remarks. — Morales (1994:95) reported the type locality as "cerca de Tingo, Rfo Utcubamba, 2391 m." My own altimeter readings at Tingo were 1810 m on 5 March 1979 and 1860 m on 24 January 1989, at which time I noted that tlie road follows the Rio Utciibamba, which in many places is in a narrow canyon and quite pictviresque. However, the hillsides have been cleared and the valley floor, what there is of it, is cultivated or in pasture. Below about 1900 m, columnar cacti are present, although at times of visits in Januaiy and Marcli the area seemed to be quite wet. In the diagnosis of this species. Morales (1994) stated that Fingers 1 and II were equal in length; in the descrip- tion of the holotype, he stated that Finger I was shorter than Finger II, whereas in the illustration of the hand of the holotype, Finger I is notably longer than Finger II. Upon my request. Morales reexamined the holotype and reported (in lift., 14 February 2004) that Finger I is longer than Finger 11. The similarities of Colostetlnis kingshiir]/i and C. iit- ciilmnibensis suggest that the two nominal taxa are either closely related or possibly conspecific. Rivero ( 1991 b) noted variation in the anterior extent of the oblique lateral stripe and the presence or absence of a ventrolateral stripe. Ac- cording to Coloma (1993), C. kiu;^dnin/i occurs in a narrow elevational belt of 1140-1300 m on the Amazonian slopes of the Andes in the Pastaza Trench and northward. Thus, a hiatus of about 550 km exists between the ranges of the two species. Until specimens become available from the intervening region or there is compelling evidence for their being conspecific, C. iitciibainhcnsis should be recognized as a distinct species. Likewise, preserved paratypes of C. poccilouotiis and C. iitciibniubeiisis reveal only subtle differences between the species (e.g., dorsolateral stripe and basal webbing in C. iitciibaiiibciifis). Status of Pliyllohatcs pcnivimuis Mclin Melin (1941) ciescribeci Pln/llohalcs pentvimuii from Roque, Departamento de San Martin, Peru; the species was placed in Coloftcthii^ by Edwards (1971). The de- scription was based on one male (NHMG 508) having a SVL of 21 mm and various characters mentioned by Melin (1941:61-62), "first finger about as long as second, Otym- paninn parth' indistinct, about one-third diameter of the eye, O discs of fingers and toes fairly small, toes webbed at base." Melin (1941:62) noted various aspects of the col- oration in preservative: "a wedge-shaped blackish medial band on the back beginning at the eyes; another dark band from the nostril through the eye and above the tympanum towards the base of the forelimb, mecliallv edged bv a light band, continuing towards the base of the hind limb; sides of the upper jaw transversely banded; flanks blackish with light spots; limbs with diffuse cross bars; hinder part of the thighs blackish with light marble and dots; feet above light; body beneath whitish with a fine blackish mottle or dotting; hinder part of bellv and tliighs beneath whitish." The "light band" extends from the orbit to the inguinal region and was defined as an oblique lateral stripe by Morales (1994), who examined the holotype and noted that dorsolateral and ventrolateral stripes were absent. Roque is at an elevation of about 1200 m on the Amazo- nian slopes of the Cordillera Central in the Departamento de San Martin; thus, it is about midway between the type localities of Colo^tcthu^ idioiiichis and C. fiiittciiiicicri at Venceremos and the type localities of C. cm^pciioct'pi^ and C. soniiiliUii^ at 30 km SW of Zapatero. Coloftctliiis niiiiciiucicri lacks pale stripes, and C. crnspcdoccps and idioiiicltis have dorsolateral stripes and short or no oblique lateral stripes; furthermore, C. cm^^pcdoccps, also differs from C. pcniviaiiiis by having Finger I longer than Finger II. Cokvtctluis sordida- tub is like C. pciuz'itDius in having an oblique lateral stripe, no dorsolateral stripe, and Fingers 1 and II equal in length, but it differs from C. pcriii'ianus in having the toes about three-fourths webbed, as contrasted to basal webbing in C. periiviaiuis. Furthermore, adult males of C. sordidatiis have SVLs of 24.2-29.9 (x = 25.7, ,V = 9) in contrast to 21 mm in the holotype of C pcnii'iaiuib, an adult male transporting tadpoles. Thus, Colostetluib pcniviniiub remains an enigma. Since the capture of the t\'pe specimen in 1925, no additional specimens have been found. DISCUSSION LlFt HISTORY AND TaDPOLES Like most other dendrobatid frogs, with the possible exception of Ammobates, members of the genus Colostethus typically deposit their eggs amidst leaf litter or soil on the forest floor; an exception is C. beebei which deposits eggs on the leaves of terrestrial bromeliads (Bourne et al., 2001 ). Eggs usually are attended by males; upon hatching, the tadpoles wriggle onto the back of the attendant male, who transports them to water. However, egg attendance and transportation of tadpoles is by females in C. iii^iiiiinlif^ and 42 SciHNTiHc Paphrs, Naturai. History Museum, Tin Um\ hrsity of Kansas C. pynitti (Grant, 2004). Both sexes of C. talanumcnc have been reported to transport tadpoles (Breder, 1946; Silverstone 1976). Of six Colostethus elachyhistus that were transport- ing tadpoles, one from Avabaca, Departamonto de Piura, is a gravid female and five from other k)calities are males. These limited data suggest that lar\ al transport (and prob- ablv egg attendance) is amphisexual in C. chuin/libtits, or that the populations represented are different species. The tadpoles of at least four species of Colostethus are nonfeeding; C. chalcopis on the island of Martinique, and C. nidkola, and C. stephcni in Amazonian Brazil are nt)t transported; instead they are nidicolous and complete their development in terrestrial nests (Caldwell and Lima, 2003; Kaiser and Altig, 1994; June- et al., 1994). The tadpoles of C. degranvillei also are nonfeeding and complete their de\'elopment on the back of an attendant male (Claessen, 2002; Lescure, 1975). Clutch size in species having transported, feeding tad- poles is 12-tO (Coloma, 1995; Lima et al., 2002; L.ddecke, 1999; Wells, 1980). Caldwell and Lima (2003) summarized data on clutch parameters in three nidicolous species of Colostethus; ovarian complements are 1-6, and diameter of ovarian eggs is 2.0-2.7 mm. The presence of only six unpigmented eggs about 2.0 mm in diameter in a female of C. oniatua in Pem indicates that this may be another nidicolous species. Growth of back-riding tadpoles has been reported for Colostctlius stihpiuictatus by Stebbins and Hendrickson (1959) and C. piviamen^i^ (as C. inguinalis) by Wells (1980). Back-riding tadpoles in 13 lots of six species (Co/os/c/Z/n.'^ cmspcdoccps. clacln/liistii^, ek'ntlwrodactylu^, idionifhi^, iiexi- ;)«s, and !^i/lz>aticiis) are in Stages 24 or 25; these tadpoles range in bodv length from 3.3- 4.9 mm and in total length from 8.7-17.4 mm. The smallest back riding tadpoles ha\e an undifferentiated gut, and the body cavity is filled \\ ith yolk; larger tadpoles have coiled intestines and commonly still contain yolk. The normal secquence of ontogenetic development of labial tooth rows in anurans is 1/0, 1/1, 1 /2, 2/2, 2/3 (Altig and McDiarmid, 1999). The smallest tadpoles of C. elacln/hiftiis, idioiuelus, and iiexipuf have LTRFsofO/0. InC. elacln/histus, ekuthewdactylus, and nexipiis, the ontogenetic sequence in LTRF in back-riding tadpoles is 0/0, 1/0, 1/1, 2/1, 2/2. (See species accounts and Table 3.) The full complement of labial teeth (2/3) apparently is not developed until the tadpoles become free swimming. In contrast, the ontogenetic sequence in back-riding tadpoles of C. idiomelus is 0/1, 0/1, 0/2, 0/3; all free-swimming tadpoles have an LTRF of 2/3. Neither of these sequences is like the "normal" sequence stated by Altig and McDiarmid (1999). It is unclear why tadpoles of Colostethus should differ in the ontogenetic sequence of tootli-row de\'elopment. 46 44 42 40 0) D) TO 55 38 2 S 36 E Q. O "53 34 > Q 32 30 28 26 C. elachytiistus C. Insulatus C. nexipus C. leucophaeus C. sylvaticus C idiomelus 10 15 20 25 30 Mean Length (mm) 35 40 45 50 Fig. 15. Growlh tr^ijectorios of t.idpok's ol si\ species of Cclostflliii^. COLOSTETHUS OF NORTHERN PkRU 43 Tadpoles of Colofitctlni$ in the Andes of northern Peru described herein are rather nondescript larvae; all taxa are similar in general shape and proportions, and none has distinctive coloration (Fig. 9). The largest tadpoles are those of C. Iciicoplincus with a body length of 20.5 mm and a total length of 50.2 mm (Stage 40); the smallest are those of C. iie.xipus with a body length of 10.3 mm and a total length of 28.2 mm (Stage 41). This is apparent in the growth trajectories of the two species; the tadpoles ot other species are intermediate in their growth trajectories (Fig. 15). ECOIOGY Most species of Colostcthub are active bv day in and along small streams. The nature of the streams differs among some species. Colostctltiis idionwlu^ and C. Iciico- phaeus were found in marshv streams or spring seepages, and C. iiiittciiiicicri was in a small mossy stream flowing from a nearby spring. Cokvtctlni^r^acnigiiwsus, craspedocepf, iiisiilntus, iiexipiis, piilchcrriiiins, soniidatus, and fi/lvatiais were found only in rocky streams, and C. cincln/histiis oc- curs in rockv streams and irrigation ditches. Coloftcthiis eleuiherodactyliis, oriiahis, and spiloiogastcr were found in areas of leaf litter on the forest floor. No ecological data are available for C. av^yrogiTT^tcr, poccilonotus, and utcubambciisib. Only seven instances of sympatry are documented among Colostcthiis in the Andes of northern Peru (Fig. 16), and for the locality 20 km south of Chiriaco, no ecological data are available. With the exception of the locality 16 km ESE of Shapaja, significant differences in snout-vent length exist between the sympatric species. At the exception, C. ncxipiis inhabits streams, whereas C. clcii- thcivdact}/liis was found on the forest floor litter. Also, at Ponga de Shilcayo, C. )iexipit$ inhabits the stream, whereas C. onintiis was found on the forest floor litter. However, Coloma (1995) noted that five C. iwxipits were under pieces of wood inside a house close to a swampv area in Limon, Ecuador; other Ecuadorian specimens were from streams. At the rocky ravine 30 km SW of Zapatero, the small C. craspcdoceps was active on small boulders in the stream by day when individuals of the larger C. sordidntiif were found only under stones at the edge of the stream. However, at night, C. fordidntii> were perched on the boulders in the stream, and no C. crnspcdoccps were observed. Tadpoles of Colo^tctliiis primarily inhabit quiet pools in streams. At least in two cases, tadpoles of sympatric species differ slightly in size. On the western slope of the Cordillera de Huancabamba, tadpoles of C. cincln/hiftiis in Stage 35 have total lengths of 32.7-36.6 mm (x = 34.58), whereas those of C. si/lvnticus in Stage 35 have total lengths of 38.1-39.3 mm (x = 38.56). Likewise, at 30 km SW of Abra Pardo de Miguel, 2180 m tctluis at seven localities in northern Peru. Horizonal lines are ranges and \ertical lines are means. Zapatero, tadpoles of C. craspedoceps in Stage 25 have total lengths of 15.9-23.7 mm (x = 20.68), and those of C. sordi- datus in the same stage have total lengths of 19.4-33.5 mm (X = 26.44). These few observations and limited data suggest that sympatric species of Colostetlius may differ in (1) size of adults, (2) size of tadpoles, (3) microhabitat, or (4) time of activity. Obviously, these possibilities need to be examined in detail. 44 Scientific Papers, Natlikai. History Museum, The University oi' Kansas Tabic 12. Geographic and elevational distribution of species of Cokvtclhiis in northern Peru. Numbers arc clc\ations in meters. Cordillera Cordillera Cordillera de Marancin Cordillera Eastern Species Occidental Colan Huancabamba Valley Central Foothills C. nerugiiwfu^ _ — 1980-2180 C. argyrogn^^tcy — — — 525-550 — 500-600 C. crnspedoceps — — — — — 500 C. elachi/hiitus 725-2760 — 710-2590 — — — C. eleutlierociach/luf — — — — — 360 C. idiomelus — — — — 1620-2840 — C. iiisulatuf — — — 1260-2600 — — C. leucopliaeu^ — — — — 2400 — C. iiiittermt'icri — — — — 1620-2050 — C. nexipus — ■ — — — 325-810 360 C. oniatus — — — — 470-680 350 C. poecilonotus — — — 500 — — C. ptilchcrninu^ 2620 — — — — — C. sordidatus — — — — 520 500 C. tipilotogaster — 2323 — — — — C. fyli'dticus 2460 — 1920-3100 — — — C. tilcubambeiidiuli/lu^) apparently are restricted to elevations of less than 500 m at the base of the mountains. Two others (C n}\^}/iogii>tcy and C. sordidntus) occupy the same elevational range at the eastern base of the mountains, and C. nexipus has a greater elevational range (520-810 m). These three species also occur at elevations of about 500 m on the drier western slopes of the Cordillera Oriental. If there is, or has been, a continuitv in their distributions, presumably it is around the northern end ot the Cordillera C'entral and not across the high elevations of the cordillera. The other four species in the Corciillera Central are restricted to higher elevaticms in the cordillera; Colostclhiis idiomelus and C. initlcnncicri descend to I(i20 m on the eastern slopes of the Cordillera Central. The former also ranges westward in the Cordillera. Colostcthusacnigiiiosiis is known only from high ele\'ations (1980-2180 m) on the humid eastern slopes, whereas C. Icucopliaciis is known on\\ from 2400 m in the highlands. One species, C. spilotogaster, is known from an ele\ation of 2325 m in the Cordillera Colan, a northern extension of the Cordillera Central. In contrast to the Cordillera Central, onh' three species of Colostethus are known from the Cordillera Occidental and its associated ranges (Cordillera de Cuchalli, Cordil- lera de Huancabamba, and Cordillera de Tabaconas). Co- lostcthiif cinchyhistiis is widespread geographically (Fig. 9) and is known from elevations of 725-2590 m. Colostetlnis si/lvaticus is more restricted geographically at elevations of 1920-3100 m, whereas C. puhiwrriimis is known from a single locality at 2620 m. Except for isolated areas at higher elevations, the western slopes of the Cordillera Occidental are dry with few permanent streams; consequenthv these slopes lack the diversity of Colostctlnis that is e\ident on the humid eastern slopes of the Cordillera Central. Colostethus citicln/liistiis occurs on the high (2020 m) Pacific slopes and eastern slopes of the Abra de Porculla, the lowest pass in the Cordillera Occidental, and at 2500 m in the upper \'al- ley of the Rfo Reque. Most areas in the arid \alle\- of the Rfo Marafion seem to be uninhabitable b\' Colostethus, but numerous small streams that plunge down the western slopes of the Cordil- lera Central and eastern slopes of the Cordillera Occidental pro\ ide adequate habitat for these frogs. Three species (C. iusulatus, poccilouotus. and utcubiuidh'iisis) are known onl\- from these tributaries to the Ri'o Marancin. As noted pre\ ioush', three other species (C. iirgi/rogaster, m'xipus, and sordidiitus) tiiat occiu' in the eastern foothills of the Cordil- lera Central presiunabh' entered the more northern part of the Rfo Marani'in \'alle\' around the noithern end ol the Cordillera Central. COI-OSTETHVS OF NORTHERN PeRT 45 C. aeruginosas C. argyrogaster C. craspedoceps C. elachyhistus — C. eleutherodactylus C. idiomelus C. insulatus C. leucophaeus C. mittermeieri C. nexipus C ornatus - C, poecilonotus C. pulcherrimus C. sordidatus C. spllotogaster C. sylvatlcus C. utcubambensis --- West 3000 2000 1000 Meters + ■ Fig. 17. Profile of tlie Andes at about 6 S Lat. in northern Peru; vertical exaggeration is about 33X. t^lack bars indicate distributions of species of Colostcthm. The ranges of C. :^piloto^astcr and C. si//i'iif(t i(s are at about 5 '30'. LITERATURE CITED Altig, R., and R. W. McDiarniid. 1999. Body plan; development and mor- phology. Pp. 24-31 111 R. W. McDiarmid and R. Altig (eds.). Tadpole^. Chicago: Univ. Chicago Press. Barbour, T., and G. K. Noble. 1920. Some amphibians from northwestern Peru, witli a revision of the genera Pln/IlobiUc^ and Tiiiiuitoluu>. Bul- letin of the Museum of Comparative Zoologv, Harvard Universitv 63:395-427. Bourne, G. R., Collins, A. C, Holder, A. M., and McCarthy C. L. 2001. Vocal communication and reproductive behavior of the frog Colostctlms beebei in Guyana, fournal of Herpetology 35:272-281. Breder, C. M Jr. 1946. Amphibians and reptiles of the Rio Chucunaque drainage, Darien, Panama, with notes on their life histories and habits. Bulletin of the American Museum of Natural Historv 86:375-436. Caldwell, J. R, A. P Lima, and C, Keller 2002. Redescnption of Colo- sti'tbiis niarchcsiauii^ (Melin, 1441) from its type locality. Copeia 2002:157-165. Caldwell, J. P., and Lima, A. P. 2003. A new Amazonian species of Colo- stethus (Anura: Dendrobatidae) with a nidicolous tadpole. Herpe- tologica 59:219-234. Claessen, H. 2002. Een niewe reprocdiemodus liij kickers van het geslacht Cohstethus. Lacerta 58:107-108. Coloma, L. A. 1995. Ecuadorian frogs of the genus Colo^tcthui. (Anura: Dendrobatidae). Miscellaneous Publication Natural Hi>-tor\- Mu- seum University of Kansas 87:1-72. Duellman, W. E. 1987. Two new species of marsupial frogs (Anura: Hy- lidae) from Peru. Copeia 1987:903-909. Duellman, W. E., and J. B. Pramuk. 1999. Frogs of the genus Eleuthero- dactylus (Anura: Leptodactylidae) in the Andes of northern Peru. Scientific Papers Natural Histor\' Museum The Universitv of Kansas 13:1-78. Duellman, W. E., and R. Schulte. 1993. New species of centrolenid frogs from northern Peru. Occasional Papers Museum of Natural Historv University of Kansas 155:1-33. Duellman, W. E., and J. E.Simmons. 1988. Two new species of dendrobatid frogs, genus Co/csfi'f/iii.s from the Cordillera del CUndor, Ecuador. Proceedings of the Academv of Natural Sciences of Philadelphia 140:115-124. Duellman, W, E., and E. R. Wild. 1493. Anuran amphibians from the Cordillera de Huancabamba, northern Peru: systematics, ecology, and blogeography Occasional Papers Museum of Natural History University of Kansas 157:1-53. Edwards, S. R. 1971 . Taxonomic notes on South American Cohstethus with descriptions of two new species (Amphibia, Dendrobatide). Proceed- ings of the Biological Society of Washington 84:147-162. Frost, D. R. 1986. Anew Colostetlius (Anura: Dendrobatidae) from Ecuador Prcxeedings of the Biological Society of Washington 99:214-217 Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica 16:183-190. Grant, T 2004. On the identities of Colostctiius in^uiiwlis Cope, 1868) and C. paihinu'iisis (Dunn, 1933), with comments on C. Intiuasus (Cope, 1863). American Museum Novitates 3444:1-24. Grant, T E., and M. C. Ardila-Robayo. 2002. A new species of Colostetlius (Anura, Dendrobatidae) from the eastern slopes of the Cordillera Oriental of Colombia. Herpetologica 58:252-260. 46 Scientific Papkrs. Natural History Museum, The University of Kansas Grant, T, and F. Castro. 1998. The cloud forest Coloslcllnif (Anura: Den- drobatidae) of a region of the Cordillera Occidental of Colombia. Journal of Herpetology 32:378-392. Grant, T., E. C. Humphrey, and C. W. Myers. 1997. The median lingual process of frogs: a bizarre character of Old World ranoids discovered In South American dendrobatids. American Museum Novitates 3212:1-40. Grant, T, and L. O. Rodriguez. 2001. Two new species of frogs of the genus Coloslctliuf (Dendrobatidae) from Peru and a redescription of C. (n'/iiicn/KS (Boulenger, 1883). American Museum Novitates 3355:1-24. Ibafiez, R., and E. M. Smith. 1995. Systematic status of C()te(f//ji(s flotu- tor and C. nubicotn (Anura: Dendrobatidae) in Panama. Copeia 1995:446-456. June-, F. A. 1998. Reproductive biology of Colostetlnis steplieni and Colo- stetliuf marcliesiaiiuf (Dendrobatidae), with the description of a new anuran mating behavior. Herpetologica ,54:377-387. June-, F. A., R. Altig, and C. Gascon. 1994. Breeding biology oi Colostctlius stephfni, a dendrobatid frog with a nontransported nidicolous tad- pole. Copeia 1994:747-750. Kaiser, H., and Altig, R. 1994. The atypical tadpole of the dendrobatid frog, Colcslcthiii chalcosis, from M (7i'i(//u'Mii/i7ch//i(S PERU: Still Mniii'ii 16 km ESE Shapaja, 360 m, Provincia de San Marh'n, KU 211813, 211814 (holotype) Colostcthuf idiomeliif PERU; Aiiiiizoncis: 2 km E Balzapata, 2220 m, Provincia de Bongara, KU 211891-905, MHNSM 6231-35; 8 km NNE Balzapata, 1850 m, Provincia de Bongara, KU 181680; 12 km NNE Balzapata, 1940 m, KU 211890, 215604 (tadpoles), MHNSM 6223-24; 11 km SSE Chachapoyas, 2840 m, Provincia de Chachapoyas, KU 21 1884; Pomacochas, 2150 m, Provincia de Bongara, KU 181681-99, 181871- 72 (tadpoles), 211885-89, 215605 (tadpoles), MHNSM 6228-30. 6272 (tadpoles); 5 km W Pomacochas, 2228 m, LSU 19576-83. Sail Martin: East slope Abra Pardo de Miguel, 1980 m, Provincia Rioja, KU 215608 (tadpoles); East slope Abra Pardo de Miguel, 2180 m, Provincia Rioja, KU 211906-39, 215606-07 (tadpoles); Venceremos, 1620 m, Provincia Rioja, MCZ A-100260 (Holotype). Colo>lcllui> iii'iulatii^ PERU: Amazoiia^: 16 km ENE Balsas, Provincia Chachapoyas, 1650 m, UF 37536-41; 17 km ENE Balsas, Provincia Chachapoyas, 1810 m, KU 211857 (holotvpe) 211858-70, 211560 (tadpoles), MHNSM 6206-14, 6274 (tadpoles); 24 km ENE Balsas, Provincia Chachapoyas, 2160 m, KU 211871-72; 37 km ENE Balsas, Provin- cia Chachapoyas, 2600 m, KU 211873-74; 33 km SE Ingenio, 1850 m, LSU 32548-51; 6 km W Pedro Ruiz, Provincia Bongar-, 1260 m, KU 211875-78, 215601 (young), 215602 (tadpoles), MHNSM 6219-21, 6269 (tadpoles), 6284 (young). Cajaiiiarcn: 11 km W Balsas, Provincia Celendin, 1600 m, KU 211853-56, 215599 (tadpoles), MHNSM 6266-68, 6276 (tadpoles); 2 km NW San Juan, 2290 m, KU 211833-42, 215598 (young), MHNSM 6257-65. C(>/(isfi'f)/»s Icucofiliacus PERU: Aniazoiia^: Molinapampa, Provincia Chachapovas, 2400 m, KU 211879 (holotype), 211880-83, 215603 (tadpoles), MHNSM 6215-17, 6271 (tadpoles). ColosUihiis iiiillcniicicn PERU: San Martin: East slope Abra Pardo de Miguel, 2050 m, Provincia Rioja, KU 211944; Venceremos, Provincia Rioja, 1620 m, MCZ A-100217 (holotype), MCZ A-100218-57. Colostetlius iicxipii> ECUADOR: Morona Saiitia;io: 5.3 km NE Plan de Milagro, 2550 m, KU 202822-24; Mendez, 500 M, KU 220664; Limon, 1090 m, KU 220661-63; LosTavos, KU 194164. PERU: Ainazwiai^: 43 km NE Chiraco, 325 m, LSUMNS 53430; 20 km SW Chiriaco, 520 m, KU 196727-28, 196730; LSUMNS 32547. San Martin: Cataratas Ahuashiyacu, 14 km NE Tarapoto, Provincia de San Martin, 730 m, KU 21 1805-10, MHNSM 6200-05; Cordillera NE of Tarapoto, Provincia de San Martin, 600 m', KU 209397-98; Ponga de Shilcayo, 4 km NNW Tarapoto, Provincia de San Martin, 470 m, KU 211830; 6 km ESE Shapaja, Provincia deSan Martin, 310 m, KU 212486 (tadpoles); 14 km ESE Shapaja, Provincia de San Martin, 360 m, KU 211815-29, KU 215594 (tadpoles); 15 km ESE Shapa|a, Provincia de San Martin, 360 m, KU 211811-12; 16 km ESE Shapaja, Provincia de San Martin, 360 m, KU 215593; 17 km NE Tarapoto, Provincia de San Marhn, 850 m, MHNSM 6199; 22.7 km NE Tarapoto, Provincia deSan Martin, 810 m, KU 211831; 215596 (tadpoles); 23.2 km NE Tarapoto, Provincia de San Martin, 800 m, KU 215595; 28 km NE Tarapoto, Provincia de San Martin, 600 m, KU 211832. Colostetlius ornatiis PERU: San Martin: Ponga de Shilcayo, about 4 km NNW of Tarapoto, Provincia de San Martin, 470 m, KU 211950.Tarapoto, Provincia de San Martin, 350 m, MHNSM 17713-17. Cotcitctlnis periivianiis PERU: San Marlhi: Roque , NHMG 508 (holotype). Coloftcthus pcccilonotuf PERU: .4;m?:('//(!s: Alva, 500 m, MCZ A-891 06-07 A-891 08 (holotype), A-89109. Colostetlius piilchcrrnniis PERU: Cajaniarni: Cutervo, Provincia de Cutervo, 2620 m, KU 211946 (holotype), 211947-49, MHNSM 6251, Colosltilius soniiiiatub PERU: Ainazonas: 20 km SW Chiriaco, 520 m, Provincia de Bagua, KU 196718-26. San Martin: 30 km SW of Zapatero ( 1 km N E of San Jose deSisa), 500 m, Provincia de Lamas, KU 211960 (holotype), 211961-72, 215611 (tadpoles). Colostctlnis spiloto^astcr PERU: Awazonas: Cordillera Colan, SE La Peca, 2326 m, LSUMNS 39341 (holotype). Coloitctlius si/limticus PERU: Cajaiiiarca: Tabaconas, Provincia de San Ignacio, MCZ 5344 (holotvpe), 5346, 5350, 5355-56. Pima: 17.7 km E Canchaque, 1920 m, UF 112484-89; 21 km E Canchaque, 2070 m, UF 112458; 24 km E Canchaque, 48 Scientific Papers, Natural History Museum, The University oh Kansas 2240 m, UF 112494-98; 30 km E Canchaque, 2530 m, UF 52040-42, 1 12499-310; 30.6 km E. Canchaque, 2590 m, UF 52141-44, 112490-93; 12.7 km E EI Tambo, 2820 m, KU 219756-58; 1 km W El Tambo, 2740 m, KU 219755; 15.5 km SVV Huancabamba, 2560 m; 181670 (tadpoles); 25.5 km SVV Huancabamba, 3010 m, KU 181668-73, 181868-69 (tadpoles); 29.3 km SVV Huancabamba, 3010 m, KU 181667; 31 km SSW Huancabamba, 3080 m, KU 181674-79; summit of Cordillera between Canchaque and Huancabamba, 3100 m, KU 138071-79. Co/os/t>f/ii(.s iilcuhambetisis PERU: Anmztmas: Tingo, MHNSM 13986 (holotype), 13987-91. APPENDIX 2 Gazetteer Following is a list of locaiitiL-s in the Andes and adjacent lowlands of northern Peru where specimens of Colo^tctlius were collected. After each place name the departamento is given in parentheses, followed by geographic coordi- nates, elevation and vegetation type. Coordinates were determined from maps (Mapa Fi'sico Politico del Peru, 1:1,000,000, 1973, and Carta Nacional del Peru, 1:100,000, 1986. Elevations were obtained from altimeter readings or from maps. When known, specific sites and habitats are given followed by names of collectors (only those responsible for field notes) and the months and years that they collected at the sites. Names of collectors are abbre- viated: ERW = Erik R. Wild, GKN = G. K. Noble, LSU = Louisiana State University ornithologists, RAM = Russell A. Mittermeier, RS = RanierSchulte, RT = Richard Thomas, THF = Thomas H. Fritts, WED = William E. Duellman. All localities are shown on the political map (Fig. 1). Ahrn de Pomilln (I'iura)— 05 50' S, 79 '30' W, 2145 m; humid montane forest. Pa.ss in the northern part of the Cordillera Occidental. Collections made on southwestern slope (2020 m) and northeastern slope (20S0 m) in March 1975 by WED. Ahrii Pindo dc Miguel ( Amazonas-San Martin)— 05 46' S, 77 42' W, 22 1 II m; very humid montane forest. A pass in the eastern range of the Cordillera Central. Collections made on the east slope just below the crest on the road to La Rioja at an elevation of 2180 m and farther down the slope at 1980 in January 1989 by WED. Alva (Amazonas)— 05"53' S, 78"56' W, 1000 m; presumably thorn forest. Site on slopes of Cordillera Central above Pedro Ruiz Gallo In Rio Utcubamba Valley. Collection in May 1974 by RAM. Ayahaca. (Piura)— 04'38' S, 79"43' W, 27.50 m; mostly cutover humid montane forest. A small town in the northernmost part of the Cordillera Occidental in Peru. Collections made in the vicinitv of the town in Febru- ary 1989 by WED. Brt/srts (Amazonas)— 06"50' S, 77"5'-r W, 1090 ni,; thorn forest. .^ vil- lage in the narrow, subhumid Marafion Valley. Collections made at small streams and waterfalls at elevations of 1810, 2160, and 2600 m along the road ascending the western face of the Cordillera Central toward Chachapoyas in January 1989 by WED. Balzapnta. (Amazonas)— 05"46' S, 77"5r W, 1 640 m. A village just west of the crest of the Cordillera Central. Collections made in disturbed and partly cutover humid forest 8 km NNE (1850 m) on the road to La Rioja in March 1979 by WED and at 2 km E (2200 m) and 12 km NNE (1940 m) in January 1989 by WED, Cajahnmba (Cajamarca) — 07 36' S, 78 02' W, 2700 m; montane dry forest. Town in the Cordillera Central. Collections made in April 1970 by THF and in at 1 km south (2650 m) in March 1989 by WFD. Camhaqw (Piura)— 05'22' S, 79-36' W, 1 120 m; tropical dry forest. A town near the western base of the Cordillera de Huancabamba. Ci>llec- tions were made in drv forest at Canchaque in May 1970 bv THF and in March 1979 by WED, 5 km W (710 m) in December 1978 by RT 6.5 km west (620 m) in January 1991 by ERW, in humid montane forest 12 km east (1770 m) on the road to Huancabamba in March 1979 by WFD, and 15 km east in December 1978 by RT. CatamhK AhuaihittacK (San Martin)— 06" 30'S, 76 '20' W, 730 m; humid subtropical forest. Waterfall and rocky stream in deep ravine 14 km NNE of Tarapoto on the road to Yurimaguas. Collections in February 1989 by WED. Clinchapo\ias (Amazonas) — 06"13' S, 77"50' W, 2360 m; disturbed humid montane forest. Town in the Cordillera Central. Collections made in humid montane forest at an elevation of 2340 m 11 km SE of town in January 1989 by WED. Chirmco (Amazonas)— 05''09' S, 78"21 ' W, ±450 m; thorn forest. Town on the Ri'o Marafion at the confluence of the Ri'o Chiriaco. Collections made in dry forests at 43 km NE in October 1978 by LSU and at 4 km SW and 20 km SVV (525 m) on the road to Bagua in December 1974 by RT. Cutervc (Cajamarca)— 06 22' S, 78 49' W, 2620 m; cultivated valley in the northern part of the Cordillera Occidental. Collection in February 1989 by WED. El Tamho (Piura)— 05°21' S, 79'"33' W, 2770 m; humid montane for- est. Settlement on the western slope of the Cordillera de Huancabamba, 31 km NNE of Canchaque on the road to Huancabamba. Collections in relatively undistvirbed forest at 1 km W (2740 m) and 12.7 km E (2820) in January 1991 by ERW. Hiumaibamhn (Piura)— 05' W S, 79"28' W, 1900 m; dry forest. Tuvn on the Rio Huancabamba at eastern base of the Cordillera de Huancabamba; area in immediate vicinity mostly cultivated. Collections made in town and at summit of Cordillera de Huancabamba (3100 m) in May 1970 bv THF and in humid montane forest at 4 km SVV (1980 m), 15.5 km SVV (2560 m), 25.5 km SW (3010 m), 29.3 km SW (3010 m) and 31 km SVV (3080 m) on the road to Canchaque in March 1979 by WED. himzii (Amazonas) — Morales and Schulte (1994) gave approximate coordinates of 04*15' S, 78'20' in "Bagua Grande, Amazonas." The.se coordinates are well within Provincia Condorcanqul; no place name of Imaza could be found in that region. The Ri'o Imaza is at approximately 05"16'S, 78" 17' Win Provincia Bagua, Departamento Amazonas. Both sites are within the Marafion Valley. April 1990 bv Padre Pedro Cashiro. In^ciiw (Amazonas)— 05"55' S, 78'55'VV, 1280 m; thorn forest. Settle- ment In the Rio Utcubamba Valley. Collection from dry forest on slopes of Cordillera Central 33 kmSE (1830m) on road to Pomacochas in December 1474 by RT. Lanms (Lhinm) (Cajamarca) — 06"3r S, 70 07' W, +2200 m; thorn forest. \illage in valley on west slope of Cordillera Occidental. Collection from roadside ditch 4 km W 2500 m) in March 1989 by WED. Im Pccii (Amazonas)— 05"36' S, 78''22' VV, + 1400 m; thorn forest. Vil- lage near western base of the Cordillera Colan. Collection from humid montane forest SE of village at 2625 m in October 1978 by LSU. Moliuapnmpn (Amazonas) — 06Tr S, 77''38' W, 2400 m; mostly cul- tivated montane dry forest. A small town in the upper Ri'o Utcubamba Valley. Collection in vicinity of tow in January 1989 by WED. 'Pcdw Ruiz Callo (Amazonas)— 05"25' S, 77-57' VV, 1280 m; thorn forest. Small village in the middle Ri'o Marancin Valley. Collection from small stream 6 km W (1260 m) on road to Bagua in January 1989 by WED. Pomacochas (= Florida) (Amazonas)— 05'49' S,' 77",55' W, 2150 m; mostly cultivated humid montane forest. Village in northern part of Cordillera Central. Collections from vicinity of village in March 1979 and January 1989 by WED, and 5 km west (2228 m) in July 1968 by RT Pon^a de SInhayo (San Martin) — 06"3r S, 76"53' W, 470 m; humid tropical forest. Deep ra\ine4 km NWV of Tarapoto. Collection In I'ebru- ary 1989 by WED. COLOSTETHVS OF NoRTHhRN PhRU 49 Rio Cumbaza Valley (San Martin)— ()6"2y S, 76"23' VV, 430 m; tropical forest. Shallow, narrow valley of a tributar\' o the Rio Mayo. Collection made in February 1989 by WED. Roque (San Martin) — 06'24'S, 76'48'W, ± 1200 m; humid montane forest. Village on the Amazonian slopes of the northern part of the Cor- dillera Central. Collection made in 1923 by Douglas Melin. San Andres de Cutervo (Cajamarca) — 06°14' S, 78"43' W, 2850 m; dry montane forest. Village in northern part of Cordillera Occidental. Collec- tion from cave near village. San Juan (Cajamarca) — 07"15' S, 78"30' W, 2290 m; montane dry forest. Town in the upper Ri'o Magdalena Valley on the west slope of the Cordillera Occidental. Collection from 2 km N on road to Cajamarca in January 1989 by WED. Santa Cruz (Cajamarca)— 0605' S, 78 51' W, ±1300 m; thorn forest. Village in the Rio Chamaya Basin. Collection from 28 km N (725 m) in December 1974 by RT. Shapaja (San Martin) — 06"36' S, 76"17' W, 310 m; mostly cutover humid tropical forest. Village just north of the Rio Huallaga. Collections made at 6 km ESE (310 m) and 14-16 km ESE (360 m) on the road to Chazuta in February 1989 by WED. Tahaeonas (Cajamarca)— 05°19'S, 79°17'W, ± 2000 m; humid tropical forest. Village in Cordillera de Tabaconas. Collection made in September 1916byC;KN. Tarapoto (San Martin) — 06'3r S, 76°23' W, 370 m; humid tropical forest. Town near the confluence of the Rio Mayo and Rio Huallaga just east of the Cordillera Central. Collections made in outlying ridge of the Andes at 12 km NE (720 m), 22,7 km NE (810 m), 23.2 km NE (800 m), 28 km NE (600 m) on the road to Yurimaguas in February 1989 by WED. Tingo Amazonas) — 06'2r S, 77"49' W, 1835 m; cultivated dry forest. Village on Rio Utcubamba. Collection made in November 1980 by RS. Venceremcis (San Martin) — 05"44' S, 77'3r W, 1630 m; humid montane forest. Former road camp and now small settlement on the road between Balzapata and La Rioja. Collection made in September 1978 by RAM. Zapatero (San Martin)— 06'34' S, 76'30' W, 320 m; humid tropical forest. Village at base of hills north of the Rio Huallaga. Collections from ravine in cutover humid tropical forest at 30 km SW (500 m) in February 1989 by WED. 2044 072 228 679 PUBLICATIONS OF THE NATURAL HISTORY MUSEUM, THE UNIVERSITY OF KANSAS The University of Kansas Publications, Museum of Natural History, beginning v\ ith Volume 1 in 1946, was discontinued with Volume 20 in 1971. 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