HERP MUS. COMP, ZOOL LIBR Ar QL JW @ 7 1981 668 as ,E257 :RSITY OF KANSAS MISGEKLANE@US> > | L93. “UM OF NATURAL HISTORY tan Ce ~ Leptodactylid Frogs of the Genus Eleutherodactylus in the Andes of Northern Ecuador and Adjacent Colombia By ‘John D. Lynch Pe PCT ee UNIVERSITY OF KANSAS LAWRENCE 198] UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly published in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publications, Museum of Natural History, began with number 1 in 1946. Longer research papers are pub- lished in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subjected to critical review by intra- and extramural specialists; final acceptance is at the discretion of the Director. Institutional libraries interested in exchanging publications may obtain the Occa- sional Papers and Miscellaneous Publications by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas 66045. Individuals may purchase separate numbers of all series. Prices for all publications of the Museum may be obtained from the Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66045. THE UNIVERSITY OF KANSAS Museum OF NATURAL HIsTOoRY July 8, 1981 Leptodactylid Frogs of the Genus Eleutherodactylus in the Andes of Northern Ecuador and Adjacent Colombia By Joun D. Lyncu Professor, School of Life Sciences The University of Nebraska, Lincoln, Nebraska 68588 Associate, Division of Herpetology Museum of Natural History, University of Kansas Lawrence, Kansas 66045 THE UNIVERSITY OF KANSAS LAWRENCE 1981 “ UNIVERSITY OF KANSAS PUBLICATIONS, MusEUM OF NATURAL HIsTORY Editor: Robert M. Mengel Managing Editor: Joseph T. Collins Miscellaneous Publication No. 72 pp. 1-46; 22 figures; 8 tables Published July 8, 1981 MuseuM or Natura History THE UNIVERSITY OF KANSAS LAWRENCE, Kansas 66045 WS -A. PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS CONTENTS ST pTG EG CO) Bw) MS) COE Te NCW) IN ese aka se oe Se Ria sh oh De oe 1 PNG let 1O oye) C1 Grn Sa eh ac ae eee ee 2 MAGIEUENIES PAINT NOP HODS 2 9 2 Ae ee ee ee 2 KEY TO ELEUTHERODACTYLINE FROGS IN PARAMOS OF SOUTHERN COLOMBIA AND NORTHERN ECUADOR _. 2 PN @OMINIS JORIS PEGI S25 Gen ee ee ee 4 Eleutherodactylus backleyt (Boulenger) 4 Eleutherodactylascuntipes. (Boulenger)0 2 ee 7 Pleutherodactylus modipeplus new species -.....- ——-— _..__.__.______._ 22 Eleutherodactylus myersi (Goin and Cochran) __--..-------------------- 25 Elgutherodactylus ocreatus new species 2 26 Pleutherodactyius orcest Viymen 2228 en ne es 29 Bigutherodactylus thymelensis Vameh 30 Eleutherodactylus anistrigatus’ (Gunther) ee 32 POOR CO SS ODI eee ee 36 BION TCAUERY,, enc ccd Nien 9, ae a i ee 2: ROR n eee) ae OA oD 40 BSS MO ANSI 90a od oe SO oi els ee ee 4) ‘Lol TO ETRE BENS GLUT RAT SR. 6] OF De met ea ee eC Nace ian 42 APPENDIX: SPECIMENS EXAMINED INTRODUCTION Although a prominent component of the fauna of most mesic environments in northwestern South America, the genus Eleutherodactylus is poorly represented in the pajonales (=subparamos) and paramos of southern Colombia and northern Ecuador. Field work in these highland areas by the late James A. Peters and his associates and by indi- viduals associated with the Museum of Natural History of the University of Kansas has yielded adequate altitudinal and geographic samples of eleuthero- dactyline frogs to permit ecological and systematic summaries of the frogs of this region. This study is the fourth in a series describing the Eleutherodactylus of Ecuador. The first (Lynch, 1979) treated the fauna of southern Andean Ecuador, the second (Lynch, 1980) sum- marized the fauna of the western Ama- zonian Basin, and the third (Lynch and Duellman, 1980) treated those faunas on the Amazonian slopes of the Andes in Ecuador. The principal area under discussion extends, from the Nudo de Pasto in southern Colombia south to the Desierto de Palmira in southern Provincia Chim- borazo of Ecuador (about 2° S). Pajo- nale and paramo habitats within this area originally ranged in altitude from 3200 to 4800 m (Acosta-Solis, 1968) but now extend to lower altitudes where human activity has removed the high Andean forests (ceja andina) or replaced them with Eucalyptus. Seven species of Eleutherodactylus and one species of Phrynopus are restricted to or primarily distributed within the pajonales and paramos. Six other species [Eleuthero- dactylus chloronotus Lynch, E. leoni Lynch, E. supernatis Lynch, E. trepi- dotus Lynch, E. w-nigrum (Boettger), and Phrynopus peraccai Lynch] are spe- cies of the ceja andina and are occasion- ally found in those pajonales that abut the high cloud forests. Most Andean Eleutherodactylus have restricted distributions. Three of the species found in the Andes of northern Ecuador are exceptions. Eleutherodac- tylus curtipes (Boulenger) ranges from the Colombia-Ecuador border (ca °1 N) south to the Desierto de Palmira (ca 2° S) over an altitudinal range of from 2750 m to at least 4400 m. Its northern limit corresponds in an interesting and sug- gestive way with the political boundary of Colombia and Ecuador but in spite of the efforts of several collectors the species has eluded discovery in Colom- bia. The collectors instead find E. buckleyi (Boulenger) there, a close rel- ative. Eleutherodactylus buckleyi is also found in extreme northern Ecuador but is distributed primarily in Colombia, in the Cordillera Central (to ca 4° N). The third widely distributed species, E. uni- strigatus (Giinther), generally occurs at lower elevations and in somewhat more xeric situations than E. buckleyi and E. curtipes. Eleutherodactylus unistrigatus occurs from the Nudo de Pasto in south- ern Colombia south primarily through interandean Ecuador to the vicinity of Riobamba. The general concordance of the southern termini of E. curtipes and E. unistrigatus coupled with the pres- ence of a distinctive suite of eleuthero- dactylines south of the Desierto de Pa- mira led Lynch (1972a) to postulate a faunal break. Lynch (1972a) suggested that E. buckleyi and E. curtipes were synony- mous. That view is contradicted, how- ever, by the sympatric occurrence of the two frogs over much of Provincia Carchi, Ecuador. The other four species of Eleuthero- dactylus and the only species of Phryn- opus (P. brunneus Lynch) found in high altitude grasslands in northern Ecuador have small distribution areas. These Eleutherodactylus are E. orcesi Lynch, E. thymelensis Lynch, and two species named below. 2 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ACKNOWLEDGEMENTS My field work in Ecuador was sup- ported by grants from the Committee on Evolutionary Biology at the Univer- sity of Kansas (1967), the Watkins Fund of the Museum of Natural History, Uni- versity of Kansas (1968), the Society of the Sigma Xi (1968), the Penrose fund of the American Philosophical Society (1970), and by the University of Ne- braska Research Council (1970, 1977). Travel to museums was supported by the University of Nebraska Research Council. The following curators pro- vided working space at their museums: S. Ayala, A. Grandison, J. Hernandez, R. Heyer, C. Myers, R. Nussbaum, the late J. Peters, P. Ruiz, the late C. Walker, E. Williams, G. Zug, and R. Zweifel. A. Leviton and H. Marx loaned critical material. William E. Duellman provided many amenities and the full facilities of the Division of Herpetology at the University of Kansas over the duration of this study. Rita Ricaurte and Jaime Villa helped me with Spanish. Tom Berger, Dave Cannatella, Pat Fritts, Tom Fritts, Bob Henderson, and especially my wife Marsha shared the disappointments, discomforts, and ex- citement of field work in Ecuador. Fer- nando Ortiz, the late Francisco Ledén Rodriquez, and Eugenia del Pino of The Universidad Catdlica de Quito provided logistic support and encouragement in Ecuador. MATERIALS AND METHODS Terminology follows that of Lynch and Duellman (1980). I have examined 3151 preserved specimens of the eight species discussed in this paper including the type specimens of all names except Hyiodes lehmanni. Throughout the text, specimens are identified by their cata- logue numbers and the appropriate acronym for the museum collection, as follows: AMNH, American Museum of Natural History, New York; ANSP, Academy of Natural Sciences, Philadel- phia; BM, British Museum (Natural History ), London; CAS-SU, California Academy of Sciences-Stanford Univer- sity Collection, San Francisco; FMNH, Field Museum of Natural History, Chi- cago; ICN, Instituto de Ciencias Natu- rales, Museo de Historia Natural, Bo- gota; KU, Museum of Natural History, University of Kansas, Lawrence; LACM, Los Angeles County Museum of Natural History, Los Angeles; MCZ, Museum of Comparative Zoology, Cambridge; MLS, Museo, Universidad de La Salle, Bo- gota; MNHNP, Muséum National dHis- toire Naturelle, Paris; MZS, Museo Zo- ologico della Specola, Fiernze; UIMNH, University of Illinois Museum of Natural History, Urbana; UMMZ, University of Michigan Museum of Zoology, Ann Arbor; USNM, National Museum of Natural History, Washington; UVMP, Museo Parasitologia, Universidad de Valle, Cali. KEY TO ELEUTHERODACTYLINE FROGS IN PARAMOS OF SOUTH- ERN COLOMBIA AND NORTHERN ECUADOR The key presented below will suffice to identify any eleutherodactyline frog encountered in paramo and interandean valley habitats from the Nudo de Pasto in Colombia (ca 1° 30’ N) south to the northern edge of the Nudo de Azuay in Ecuador (ca 2° 30’ S) even though some of the included taxa rarely penetrate paramos. Figures 1 and 2 are provided as aids to the key. 1. Toes bear discs on ventral surfaces Toes lack discs 2... - 2) 2. Adults less than 22 mm SVL; outer metatarsal tubercle about same size as inner metatarsal tubercle _. oe Ae wees nce Phrynopus peraccai Adults more than 25 mm SVL; outer metatarsal tubercle much smaller than inner metatarsal tubercle a ee eee Phrynopus brunneus 3. Skin of venter smooth; first finger longer ‘than second. .2 43. 23 3 Wee feet Eleutherodactylus w-nigrum 10. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 3 Skin of venter areolate; first finger shorter than second 2-2 =... 4 . Tympanum concealed beneath skin ee 5 ee eee eee 5 iympanumevisible 24. vk . Digits bearing dilated pads (Fig. 1B) — Eleutherodactylus thymelensis Digital tips narrow (bulbous) (Fig. 1A) 6 . White stripe on lip; cranial crests evident — Eleutherodactylus curtipes Lip barred; no cranial crests __..___. Eleutherodactylus ocreatus . White or cream stripe on upper lip eevee eas 2 Eleutherodactylus buckleyi No cream stripe on lip —__________ 8 . Digital pads narrow (Fig. 1A) __ 9 Digital pads dilated (Fig. 1B) _— 11 . Vomerine teeth present on elevated odontophores Vomerine teeth usually absent, no pmomiophore, 24.22 2e0 Ad ei F 3 Eleutherodactylus myersi Upper eyelid bearing several conical NWAEUS) 2-2 Eleutherodactylus leoni HIE 12. 13. 14. Upper eyelid smooth to shagreened st ee Eleutherodactylus trepidotus Canthus rostralis sharp, strongly concave (Fig. 2) Eleutherodactylus chloronotus Canthus rostralis round to sharp, SLU Ose ees ceo ee ae 12 Skin of back and flanks coarsely areolate _. Eleutherodactylus orcesi Skin of back and flanks smooth or shagreened with or without scattered RVG DES oe oe eee ere te eel ee 13 Distinct tubercle on heel; males lack VOKCall slits heute. se 5ecs A) Bo Ce eee. eee Eleutherodactylus supernatis Heel lacking distinct tubercle; males with vocal slits and external vocal SAC fiend Le oe ee 14 Posterior surfaces of thighs brown with or without cream reticulation; no tubercles on upper eyelid — Eleutherodactylus unistrigatus Posterior surfaces of thighs colorless in preservative (vermilion in life); low tubercles on upper eyelid —__ ees: Eleutherodactylus modipeplus Fic. 1. Hands of (A) Eleutherodactylus curtipes, 10 km SW Mocha, and (B) E. sp., KU 169030. Scale equals 2 mm. MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 2. Dorsal and lateral views of Eleutherodactylus sp. (KU 169030), scale equals 5 mm. ACCOUNTS OF SPECIES In the following accounts I attempt to provide comparable statements in the diagnoses and descriptions. The diag- noses consist of 14 numbered statements (or sets of statements) concerning the important characteristics of eleuthero- dactyline frogs followed by a phenetic diagnosis where I distinguish the taxon from those species it most nearly resem- bles. Measurements and proportions are given for each sex when the means differ significantly (p < 0.05); otherwise these data are combined. Descriptions are not provided for E. myersi, E. orcesi, or E. thymelensis; the original descriptions remain adequate (Goin and Cochran, 1963; Lynch 1972b). Eleutherodactylus buckleyi (Boulenger) Fig. 3A-B Hylodes buckleyi Boulenger, 1882:217 (syn- types, BM 78.1.25.40-44, collected at Intac, Provincia Imbabura, Ecuador, by Mr. Buck- ley; BM 78.1.25.40, an adult female, is here designated the lectoholotype). Diagnosis.—(1) skin of dorsum bear- ing low, flat warts, that of venter areo- late; dorsolateral folds indistinct; (2) tympanum prominent, its length 4-2 eye length; (3) snout subacuminate in dorsal view, round in lateral profile; canthus rostralis sharp; (4) upper eyelid nar- rower than IOD, lacking pungent tuber- cles; cranial crests present; (5) vomerine odontophores oblique in small individ- uals, triangular in outline in large fe- males; (6) males lacking vocal slits and vocal sac; males bearing non-spinous nuptial pad; (7) first finger shorter than second; fingers bearing broad discs on narrow, truncate pads; (8) fingers bear- ing lateral keels; (9) ulnar tubercles lacking; (10) heel, outer edge of tarsus lack tubercles; tubercle on inner edge of tarsus; (11) two metatarsal tubercles, inner oval, 3-4 times size of round outer; supernumerary plantar tubercles lack- ing, or few and indistinct; (12) toes bearing narrow lateral fringes, no web; toe pads small, as large as those of fin- gers; (13) brown above with darker blotches; labial stripe white to cream; venter white to gray, spotted or reticu- lated with brown or not; posterior sur- faces of thighs dark brown with cream spots; (14) adults moderate-sized, males 24.5-28.7 (x = 30.0 + 1.3) mm, females 37.1-48.8 mm SVL. Eleutherodactylus buckleyi differs from E. curtipes in having exposed tym- pana, subacuminate rather than rounded snout, less distinct dorsolateral folds, LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 5 Fic. 3. (A) Eleutherodactylus buckleyi, KU 177225, 43.1 mm SVL; (B) E. buckleyi, KU 167945, 45.1 mm SVL; (C) E. curtipes, KU 130596, 35.2 mm SVL; (D) E. curtipes, Laguna de Papallacta, not preserved; (E) E. curtipes, KU 130597, 35.0 mm SVL; (F) E. curtipes, AMNH 104719, 32 mm SVL. larger digital pads, and in greater size where the two are sympatric. The ven- ter of E. buckleyi bears dark brown or black spots or reticulation whereas in sympatric populations of E. curtipes the venter is cream without dark markings. Eleutherodactylus buckleyi also resem- bles E. devillei (Boulenger) and E. ver- tebralis (Boulenger), species found in high cloud forests on the Amazonian and Pacific Andean versants of northern Ecuador. Both differ from E. buckleyi in having distinct dorsolateral folds, longer hind legs, and in coloration. Description—Head narrower than body, wider than long; snout subacumi- nate in dorsal view, rounded in lateral profile; snout moderately short; canthus rostralis moderately sharp, _ straight; loreal region weakly concave, sloping to lips; lips not flared; interorbital space broader than upper eyelid; cranial crests heavy; upper eyelid lacking tubercles; supratympanic fold obscure, concealing 6 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY upper edge of tympanum; tympanum moderately to quite distinct, round to slightly higher than long, separated from eye by 1% times tympanic length; cho- anae small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores median and posterior to choanae, generally oblique in males and juvenile females, becoming triangular in outline in adult females; odontophores bearing row of 3-5 teeth along posterior edge, separated by distance equal to choanal width; males lack vocal slits and vocal sac. Skin of dorsum bearing low, flat warts (most evident on posterior back and flanks); dorsolateral folds not prom- inent; anal opening not extended in sheath; venter coarsely areolate; dis- coidal folds prominent; ulnar tubercles not evident; palmar tubercle bifid, larger than oval thenar tubercle; supernumer- ary palmar tubercles low, flat; subartic- ular tubercles round, nonconical; fingers lacking distinct fringes but bearing keel- like lateral ridges; all digits bearing discs (broader than long) on narrow pads; disc on thumb not distinct; first finger shorter than second. No tubercles on knee, heel, or outer edge of tarsus; indistinct tubercle on inner edge of tarsus; inner metatarsal tubercle oval, its length twice its width, non-compressed, 3-4 times size of round, non-conical outer; few or no_ super- numerary plantar tubercles; subarticular tubercles round to longer than wide; toes bearing narrow lateral fringes; all toes bearing discs (broader than long) on narrow pads; hind limbs short, heel of adpressed leg reaches eye. Brown above with indefinite darker blotching; lip stripe white to cream; venter white to pale gray, spotted or reticulated with brown or not; throat usually not spotted or reticulated in males; posterior flank, groin, anterior and posterior surfaces of thigh, con- cealed shank dark brown with cream spots. The testes are white in all males except those from the paramos de las Hermosas and Puracé in Deptos. Valle and Cauca in Colombia (those frogs have black testes). In life E. buckleyi is gray-tan to yel- low-brown through darker browns to reddish brown or black. The dorsum is flecked with black. The toe tops are black. The pale areas on the flanks, groin, and hidden limb surfaces are creamy yellow (rarely off-white). The venter is dirty cream with brown spots or marbling (to nearly uniform black). The iris is dark chocolate-brown with a reddish cast (or copper-bronze with black reticulation and a brown horizon- tal streak). Variation.—Males are smaller than females and have longer hind limbs. Specimens from the paramos de Puracé (Departamento Cauca) have larger tym- pana than specimens from the Nudo de Pasto and Ecuador and also have broader heads (Tables 1 and 2). The Cauca specimens are also distinctive in having darker venters than do more southern examples (Table 3). Remarks.—Only 3 of the syntypes of buckleyi (BM 78.1.25.40-41, 78.1.25.43) are E. buckleyi. BM 78.1.25.42 is an adult female (32.3 mm SVL) E. uni- strigatus exhibiting the striped pheno- type of that species (see account of E. unistrigatus ). The other syntype (BM 78.1.25.44) is a juvenile Gastrotheca riobambae (18.5 mm SVL). BM 78.1.- 25.40 is here designated the lectoholo- type. Peracca (1904) reported this spe- cies from Quito. His specimens are al- most certainly E. unistrigatus. Peracca cited Boulenger’s (1882; plate 14, fig. 5) figure of buckleyi (based on BM 78.1.- TaBLE 1. Variation in body size in male Eleutherodactylus buckleyi. Locality Range SE2SE a aN P. Puracé DAI5eSSu) | oodles La Cocha OF ASB ds) 302) 3 Santa Barbara Daeei(dl) — eX0)fofeA ts 8 13 km SE Tulean 25.6-34.2 29.0+1.9 11 P. Angél 93.8-34.4 30.4+2.8 8 Intag D(seeyo — Sil Seely! 8 LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS a TABLE 2. Variation in proportions in Eleutherodactylus buckleyi. (First line, range; second line, mean + 2 standard errors ) Shank/ HW/ Upper eyelid/ Tympanum/ Locality (N) SVL SiVie IOD Eye Puracé } 22 43.3-48.6 38.7-42.4 58.9-83.0 38.1-55.6 46.1+0.6 40.3+0.4 72.5+3.4 48.2+1.7 co) 4 43.7-46.4 38.7-41.4 63.7-90.3 40.6-49.5 44.8 78.1 46.8 P. Angél 6 7 41.0-46.0 36.8-40.1 69.6-100.0 32.9-40.9 43.5+1.0 38.8+1.0 78.6+7.4 35.8+1.8 ©) 4 35.8-41.7 35.5-39.1 80.0-100.0 31.1-39.0 39.0 37.0 88.5 ys) Intag é tf 38.5-46.8 35.1-39.2 60.5-90.1 30.0-42.1 43.3+2.4 37.6+0.8 71.6+6.6 Sowa ow, Q 5 42.2-46.2 37.2-38.9 60.5-85.1 30.0-39.3 44.6 37.9 68.4 34.8 TaBLe 3. Belly pigmentation (see Fig. 6) in Eleutherodactylus buckleyi Locality N Range *£+2SE P. Puracé 28 1-6 4.75+0.83 P. Angél 9 1-6 3.56+0.95 Intag 24 0-7 3.02+0.93 25.42) as identical with a pattern exhib- ited in his samples. The specimens from Departamentos Cauca and Valle (Colombia) differ from specimens from more southern localities in having darker venters, black pigment on the testes, and larger tympana. Nev- ertheless, I consider them conspecific with material from the Nudo de Pasto. Some frogs from the eastern face of the Llanganati mountains, although su- perficially resembling E. buckleyi in size, skin texture, coloration, digital pad size, and in having cranial crests, appear to represent a new species, description of which is deferred pending acquisition of additional material. Involved are the following specimens: CAS-SU_ 13190, 17429-37, “El Golpe,” 3000 m; CAS-SU 17439-41, Paracayacu, 2700 m; CAS-SU 17443, 17446, near Rio Jorge, 3000 m; CAS-SU_ 10390, 10392-99, at Rio San Jose, 10000 feet: all Ecuador, Prov. Napo, east face of Llanganati Mts. The sample consists mostly of juveniles al- though CAS-SU 17437 is an adult female (42.2 mm SLV). The undescribed form, unlike E. buckleyi, has the tympana con- cealed. The digital pads in the Llanga- nati form are larger than those seen in the allied E. cryophilius and E. curtipes. Distribution.—Cordillera Central of Colombia (ca. 3°30’ N) south to the Nudo de Pasto; Cordillera Occidental in Ecuador (Cordillera de Intac); Cordil- lera Oriental in Ecuador south to Nevado Cayambe (Fig. 4). Known at elevations between 2400 and 3700 m, primarily above 3200 m. Eleutherodactylus buckleyi is found in cloud forests as well as in paramo habitats. Eleutherodactylus curtipes (Boulenger) Fig. 3C-F Hylodes curtipes Boulenger, 1882: 218 (syn- types, BM 78.1.25.29-36 plus 3 others ex- changed to Werner C. A. Bokermann, Marquis Doria, and the National Museum of Natural History, collected at Intac, Provincia Imbabura, Ecuador, by Mr. Buck- ley). BM 78.1.25.29 is here designated the lectoholotype. Hylodes whymperi Boulenger, 1882:218 (syn- types, BM 82.7.13.3 [registered as BM 1947.2.17.24], obtained at Cotocachi, Pro- vincia Imbabura, Ecuador, 3963 m, by Ed- ward Whymper, and BM 82.7.13.4-5 [reg- istered as BM 1947.2.17.22-23], collected at Tortorillas, Provincia Chimborazo, Ecua- dor, 4024 m, by Edward Whymper). BM 82.7.13.4/ RR 1947.2.17.22 is here desig- nated the lectoholotype. New synonym. Hyla chimboe Fowler, 1913:157 (holotype, ANSP 18137, obtained from mountains 8 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 4. Distribution of Eleutherodactylus buckleyi. Stippled areas lie at or above 3000 m. above Chimbo (= San Jose del Chimbo) (probably on the Bolivar-Chimborazo fron- tier), Ecuador, 10000 feet, by Mr. Rhoads). New synonym. Diagnosis.—(1) skin of dorsum bear- ing low, flat warts and dorsolateral folds (some paravertebral folds as well), that of venter coarsely areolate; (2) tym- panum concealed beneath skin; (3) snout round to obtuse in dorsal view, round in lateral profile; E-N < eye length; canthus rostralis obtuse; (4) up- per eyelid narrower than IOD, lacking pungent tubercles; cranial crests present; (5) vomerine odontophores oblique in small individuals, triangular in outline in large females; (6) males lack vocal slits and vocal sacs; males have non- spinous nuptial pads on swollen thumbs; (7) first finger shorter then second; fin- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 9 gers bearing discs (about as wide as long) on scarcely expanded pads; (8) fingers bearing lateral fringes; (9) ulnar tubercles normally indistinct; (10) no enlarged tubercles on heel or tarsus; in- ner edge of tarsus bearing thickened tubercle; (11) two metatarsal tubercles, inner oval, 1% to 2 times size of outer; numerous supernumerary plantar tuber- cles; (12) toes bearing lateral fringes, no web; toe tips as narrow as finger tips; (13) pattern and coloration variable; pale stripe on lip; dorsum usually lack- ing distinct markings, limbs not barred; venter cream to heavily marbled with brown (Fig. 5); (14) adults moderate- sized, males 16.0-32.5 mm SVL, females 25.9-42.9 mm SVL (see Table 4). Eleutherodactylus curtipes differs from E. buckleyi in having the tympana concealed, a shorter, more rounded snout, more distinct dorsolateral folds, and in lacking distinct digital pads. Description—Head narrower than body, head wider than long; head width 29.9-41.8 (x = 37.0, N = 485) percent SVL; snout rounded to obtuse in dorsal view, rounded to sloping in lateral pro- file, slightly overhanging lower jaw, short, E-N less than eye length; canthus rostralis obtuse (rarely moderately sharp), slightly concave; loreal region concave, sloping abruptly to lips; lips not flared except in very large females; nostrils weakly protuberant, directed dorsolater- ally; interorbital distance greater than upper eyelid width; upper eyelid width 61.02121.4(x% = 89, N= 291) percent IOD in males, 60.5-108.6 (x = 77.4, N — 187) percent in females; interorbital distance narrower in males than in fe- males; edges of frontoparietals forming crests, crests more distinct in larger in- dividuals; frontoparietals complete, no fontanelle; tympanum concealed beneath skin; supratympanic fold prominent, obese; tongue large, thick, not notched posteriorly, posterior one-fifth to one- third free; choanae round, completely visible on roof of mouth when viewed TABLE 4. Variation in body size in adult Eleutherodactylus curtipes. Data given are: Range in mm (N) « + 2SE. Localities are numbered as in Fig. 8. Locality Males Females 1. Volcan Chiles 26.9-28.9(2) 2. 13 km SE Tulcan 96.9-31.5(3) 3. 14 km SW Tulcan 16.6-26.3(10)22.0+1.8 27 .6-33.2(13)30.2+0.9 4. 20 km SW Tulcan 18.8-22.9(9)21.3+0.9 27.4-32.0(16)29.7+0.8 5. Intag 29.6-32.8(2) 6. Nudo de Mojanda DIED eT (18) 23.9057 29.5-30.6(2) 7. Lago de Mojanda 19.2-27.5(35)23.7+0.7 25.7-36.7(20)31.4£1.0 8. V. Pichincha 21.6-30.6(12)26.3+1.6 35.3-42.9(8)40.4£1.5 9. Guamani 3800 m DIRQ=D 54103) 34.6-37.2(2) 10. Guamani 3600 m 22,.2-29.2(20)24.8+0.8 31.2-36.5(4)34.1 11. Lago Papallacta 23.7-28.4(10)26.1+0.8 33.8-35.4(4)34.5 12. vic. Papallacta 17.8-28.8( 49 )25.2+0.6 29.9-37.1(17)33.7+1.2 14. P. de Milin 21.8-29.6(4)24.8 30.2-40.5(10)35.9+2.2 15. P. de Apaqua 21.5-28.5(13)24.7+1.2 32.6-41.4(7)36.8+2.3 16. Guilo 3500 m 17.3-27.3(29)22.9+1.0 30.8-33.4(6)32.1+0.8 17. 12 km SW Sta. Rosa 16.7-27.0(15)23.2+1.4 32.9-37.0( 2) 18. 18-20 km SW Sta. Rosa 16.3-26.2(16)22.5+1.7 19. Arenal 16.0-26.8(11)22.5+2.0 28.2-34.4(6)31.0+2.0 20. 7-10 km W Banos 20.6-32.5( 26 )26.6+1.0 32.4-41.8(15)37.4+£1.4 21. 10 km W Cotald 26.6-31.8(4)28.8 32.8-40.2(3)36.5 22. 10 km SW Mocha 21.2-27.8(8)24.4+1.8 31.6-34.7(9)33.5+0.8 23. Urbina 17.4-28.9( 22 )25.1+1.2 27.2-39.0(38 )34.6+0.7 24. W San Juan 22.6-25.6(10)24.3+0.8 30.2-35.5(12)32.3-0.9 25. E Quaranda 17.0-25.2(15)23.2+1.0 30.0-39.5(12)34.0+1.6 26. SW Cajabamba DNeF-25.9'01) )23:622057 32.5-37.7(5)34.5 27. SE Pungala 19.1-21.2(2) 35.1-40.1(2) 10 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY from directly above; vomerine dentiger- ous processes present, median and pos- terior to choanae, obliquely oriented, each process two to three times size of a choana, triangular in outline in larger individuals, vomerine teeth in a trans- verse series across posterior edge of process, 4-7 teeth/process; dentigerous processes separated by distance equal to one-half width of a choana in large spec- imens, separation equal to two or three times width of choana in small speci- mens; males lack vocal slits and sac. Skin of dorsum bearing a pair of ill- defined dorsolateral folds (more promi- nent in living than preserved examples); some specimens have faint paravertebral folds as well, these are usually striped individuals (see below); center of back pustulate, bearing low, flat warts; skin of head weakly pustulate except on up- per eyelids which have low, flat warts; skin of flanks with large, low, flat warts interspaced among small warts; skin of lower flanks coarsely areolate, as is skin of venter, including throat; discoidal folds prominent to obscure, ending an- terior to groin; skin of face and con- cealed surfaces of limbs smooth, that of exposed surfaces of limbs bearing low, flat, warts; skin on knee, heel, and elbow bearing numerous small warts; ulnar surface bearing row of tubercles, vary from indistinct to prominent, coalesce into a ridge distally extending onto edge of palm and outer edge of fourth finger; all fingers bearing lateral fringes, fingers relatively slender in females, somewhat thicker in males; first finger shorter than second; subarticular tubercles round, non-conical, simple; thenar surface with few supernumerary tubercles, outer pal- mar tubercle largely fused with median (= bifid palmar tubercle), inner palmar tubercle (= thenar) largest; all fingers bear apical pads with circumferential grooves; pads of third and fourth fingers wider than long, truncate, not expanded; pads of inner two fingers more rounded and smaller than those of outer fingers. Tarsus bearing row of poorly defined tubercles along outer edge which coa- lesce distally and continue as a fold along sole and outer edge of fifth toe; all toes fringed, fringes coalesce as basal web; inner edge of tarsus bearing thick inner tarsal tubercle lying just proximal to inner metatarsal tubercle; two meta- tarsal tubercles, inner larger, oval, not compressed, outer round, low; outer metatarsal tubercle is one-half to two- thirds size of inner in large specimens, in small specimens the outer is as much as four-fifths size of inner; subarticular tubercles of toes smaller than those of fingers, round, non-conical, simple; no supernumerary tubercles on toes; plantar supernumerary tubercles numerous, low, flattened; shank 34.0-45.8 (x = 39.2, N = 305) percent SVL in males, 31.9-47.0 (x = 39.0, N = 206) in females. Males have large testes which are usually darkly pigmented with brown or black. The pigmentation was not found in 5.8 percent of the males examined, 17.6 percent had white testes with a black or brown network of pigment, and 76.6 percent had more or less uniform black or brown pigmentation on the testes. Some females, especially young females, have brown peppering on the peritoneum covering the ovaries. The coloration of E. curtipes is highly variable. The ground color is usually dark gray or brown but specimens with yellow or gold ground color are rela- tively abundant in central Ecuador and specimens with reddish ground color occur throughout the range of the spe- cies. Green is rarely seen in the color pattern. The majority of specimens have indefinite darker mottling on the dorsum but some specimens have _ distinct blotches, usually brown, on the back. The limbs are very rarely barred. The face is not barred but a distinctive cream or white stripe is found at the edge of the upper lip. The axilla, groin, and concealed surfaces of the thigh and shank are usually unpigmented or faintly pigmented with brown. Occasional ex- amples, mostly from central Ecuador, have pale red washes on the concealed surfaces. The dorsal pattern in some LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 11 specimens from most populations is striped—a combination of dark and cream stripes between the dorsolateral folds; the pattern is not polymorphic in E. curtipes but I will refer to the “striped pattern” and “normal pattern” in the dis- cussion of interpopulation variation. Variation.—As mentioned above, E. curtipes often exhibits a pattern of dor- sal stripes. Striped individuals occur in every sample examined except one from the paramo de Apaqua (N = 24) and another from Urbina (N = 71). In each case, adjacent samples exhibit low fre- quencies of striped frogs (Guilo, 8 km W Apaqua, 13%, N = 46; 10 km SW Mocha, 12%, N = 30). Elsewhere (Table 5), striped frogs occur at frequencies between 3.9% and 69.5% but with no sug- | gestion of a geographic pattern (Fig. 5, 6). In addition to the two cases of distinctly different frequencies in adjacent popu- lations cited above, the two sites on the Nudo de Mojanda differ appreciably in stripe frequencies (3.9 vs 33.8%) and the two sites on the southern flank of Ne- vado Chimborazo are markedly different (34.0 vs 69.5%) in spite of less than 10 km separations. Eleutherodactylus curtipes also var- ies considerably in the extent of dark pigment on the venter (Table 5). Vari- ation is continuous from completely cream or white venters to entirely black venters. To assess populational variation in this attribute, I coded a series of in- dividuals for increasing degree of pig- mentation (Fig. 6) and scored specimens to the best fit with the graded series. Ventral pigmentation varies geograph- ically in that the most northern samples consist of frogs with little or no dark pigment on the venter, whereas southern samples are made up of frogs with more dense pigmentation of the venter (Fig. 5). Adjacent populations do not exhibit pronounced differences (in contrast to the case in dorsal striping). The popu- lations with the darkest venters are found on Volcan Pichincha and SW of the Riobamba hoya. Eleutherodactylus curtipes exhibits TABLE 5. Pattern variation in Eleutherodactylus curtipes (see Fig. 6 for belly pattern code). Lo- calities arranged from N to S (see Fig. 8). Belly Pattern Locality % striped Range £+2SE N 1. Volcan Chiles 0.0 0-1 0.50 2, 3. 14 km SW Tulcan 56.5 0-4 0.52+0.43 23 4. 20 km SW Tulcan 33.3 0-3 0.22+0.24 27 5. Intag 0.0 0-1 0.50 2 6. Nudo Mojanda 3.9 0-6 2.31057) 26 7. Lago Mojanda 33.8 0-7 2.54+0.49 Thi 8. V. Pichincha O34 1-8 5.18+0.69 39 9-10. Paso Guamani 47.7 0-8 2.70+0.67 43 11. Lago Papallacta 57.9 0-8 3,241.94 19 N slope V. Cotopaxi 55.5 1-8 4.06+1.47 9 13. Nudo Tiopullo 24.4 0-7 2.11+0.48 78 14. Paramo de Milin 30.0 0-8 2.15+0.83 34 15. Paramo de Apagua 0.0 0-6 2.79+0.78 24 16. Guilo 3500 m 13.0 0-8 3.72+0.56 46 17. 12 km SW Sta. Rosa 40.0 0-7 2.84+0.9 25 18. 15-18 km SW Sta. Rosa 38.5 0-6 2.45+0.8 oF 19. Arenal 41.7 1-7 3.96+0.8 24 21. 10 km W Cotalé 14.3 0-7 2.54+1.9 1 22. 10 km S Mocha 12.0 1-6 2.42+0.65 30 23. Urbina 0.0 0-6 1.80+0.38 7 24. W. San Juan 69.5 0-9 3.48+0.7 59 25. E Guaranda 34.0 0-7 3.86+0.8 29 26. SW Cajabamba 16.7 2-9 5.90+0.8 23 27. SE Pungala 25.0 1-6 3.25 4 12 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 5. Geographic variation of Eleutherodactylus curtipes in (A) frequency of dorsal striping and (B) ventral pigmentation. 3000 meter contour is outlined for geographic reference. considerable variation in body size (Table 4). The smallest frogs are found in the most northern samples although small frogs also occur on the NW slopes of Nevado Chimborazo. The largest frogs are found on ridges west of the junction of the Rio Chambo and Rio Patate in Central Ecuador and on Volcan Pi- chincha. The broad overlap in sizes of adult males and adult females (24.5% of total size range), indicated in the diag- nosis, is a product of marked interpopu- lational variation in body sizes. In only two populations (Laguna de Mojanda and W of Banos) is there actual overlap (8.0% and 0.5% respectively ). Eleutherodactylus curtipes is a short- legged frog. Except for the northern- most populations (in which legs are quite short), leg length decreases from north to south (Table 6). The cline from the Desierto de Palmira north to the Nudo de Mojanda breaks sharply in E. curtipes but the sympatric E. buck- leyi has long legs as might have been predicted for the northern curtipes pop- ulations. Head width does not exhibit so marked a cline but nonetheless de- creases slightly from north to south (Table 6). Eyelid width relative to IOD exhibits almost no variation except that males have larger eyes than females. The high variance in males of the 10 km SW Mocha population is inexplicable (Table 6). Natural history—E. curtipes is an abundant, lapidicolous frog living in the prairie-like paramos (Fig. 7) of the northern two-thirds of Andean Ecuador. Over this geographic area, the paramo LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 13 Fic. 6. Standards of ventral pigmentation in Eleutherodactylus curtipes. (A) KU 119604; (B) KU 119632; (C) KU 119644; (D) KU 119608; (E) KU 119656; (F) KU 119645; (G) KU 119634; (H) KU 119592. A-H are pigmentation standards 1-8. Standards 0 (completely white venters) and 9 (completely black venters) are not illustrated. habitat occurs between 3200 and 4700 m; above 4700 m the average annual temperature is 5°C or below and the soil frequently frozen or snow-covered (Acosta-Solis, 1968). The lowest eleva- tion at which E. curtipes has been found is 2750 meters (1 km W Latacunga, Provincia Cotopaxi, Ecuador). The re- maining localities range from 3200 m to 4400 m. The species no doubt occurs above 4400 m but to date I have been unsuccessful in attempts to scale Nevado Cayambe or N. Chimborazo to such altitudes to search for it. At altitudes above 3200 m, E. curtipes is not found in the few remaining high altitude for- ests although other species occur there. At elevations below 3200 m in Ecuador, the predominant “forests” are Eucalyp- tus groves. In these habitats, the only eleutherodactyline frog encountered is E. unistrigatus. The grasslands extend below 3000 m in many areas in Ecuador but the microenvironments are percep- tibly more xeric than at higher eleva- tions and E. curtipes is replaced by E. unistrigatus (Figs. 8-9). Eleutherodac- tylus unistrigatus is usually parapatri- cally distributed relative to E. curtipes but I have collected them syntopically at five localities, all but one above 3200 m (“Intag”’, Imbabura Proy.; slopes of Volcan Pichincha, Pichincha Prov.; Nudo de Tiopullo, Cotopaxi Prov.; and 10 km W Cotalé, Tungurahua Prog.) in Ecua- dor. In July 1977, two E. curtipes were found beneath rocks beside the Rio Latacunga (1 km W Latunga, 2750 m) where E. unistrigatus is extremely com- mon. The microhabitat was very wet; MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 14 (3) TOL (3) 108 (8) FFOFESS (11 )6Z0+088 (IT) 8@O*LSL (FI )6F0+008 (03) 1TFOFFI8 (OT )9E0098 (81 )9ZO+9SL (ZL )SPOFETS (OL )9S006L (8)ELE+9L8 (SEE (7 )€08 (€1)0SO*6SL (OL)EZO+0E8 (FI )0G0+Z08 (F)O8L (ST )SZ0+98L (9)SZ0*ZS9 (LZ )EZOFELL (L)6F0+699 (€T)9Z0*008 (OL )FFOFSSS (F)1L8 (LI )9€0*S08 (6F ) 81008 (9 )SFOFESL (11)980+828 (11) LSOO0P8 (03) 1006 (3)8SL (€)PS8 (L)SZO+FF9OL (11 )Se00E8 (0% )SZ0O+60L (SE) FZOF9SL (8)880+F9OL (81)TS0+8T8 (3)998 (3) #S6 So[PULO Sol[P IN COI/PH949 (3)36E (3)€6E (3)O8E (3)OIF vyesung AS “LZ (8)L00+89¢ (IT )O10+998 (8)910+9LE (IL)Z10O=LLE vquieqeley) AAS “9% (ZI )OIOFZLE (ST )900+€8E (ZI )FIOFZSE (ST )O1O+06E vpuvieny WY “GZ (03) FOOFSSE (OL )600+LSE (0) 800+88¢ (OI )900+E€8E uvn{ Urg AA “FZ (81) L00+08E (31 )600+F6E (SZ) S00+FLE (ZI )600+€8E vUIqI, “EZ (OT )900+ELE (8)FIO=08E (OL) ZIO+ L0F (8)FIO+96E vYDOW MS “GS (¢)69¢ (F)99€ (€) FPP (F)SGP Q[RIOD M “TZ (€T)LOOFSLE (11 )800+996 (9 )91O*S6E (IL) ZIOFESE [vuc1y “61 (91) LOOFOLE (91) Z10O*Z9E Bsoy "kIS AAS ‘ST (F)Z9E (SI )900+F9E (FP) PLE (ST) L00+89¢ vsoy ‘RIS AAS ‘LI (9)900+69¢ (9%)SOO*OLE (9) 11O+S8IF (13) LOO*SIF o[mMy ‘9T (L)800+998 (€T)SOOFLOE (L)800+88S (€1 )600+ OF vensedy ‘d ‘cy (OL )€1O+F9E (F) LOE (TT )STO*6LE (+) 90F uy ‘dP (LT )900+998 (6%) FOOFEIE (LT) 110+€6E (6F ) LOO#06E oy[ndory, “ET (9) TIOF9E (OL) FIOFFSE (9) FLIOF FSP (O01 )600+66E vjoryedeg ose] “TT (IT )OLO+6LE (0%) FOOFGLE (IL) FIO+OLF (0%) 900+ 66E TuvURNny) “OT (3)EePrE (€)P9E (3)3LE (€)€OF TuvUeny) "6 (8) L00+SS¢ (ZI) 800+ GSE (8)ZIO#FLIF (ZL) FIO+S80F vyouryorg urofoA 8 (33) SOO#LSE (SE )SOOFESE (0% )S800+00F (SE )SOO+F68E vpurloyy ose'y "py (8)600+68E (8T)LOOFOLE (8)O01IOFLGF (ST )OLO+OIF vpurloyyY OpnN “9 (3)6SE (3) 1gS¢ Bey “GC (91 )O1O+Ss¢E (OL )€1O*6LE uvgznL MAS ‘FE (3) SSE (3)F9E soy UvofoA ‘T so[vulo J solv So[VUld Solve A}T[ROO'T TAS/MH TAS/PIGEL ‘9 BIY Ul se poloquinu puv § 0} N Woy posure sanyeooyT (NV) ASG UvoUT o1v UDAIS son[eA ‘sadyina snjhjopposayinayy JO (OOOT x) suoqodoid ut suoyeura orydeiso9s ‘g ATAV], LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 15 Fic. 7. Eleutherodactyline frog habitats. (A) Paramo del Angél, ca. 3800 m; (B) paramo 13 km SE Tulcdn, 3200 m; (C) Paso de Guamani, ca. 4000 m; (D) Paso de Guamani, ca. 3600 m; (E) Paéramo de Apagua, 3860 m; (F) padramo 12 km SW Cajabamba, 3800 m. when a rock was overturned, water flowed from the soil into the cavity. Additionally, some museum records sug- gest sympatry but without precise col- lecting data, altitudinal parapatry or ecological parapatry (such as E. uni- strigatus in a Eucalyptus grove and 50 m away E. curtipes in grassland) give an illusion of sympatry. Dra. E. del Pino found both species in Machachi, Prov. Pichincha. My analysis suggests that E. curtipes prefers higher, more mesic habitats (= grasslands, paramo) whereas E. uni- strigatus prefers (or is forced to occupy) lower, more xeric habitats (= Eucalyp- tus groves, grassland below 3200 m). In dry paramo, such as paramo del Angél or Arenal (west falda of Nevado Chim- borazo), E. curtipes is very locally dis- tributed and found only along stream courses. In the paramos del Angél, most of the habitat away from the streams is occupied by E. thymelensis, a species smaller than E. curtipes but about the size of E. unistrigatus (Lynch, 1972b). The paramo Arenal is very much drier (Fig. 10) and all frogs occurring there 16 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 8. Distribution of Eleutherodactylus curtipes (@). Numbers in circles identify localities listed in tables 4-6 and text. Six Andean towns (™) are named. Hexagons represent localities at which E. curtipes is sympatric with E. unistrigatus (Fig 9). Area above 3000 m is stippled. Major Nevados are indicated by A (named on Fig. 9, facing page). LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 17 CcAYAMBE rN 2 Ne Zen) ‘,~COTOPAXI: « Fic. 9. Distribution of Eleutherodactylus unistrigatus (@). Some rivers and Nevados (A) are identified as are certain towns (™) for geographic reference. Area above 3000 m is stippled. 18 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY are restricted to stream courses. The only Eleutherodactylus sympatric with E. curtipes at Arenal is E. orcesi which is about the size of E. thymelensis and E. unistrigatus (Lynch, 1972b). In all my work in the Ecuadorian paramos I have never been certain that I heard E. curtipes call. Colostethus and Gastrotheca frequently are heard calling and other high altitude Eleutherodac- tylus have been traced to calling males. I have frequently heard short trilling Fic. 10. Habitats of Eleutherodactylus curtipes. Mesic areas on east slope of N. Chimborazo, 3700 m—(A) cushion plants, (B) aspect. Xeric areas, west slope of N. Chimborazo, Arenal, 4150 m—(C-D) dry areas, (E) slightly more moist area. (F) Desierto de Palmira, N of Palmira. noises in the paramos but cannot say that the calls were not produced by insects. I have found egg masses on only four occasions. Eggs are adherent. A mass of 19 eggs (5.2-5.4 mm diameters) was found February 23, 1968, beneath a rock at about 3600 m on the south slope of Volcan Chiles, Prov. Carchi, Ecuador (KU 118107). No frogs were beneath the rock. The eggs are early in development. The egg mass is almost certainly that of LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 19 E. curtipes although E. thymelensis was more abundant at the locality. On 15 July 1970, a second egg mass (KU 130466) consisting of 79 eggs (4.3-4.8 mm diameter) was dug out of an earthen bank where it was under a well-buried rock (Fig. 11). This egg mass was found at 3700 m near the frontier between Provincias Bolivar and Chimborazo (Guaranda-Riobamba road). Although E. orcesi probably occurs at the same local- ity, the number of eggs and the abun- dance of E. curtipes with ripe eggs sug- gest that the mass is of E. curtipes. These eggs are also in early stages of development. On 14 July 1977, while marking E. curtipes, I found four egg masses be- tween stones 4.7 km W Papallacta, Prov. , Napo, Ecuador, 3360 m. The clutches were within 2 m of one another. KU 180286 consists of 38 eggs (diameters ca. 3.2 mm) and KU 180287 of 17 eggs (diameters ca. 4.6 mm). The other two clutches were parasitized by an un- known dipteran. KU 180288 consists of 37 eggs plus four or five capsules fused together; egg diameters are about 5.0 mm. KU 180289 consisted of more than 60 eggs, mostly destroyed by maggots. On 27 July 1977, at 10 km SSW Mocha, frontier between Provincias Chimborazo and Tungurahua, 3450 m, I found a clutch of 30 eggs (diameters ca. 5.0 mm) and three jelly capsules (KU 180290) beneath a rock. A few centi- meters away was an adult female E. curtipes. The eggs were in a cavity beneath the rock. I have collected E. curtipes in Janu- ary through March and June through August and detect no evidence of sea- sonality as judged from the ripe ovarian eggs present in the vast majority of adult females from each month. Young E. curtipes (under 13 mm SVL) have been collected at most localities and during all months collections were made. Thus reproduction is apparently aseasonal. Eleutherodactylus curtipes is most readily collected along road cuts where there is an abundance of rocks (Fig. 11). However, careful collecting, frequently induced by a scarcity of rocks and the crisp paramo climate, has revealed that the frogs occur at the bases of bunch grasses (Festuca and Stipa) as well as in fissures in the soil (Fig. 11). When many rocks are present, E. curtipes may be very abundant. For example, on 7 August 1970 I caught 80 E. curtipes in 30 minutes. Lynch (1969, 1970) demonstrated that collecting habits can result in biased sex ratios in collections, so care was taken to collect in diverse sites within a locality. The combined localities yield the ratio of 263 males to 169 females among the adult E. curtipes measured. The higher number of males may reflect a real bias in the sex ratio or may reflect my age determination technique. Fe- males with small ovarian eggs and thin, straight oviducts were classed as im- mature. Their inclusion in the female count results approximately in a 50:50 sex ratio. Eleutherodactylus curtipes has been collected at many localities in the Andes of northern Ecuador. About 90 per cent of the extant specimens were collected in the past decade. At every locality where I found E. curtipes, the vegeta- tion was largely grassland (paramo) with few scrubs or trees. The most il- luminating case involves collections on the Nudo de Mojanda (Imbabura Prov.). Collecting in the vicinity of the Lagos near the top of the Nudo (3680 m) yielded numerous E. curtipes as well as Atelopus ignescens and a few Gastro- theca. The specimens were taken be- neath rocks and grass as well as from fissures in dirt banks. The vegetation consists of bunch grasses and scattered stands of a shrub (Polylepis) which reaches a height of 2.5-3.0 m. Tree-line in 1970 was about 3600 m; the predomi- nantly grass paramo abruptly gave way to a relatively dense forest festooned with epiphytes and llianas. The trees reached heights of between 6 and 10 m. No Eleutherodactylus were found in the upper forest but at 3400 m several spe- 20 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY BJisiinaigs NOSaHiwvo Fic. 11. Eleutherodactylus curtipes microhabitats: (A) bunch grass, paramo on frontier on Guaranda-Riobamba road, 3700 m; (B) rocky area, near Laguna de Papallacta, 3360 m; (C) closeup of part of B where four nests were found; (D) closeup of one nest; (E) egg mass, KU 180288; (F) egg mass, KU 130466, in situ, above Guaranda, 3700 m. cies (but not E. curtipes) were found. At the same altitude there was a large patch (several hectares) of grassland environment. Few rocks were present but in fissures in the soil I found many specimens of E. curtipes. The experience of intensive collecting in distinct but adjacent communities leads me to con- clude that E. curtipes is a paramo frog and cannot (or prefers not to) invade forested habitats. The widely intro- duced Eucalyptus forests occur in Ecua- dor to about 3200 m; E. curtipes invari- ably occurs above the Eucalyptus forests and not infrequently is found in grass- lands adjacent to them. The paramos of Ecuador are prob- ably much more widespread today than in the comparatively recent past. Before extensive deforestation, paramos evi- dently did not occur at elevations lower than 3600 m, if the forests on the slopes of the relatively inaccesible (until re- cently) Nevado Cayambe and Nudo de Mojanda are representative. If the pref- erence for grassland habitats by E. cur- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 21 tipes has not changed in response to the recent expansion of grassland habitats the distribution area of E. curtipes may have been significantly increased and become more continuous in the recent past through the activities of wood- cutters and carboneros. This increase of area and development of a continuous habitat would have the effect of swamp- ing the distinctions acquired by popula- tions previously isolated on mountain peaks above 3600 m. Often, samples from relatively near-by localities differ appreciably in proportions and _ color- ation (e.g., frequency of striped individ- uals at localities 24 and 25 or 6 and 7; mean values of belly pigmentation at localities 24 and 26); such disparities may well reflect recent isolation of pop- | ulations. Remarks.—The type-series of Hy- lodes curtipes consists of 11 specimens, of which 8 (BM 78.1.25.29-36) have been examined; 3 others were exchanged (Werner C.A. Bokermann, Marquis Doria, and USNM). The 8 are conspe- cific; BM 78.1.25.29, an adult female 32.8 mm SVL is here designated the lecto- holotype. The type-series of Hylodes whymperi includes 3 specimens; 2 (BM 82.7.13.4-5/ RR_ 1947.2.17.22-23) are adult males (each 25.2 mm SVL) from Tortorillas, Prov. Chimborazo, Ecuador, 4024 m. The third syntype is a juvenile female, 25.0 mm SVL, from Cotocachi, Imba- bura Prov., Ecuador, 3963 m (BM 82.7.12.3/ RR 1947.2.17.24). BM 82.7.- 13.4/ RR 1947.2.17.22 is here designated the lectoholotype. Boulenger (1882) dis- tinguished whymperi from curtipes be- cause the tympanum is partially visible in the latter, the digital pads of the lat- ter are smaller than those of the former, and the hind limb of the former is slightly longer than that of the latter. Additionally, Boulenger (1882: 218-19) recorded the presence of an extensive frontoparietal fontanelle in curtipes. The frontoparietal fontanelle is not ex- posed in the BMNH specimens but may be in one or more of the three specimens exchanged to other collections. The dif- ferences in distinctness of the tympa- num, digit pad size, and limb length (used by Boulenger to distinguish taxa) vary geographically in Ecuador and so long as one has small sample sizes ap- pear distinctive. Another geographically variable feature not clearly noted by Boulenger is color of the venter. He re- corded the venter of curtipes as immac- ulate and that of whymperi as immacu- late or reticulated with brown. The whymperi syntype having an immaculate venter is from Cotocachi near the Cor- dillera de Intag in Imbabura Province. The two with spotted venters are from Tortorillas on the slopes of Nevado Chimborazo, Chimborazo Prov., Ecua- dor. I consider the two names synony- mous and in spite of line priority select curtipes as the name for the species (acting as first reviser). Hyla chimboe was named by Fowler (1913) on the basis of ANSP 18137, an adult female 36.3 mm SVL. His mis- identification of the specimen as a hylid insulated the name from comparison with Andean Eleutherodactylus. The type is a short-legged, stocky frog of the E. unistrigatus group having narrow dig- ital pads, prominent cranial crests, par- tially concealed tympana, and a pale stripe on the upper lips. The venter is not reticulated with brown. The visibil- ity of the tympanum may be a result of preservatives. The tympanum lies be- neath the skin, now partially transparent in the temporal region, as in the type- series of E. curtipes. Hyla chimboe is here considered a synonym of E. cur- tipes. Distribution.—E. curtipes occurs at elevations between 2750 m and 4400 m but is primarily distributed above 3200 m. Specimens are known from three localities virtually on the Colombian border but no specimens are available from Colombia. In Ecuador, E. curtipes is found on the crests of the eastern and western cordilleras (and on the connect- ing nudos) south to the Desierto de Palmira (Fig. 8). Peracca’s (1904) re- 22 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ports of Hylodes curtipes from E] Troje and Tulcan (Prov. Carchi) and of H. whymperi from Papallacta (Prov. Napo) probably represent this species. His rec- ords of H. buckleyi from Quito (Prov. Pichincha) and El] Troje, Huaca (Prov. Carchi) probably represent E. unistri- gatus whereas those for Tambo (Prov. Cafiar) probably represent E. riveti. Peracca’s (1904) report of H. curtipes from Paredonas, 4042 m (Azuay Prov.) probably applies to E. riveti. Eleutherodactylus modipeplus new species Fig. 12A Holotype—KU 131283, an immature female taken 1 km S of the Urbina rail- way station, Provincia Chimborazo, Ecuador, 3650 m, on 29 July 1970 by John D. Lynch. Paratypes.—KU 131284-85, taken syn- topically with holotype. Diagnosis——(1) skin of dorsum sha- Fic. 12. Andean Eleutherodactylus. (A) E. modipeplus, KU 120021, 34.2 mm; (B) E. myersi, KU 168436, 18.9 mm; (C) E. orcesi, AMNH 104720, 32 mm SVL; (D) E. orcesii, KU 177814, 35.2 mm; (E) E. thymelensis, topotype, not preserved; (F) E. unistrigatus, KU 119750, 28.4 mm. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 23 greened with scattered warts, that of venter areolate; no dorsolateral folds; (2) tympanum prominent, round, its length % to % that of eye; (3) snout subacuminate in dorsal view, rounded in lateral profile; E-N slightly less than eye length; (4) interorbital space slightly greater than upper eyelid width; most individuals have flat interorbital region but low crests develop in large females; (5) vomerine teeth and odontophores present, odontophores slanted (teardrop- shaped), posterior and median to cho- anae; (6) males with vocal slits and subgular vocal sac; (7) first finger shorter than second; all digits bearing discs on dilated pads; dilation ratios I: 1.4, II: 1.7, III: 2.1, IV: 2.1; (8) fingers bearing lateral fringes; (9) ulnar tubercles poorly | defined; (10) inner tarsal surface bear- ing one or two tubercles, outer edge of tarsus with a row of ill-defined tubercles; no prominent heel tubercles; (11) two metatarsal tubercles, outer round, coni- cal, 4 to 4 size of elongate inner; numer- ous supernumerary plantar tubercles; (12) toes bearing lateral fringes, no basal webbing; all toes bearing discs on di- lated pads, pads as large as those of fingers; (13) dorsum gray to brown with brown chevrons, interorbital bar; limb bars narrow, oblique; venter cream to nearly black; concealed limb surfaces, groin, and lower flanks pigmentless in preservative (red in life); several indi- viduals have pale interorbital bar in front of dark bar and pale blotch edged with black in the center of the back; (14) adults small, males 24.6-28.5 mm, females 28.0-34.7 mm SVL. Eleutherodactylus | modipeplus was confused with FE. cajamarcensis by Lynch (1969:269) who noted that modi- peplus lacks the spotting of the venter characteristic of cajamarcensis. Eleu- therodactylus modipeplus likewise lacks white spots on the black field on the posterior surfaces of the thighs. Lynch (1969) noted some partially distinguish- ing differences between E. modipeplus and E. unistrigatus (viz., tubercles on eyelids, low, pustular ridges on the dor- sum, and supernumerary plantar tuber- cles of E. modipeplus). In life, E. modipeplus and E. uni- strigatus are easily separated because the former has red flash colors in the groin, anterior and posterior surfaces of the thighs, and concealed shank (cream to yellow in E. unistrigatus). Preserved examples of E. unistrigatus are subject to confusion with E. modipeplus. Ex- cept for the northermost populations of E. unistrigatus, the concealed surfaces of the thighs and groin are finely pep- pered with melanophores (lacking in E. modipeplus). The snout is more acumi- nate in E. modipeplus than in E. uni- strigatus. Description—Head narrower than body, slightly wider than long; head width 36.0-39.6 (x = 37.3) percent SVL; snout subacuminate in dorsal view, bluntly rounded in lateral profile; no keel on snout tip; canthus rostralis mod- erately sharp, concave; loreal region weakly concave, sloping abruptly to lip; lips not flared; nostrils directed dorso- laterally, not protuberant; eye length slightly greater than E-N distance, E-N 82.8-100.0 (x = 93.0) percent eye length; interorbital space flat in most individ- uals but low lateral crests occur in large females (KU 120021, 34.2 mm SVL, and 142134, 34.7 mm SVL); no frontoparietal fontanelle; upper eyelid width 71.1-83.1 (x = 78.0) percent IOD; tympanum prominent, round, its upper edge con- cealed by thick supratympanic fold; tympanum length 35.3-45.8 (x = 41.1) percent eye length, not sexually di- morphic; choanae small, round, situated well lateral on palate, completely visible when palate is viewed from directly above; vomerine odontophores between and posterior to choanae, each twice as large as a choana; odontophores not ele- vated, slanted (teardrop-shaped), bear- ing 2-6 teeth in a slanted row, separated by distance less than choanal width; tongue large, longer than wide, weakly notched posteriorly, posterior one-third to two-fifths not adherent to floor of 24 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY mouth; males with vocal slits and sub- gular vocal sac. Skin of dorsum shagreened with scattered low warts; enlarged warts most numerous on posterior portion of back; upper eyelid tuberculate; no dorsolateral folds; flank bearing large, flat warts; venter coarsely areolate; discoidal folds prominent. Series of indefinite ulnar tubercles present; one bifid palmar tubercle or two palmar tubercles (median larger); larger palmar tubercle larger than oval thenar tubercle; a few low supernumer- ary tubercles on thenar surface; sub- articular tubercles large, round, non- conical, simple; digits fringed (includ- ing outer edge of hand and fourth fin- ger); all digits bearing discs on dilated pads, pads broader than long on all digits but pad of thumb smallest; dila- tion ratios I: 1.2-1.6 (x = 1.4), I: 1.5- De eye de), Te 8-216) (4 = 2), IV: 1.7-2.4 (x = 2.1); tarsus bearing one to two small tubercles along inner edge; outer edge of tarsus bearing a row of indefinite tubercles (none prominent); no enlarged tubercles on heel; hind limbs short, heel of adpressed leg ex- tends to between posterior corner of eye and tympanum; shank 43.2-51.1 (x = 46.9) percent SVL; two metatarsal tu- bercles, outer rounded, conical, one-fifth to one-third size of elongate (length twice width), non-compressed inner metatarsal tubercle; numerous super- numerary plantar tubercles; subarticular tubercles of toes like those of fingers; toes bearing lateral fringes, no basal webbing; toes bearing discs on dilated pads, pads broader than long, smaller than those of outer fingers. In preservative, the ground color (all dorsal surfaces but limited to an inter- rupted line atop thigh) gray to gray- brown with brown chevrons and elon- gate spots edged with cream; some spots, especially on lower back, are spotted with black; interorbital band prominent, black or dark brown with an off-white to cream band anterior to the black band; some individuals have an off-white to cream blotch above sacrum; the spot is irregularly edged with black; the flanks are not spotted or barred; shank and forearm bear oblique bars, slightly nar- rower than pale interspaces; canthal and supratympanic stripes not prominent; lips barred; venter cream with white in- fusion in most individuals; some have faint gray reticulation on belly; groin, anterior and posterior thigh not pig- mented but the thigh may have some brown reticulation on the upper portion of the posterior surface. The individuals in the type-series have a dark-brown ground color and little pattern is evident beyond the interorbital bar and sacral spot. These individuals also have brown venters with some mottling on the throat. In life, E. modipeplus is brown to reddish-brown with creamy-tan inter- orbital bar and sacral spot (both edged with black). The groin, axilla, concealed thigh and concealed shank are reddish- orange to blood red. The throat of males is pale yellow. The venter is dirty cream with or without gray and brown mar- bling. The iris is gray marbled with brown and having a medium reddish- brown horizontal streak. Measurements of the holotype in mm.—SVL 25.7, shank 12.0, head width 9.5, upper eyelid width 2.4, IOD 3.1, tympanum length 1.2, eye length 2.8, E-N 2.7. The holotype is an immature female with straight (non-convoluted) oviducts and small cream ovarian eggs. Etymology.—Latin, modicus (mod- est) and peplum (robe or tunic), in reference to the less ornate flash-colors compared with E. cajamarcensis (no black and white marbling). Natural history—All specimens of E. modipeplus I found were taken by day beneath rocks, clumps of dirt, or bunches of paramo grass. Gravid fe- males have been found in June and August. Specimens captured in July 1977 (10 km SW Mocha, Prov. Tungu- rahua, 3700 m) and kept live in aquaria regularly called until late August when they died. The call is a sharp whistle and markedly unlike the call of E. uni- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 25 strigatus. James A. Peters secured call- ing males on bushes 2 km S Mocha, Prov. Tungurahua, 3000 m. The type-series was found beneath a large flat rock beside a stream. Three individuals were captured but two oth- ers jumped into the stream and escaped. Specimens collected in 1968 (10 km SW Mocha) were found beneath clumps of dirt in a dried-up vernal pond. Eleutherodactylus wunistrigatus has not been taken sympatrically with E. modipeplus but the two surely occur to- gether. The usual congeneric sym- patrant is E. curtipes. The breeding season of E. unistrigatus is from late December until February. E. modi- peplus appears to breed during June and July but too little data are available . to be conclusive. E. modipeplus, like E. curtipes, may breed all year long at higher (3200-3700 m) altitudes. Remarks.—In my redescription of E. cajamarcensis (Lynch, 1969), it was dis- tinguished from wunistrigatus in part by the “slightly smaller” digital pads of cajamarcensis (p. 269). That statement is in error. The dilation ratios of the hand of cajamarcensis are as follows: I: fee (eA) 6-210" 8); 92022) (2 =o) AV 2922.3 (es = 2.2). The values for unistrigatus are: I: ies (a= V2) 1s) PAE. 8) (4-1-5), ME tA. 0nd Xe IY), TV = VA-1 9n(e = 1.7). The dilation ratios of modipeplus are not appreciably different from those of cajamarcensis, but both have slight larger digital pads than does E. uni- strigatus. Distribution—Known only from the edges of the Ambato-Riobamba hoya (Fig. 13) in central Ecuador at eleva- tions between 2560 and 3700 m. Eleutherodactylus myersi (Goin and Cochran) Fig. 12B Trachyphrynus myersi Goin and Cochran, 1963: 502 (holotype, CAS 85177, an adult female obtained “20 miles east of Pasto,” Depto. Narifo, Colombia, on 1 March 1955 by E. S. Ross). Eleutherodactylus myersi: Lynch, 1968:295. Diagnosis.—(1) skin of dorsum pus- tulate, bearing paravertebral and sinuate dorsolateral ridges, that of venter areo- late; no anal sheath; (2) tympanum vis- ible, its length %-% eye length; (3) snout subacuminate in dorsal view, rounded in lateral profile, bearing pa- pilla at tip; canthus rostralis sharp; (4) upper eyelid bearing numerous low tu- bercles, its width 51.7-79.2 (x = 65.9, N = 18) percent IOD: no cranial crests; (5) vomerine odontophores absent, 0-2 vomerine teeth; (6) males with vocal slits, white nuptial pad on thumb, black lining of testes; (7) first finger shorter than second; fingers bearing discs and small pads; (8) feeble lateral keels on fingers; subarticular tubercles flat; (9) low ulnar tubercles present; (10) non- conical tubercles on outer edge of tarsus; short inner tarsal fold; (11) two meta- tarsal tubercles, inner oval, 3-4 times size of outer; numerous supernumerary plantar tubercles; (12) toes bear narrow lateral fringes; toes with narrow pads and discs; (13) brown above, black be- low; pale spots in groin and anterior and posterior surfaces of thighs (spots red in life); inner digits cream; (14) adults small, size geographically variable (see below), males 13.7-17.5 mm, females 17.5-23.2 mm SVL. Eleutherodactylus myersi is most similar to E. trepidotus but differs in lacking vomerine odontophores and in having outer tarsal tubercles and distinct ridges on the dorsum. Description—Goin and Cochran (1963); Cochran and Goin (1970). Variation —Although known from few specimens and only nine localities, geographic variation in size is apparent. Specimens from the Pasto region are larger than those from the Péramo de Puracé: Pasto male 17.8 mm SVL, fe- miales/ 199-23: (2.— 21-65-2903. .N 7) mm SVL; Puracé males 13.7-13.8 mm SVE (N= 2), females 17:5-20:9 (x = 18:8 2-512. Ne] 5) mm) SVE. Distribution—Known from paramos on the Nudo de Pasto and the southern end of the Cordillera Central in Co- 26 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Kilometers Fic. 13. Distributions of Eleutherodactylus modipeplus (@) and E. ocreatus (A) in Ecuador. lombia at elevations between 2900 and 3275 m. Eleutherodactylus ocreatus new species Eleutherodactylus myersi (part): Lynch, 1968, 295, 289-99. Holotype—KU 117573, an adult fe- male collected on the west slope of Vol- can Chiles, 10 km W_ Tufifio, Prov. Carchi, Ecuador, 3500-3800 m, on 23 February 1968 by John D. Lynch. Paratypes.—KU 117574-81, taken with the holotype. Diagnosis.—(1) skin of dorsum bear- ing low ridges, finely areolate, subconi- cal warts on flanks; upper eyelid bearing low tubercles; (2) tympanum concealed; (3) snout round in dorsal and lateral profiles, short; (4) interorbital space flat (no cranial crests), broader than upper eyelid; (5) vomers lacking odontophores; (6) males with short vocal slits, internal vocal sac; testes dark brown; (7) first finger shorter than second; fingers lack- ing pads; (8) fingers lacking lateral fringes; subarticular tubercles non-pun- gent; (9) ulnar tubercles not distinct; (10) no enlarged or distinctive tubercles on heel or tarsus; (11) two metatarsal tubercles, inner large, not compressed, two to three times size round, non-pun- gent outer; (12) toes lacking lateral fringes, webbing, pads; (13) brown to dark-brown with pale spots in axilla and groin (white with pink cast in life); venter dark brown, spotted with cream or not; (14) four males 12.4-17.2 (* = LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 27 14.4) mm SVL, two adult females 19.6- 20.2 mm SVL. Eleutherodactylus ocreatus is most similar to E. trepidotus and E. vidua but differs from both in lacking vomerine odontophores. Eleutherodactylus myersi, with which E. ocreatus was previously confused by me, is readily distinguished in having small, albeit perceptible, dig- ital pads (Fig. 14) and a larger, more obvious tympanum (Fig. 15). Description —Head as broad as body except in adult females (where narrower than body), wider than long; head width 34:6-37.9 (x = 36.6, N = 6) percent SVL; snout rounded in dorsal and lat- eral views; nostrils weakly protuberant, directed dorsolaterally; snout very short, B-N 51.5-71.4 (x = 62.0, N = 6) per- cent eye length; canthus rostralis rounded to obtuse; loreal region weakly concave, sloping gradually to lips; lips not flared; interorbital space flat, no cranial crests; upper eyelid 79.1-93.8 (x = 85.1, N = 6) percent IOD, bearing rounded, non- conical warts; tympanum concealed be- neath skin; upon drying, annulus visible, its length 32.5-48.5 (x — 40.4, N = 5) percent eye length, separated from eye by one annulus length; supratympanic fold not prominent; tympanic region not vertical; temporal region not swollen; postrictal tubercles not conical; choanae small, round, not concealed by palatal shelf of maxillary arch when roof of mouth is viewed from directly above; vomers lacking odontophores; tongue elongate (length 2% times width), not notched posteriorly, posterior % not ad- herent to floor of mouth; males with short vocal slits near angle of jaws, vocal sac internal. [TOL'80 Fic. 14. Palmar views of hands of (A) E. myersi, AMNH 86752, and (B) E. ocreatus, KU 117574. Scale equals 2 mm. MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 15. Dorsal and lateral views of heads of (A) Eleutherodactylus myersi, AMNH 86752 and 86753 and (B) E. ocreatus, KU 117574. Scale equals 2 mm. Skin of dorsum smooth anteriorly, finely areolate posteriorly and laterally, bearing low paravertebral and dorsolat- eral folds; skin of lower flanks coarsely areolate (warts subconical), that of ven- ter coarsely areolate; discoidal folds prominent; anal opening not modified; no inguinal glands; limbs smooth with subconical warts; ulnar tubercles not prominent; palmar tubercle bifid, as large as oval thenar tubercle; super- numerary palmar tubercles indistinct; subarticular tubercles non-pungent, broader than long; fingers lacking fringes and pads; discs (as long as wide) on fingers III and IV; first finger shorter than second. No tubercles clearly identifiable on heel or tarsus (tarsus is tuberculate, as is leg); inner metatarsal tubercle twice as long as wide, large, not compressed, two to two and one-half times size of round, non-pungent outer metatarsal tu- bercle; supernumerary plantar tubercles non-pungent; subarticular tubercles non- pungent, round or weakly elongate; toes lacking fringes and pads; hind legs very short, shank 32.8-41.1 (x = 369, N = 6) per cent SVL. Above brown with diffuse darker brown mottling; dark tan interorbital bar present or not; middorsal raphe from scapula to vent present or not, if present, cream edged with reddish-brown; indi- viduals having dorsal raphe have cream line down midline on venter and another (perpendicular) across breast; groin and axilla bearing cream spots; posterior sur- faces of thighs brown with cream mar- bling; venter dark brown, usually spotted or marbled with cream; hands and feet, except outer edge, white, as are top of foot and tarsus and underside of shank. In life, E. ocreatus was colored as follows: “Dorsum and upper flanks dark brown, in some a reddish-brown mid- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 29 dorsal stripe or broad dorsal band (= raphe); most have pale orange spots along dorsolateral folds; flanks and ven- ter black with white spots; posterior sur- face of thigh fuscous brown with pale yellow spots; the spots in axillae tend to pink; in some specimens, there is a cross on the venter, very pale pink; iris blue- gray; inner digits pink to cream” (J. D. Lynch field notes 23 February 1968). Measurements of holotype in mm.— SVL 19.6; shank 7.2; head width 6.8; head length 6.2; upper eyelid width 1.7; IOD 2.2; tympanum length 1.05; eye length 2.2; E-N 1.45. The holotype is a gravid female with extensively con- voluted oviducts. Etymology.—Latin, ocreatus, mean- ing wearing leggings or booted, in ref- erence to the pale hands and feet which appear clothed in gloves and socks. Natural history—The nine specimens were collected by day beneath rocks and clumps of dirt in non-burned areas of the pdramo. The area was recently burned but some enclaves of green grass remained; presumably these areas were wetter than the burned areas. Both adult females are gravid. The thumbs of the males are swollen but do not possess nuptial pads. The testes are dark brown (presumably black in life). The ovarian linings are peppered with brown in the three juvenile females. Remarks.—E. ocreatus is mostly closely related to E. trepidotus found at comparable or slightly lower elevations on the Amazonian versant of the Andes in northern Ecuador. The two are sim- ilar in size and proportions, morphology of the hands and feet, and in color pattern. Eleutherodactylus ocreatus is sym- patric with E. curtipes and E. thymelen- sis; both are larger frogs. Further col- lecting may show that the still larger E. buckleyi is sympatric with E. ocreatus in the Paramos del Angel. Distribution—Known only from the Paramos del Angél (3500-4150 m) in extreme northern Ecuador (Fig. 13). Eleutherodactylus orcesi Lynch Fig. 12C-D Eleutherodactylus orcesi Lynch, 1972:142 (holotype, KU 130316, an adult female ob- tained 5 km SW Bolivar-Chimborazo fron- tier on Guaranda-Ambato road [= 24.8 km by road N Guaranda], SW slope of Nevado Chimborazo, Prov. Bolivar, Ecuador, 3800 m, on 17 July 1970 by J. D. Lynch). Diagnosis.—(1) skin of head smooth, that of dorsum, flanks, and venter areo- late; no dorsolateral folds; (2) tympanum visible, its length % eye length; (3) snout short; snout round in dorsal and lateral profiles; canthus rostralis sharp; (4) LOD broader than upper eyelid; no cranial crests; no tubercles on eyelid; (5) vomer- ine odontophores absent; (6) males with vocal sac and slits; white nuptial pad on thumb in males; (7) first finger shorter than second; pads of outer fingers as large as tympanum; (8) fingers bear narrow lat- eral fringes; (9) ulnar tubercles absent; (10) no tarsal tubercles; a small heel tubercle; (11) two metatarsal tubercles, inner oval, 3-5 times size of outer; nu- merous supernumerary plantar tubercles; (12) toes bear lateral fringes; toe pads as large as those of fingers; (13) brown to black above with or without white warts on flanks; venter pale brown; no limb or lip markings; (14) adults small, males 23.9-29.6 (x = 26.5, N = 7) mm, females 35.2-36.1 (x = 35.8, N = 3) mm SVL. Eleutherodactylus orcesi is not easily confused with other species. Its nearest relative is E. thymelensis which is easily distinguished in having the tympana concealed and in having vomerine teeth. Description—(Lynch, 1972b). Variation.—Little variation is evident among the 17 available specimens. Fe- males as large as 31.2 mm (KU 130315) are immature (small eggs, narrow ovi- ducts). Specimens from the Paso San Juan (northerly locality, Fig. 12C) differ from those from the slopes of the Nevado de Chimborazo in lacking white spots on the flanks (Fig. 12D). Natural history —E. orcesi has been found beneath rocks and in terrestrial 30 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY bromeliads in paramos by day. No data on nocturnal activity are available. On the slopes of Nevado Chimborazo, E. orcesi is sympatric with E. curtipes. On the crest of the Cordillera Occidental SW of Quito (San Juan area) it is sympatric with E. curtipes and E. uni- strigatus. In July 1970 the type-series of three individuals was secured under a single rock. In January 1978 I turned over the same rock and found three more indi- viduals. In each instance, a gravid fe- male and a reproductively active male (swollen testes, white nuptial pads ) were found. Distribution—The Cordillera Occi- dental of central Ecuador between 3160 and 3800 m (Fig. 16). Eleutherodactylus thymelensis Lynch Fig. 12E Eleutherodactylus whymperi: Lynch, 1971:45, 146. Eleutherodactylus thymelensis Lynch, 1972:144 (holotype, KU 117719, an adult male ob- tained in the Paramo del Angél, ca. 23 km [airline] SW Tulcdn, Prov. Carchi, Ecua- dor, 3700 m, by J. D. Lynch on 24 Febru- ary, 1968). Diagnosis.—(1) skin of dorsum bear- ing large, flat granules, that of venter are- olate; paravertebral but not dorsolateral folds present; (2) tympanum concealed beneath skin; (3) snout round in dorsal and lateral profiles; canthus rostralis sharp; (4) no cranial crests; no enlarged eyelid tubercles; (5) vomerine odonto- phores oblique; (6) males with vocal slits, subgular vocal sac; (7) first finger ———! 76 Fic. 16. Distribution of Eleutherodactylus orcesi (@) and E. thymelensis (A). LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 31 shorter than second; pads and discs pres- ent; (8) fingers bear lateral fringes; (9) ulnar tubercles prominent; (10) an inner tarsal tubercle; less prominent tubercles on heel and outer edge of tarsus; (11) two metatarsal tubercles, inner oval, 5-6 times size of outer; numerous super- numerary plantar tubercles; (12) toes bear lateral fringes; toe pads and discs as large as those of fingers; (13) gray to rust-brown above with gray, tan, or black speckling, no color pattern; venter gray to brown, flecked or marbled with black; posterior surfaces of thighs brown; (14) adults small, males 21.4- Panemex) ==-23-6 a) 0.5, N=. 18). mm) females 28:0-33.5.(x = 31.1 = 14, N =i) mnass'V Li: Eleutherodactylus thymelensis is most similar to E. obmutescens from the Paramo de Puracé but differs in having a vocal sac and vocal slits, an exposed frontoparietal fontanelle, and smaller outer metatarsal tubercle. Description—Lynch (1972b). Natural history—E. thymelensis re- mains inexplicably rare. Although it was abundant on 23-24 February 1968 when first collected, subsequent efforts at other localities have yielded only four addi- tional specimens. All specimens known were collected by day beneath rocks. Five localities lie in Espeletia paramos (Fig. 7A-B) but the locality in Prov. Pichincha lacks Espeletia (Fig. 7C-D). DistributionKnown from five pa- ramo localities in southern Colombia and northern Ecuador at elevations between 3310 and 4150 (Fig. 16). Fic. 17. Eleutherodactylus unistrigatus pattern morphs. (A) spotted morph, Quito, not pre- served; (B) mid-dorsal line morph, KU 177476; (C) striped morph, Quito, not preserved; (D) striped morph, KU 138852. 32 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Eleutherodactylus unistrigatus (Ginther ) Figs. 12F, 17 Hylodes unistrigatus Giinther, 1859, Proc. Zool. Soc. London, 27:416 (cotypes, BMNH 60.6.16.97(2) and 60.6.16.105 [reregistered as 1947.2.17.7-9] from “western Ecuador” collected by Mr. Fraser). Hylodes lehmanni Boettger, 1892, Kat. Batr. Mus. Senckenburg. p. 26 (cotypes, Mus. Senckenburg 1200, la [not examined], col- lected by C. F. Lehmann, from the paramo del Bordoncillo, 3500 m [probably along the frontier between Departamento de Narifio and Intendencia Putumayo], Colom- bia, in 1885). New synonym. Phyllobates equatorialis Barbour, 1908, Bull. Mus. Comp. Zool., 51:320-21 (cotypes, MCZ 2261, 85082, from “Equator’’). Syrrhophus coeruleus Andersson, 1945, Arkiv for Zoologi, 37A(2): 24 (syntypes, NHRM 1913 (5 examples), “Eastern Ecuador” col- lected by Wm. Clarke-Macintyre 1937- 1938). New synonym. Diagnosis —(1) skin of dorsum sha- greened, with scattered flat warts, that of venter aerolate; no dorsolateral folds; (2) tympanum partially concealed, its length 4-% eye length; (3) snout short; snout subovoid to subacuminate in dor- sal view, round in lateral profile; canthus rostralis moderately sharp; (4) TOD broader than upper eyelid; no cranial crests; upper eyelid lacking obvious tu- bercles; (5) vomerine odontophores oblique, moderately prominent (absent in young individuals and some southern populations); (6) males with vocal slits, external subgular vocal sac; small white nuptial pad on thumb of males; (7) first finger shorter than second; pads of outer fingers as broad as tympanum; (8) in- distinct lateral fringes on fingers; (9) ulnar tubercles not evident except ante- brachial; (10) no tubercles on heel or outer edge of tarsus; indistinct fold or tubercle on inner edge of tarsus; (11) two metatarsal tubercles, inner oval, 3-4 times size of elongate outer; supernu- merary plantar tubercles present, most distinct at bases of toes; (12) toes bear narrow lateral fringes and distinct pads; (13) most pale brown (to cream) above with brown markings (interorbital tri- angle, dorsal chevrons [scapular, sacral, and suprainguinal], canthal-supratym- panic stripe); venter cream; groin finely stippled with brown (or unpigmented); anterior and posterior surfaces of thighs finely stippled with brown; polymorphic —see text; (14) adults small, males 14.9- 28.8 mm, females 22.5-38.5 mm SVL; geographically variable, see text. Eleutherodactylus unistrigatus is most similar to E. cajamarcensis and E. modi- peplus but differs in having the posterior surfaces of the thighs brown with (or without) cream reticulation, in lacking tubercles on the upper eyelid, usually lacking tubercles on the dorsum (present on the flanks), and in having less distinct supernumerary plantar tubercles. Description—Head narrower than, to as wide as, body in males, broader than body in adult females, wider than long; snout subovoid to subacuminate in dorsal view, round in lateral profile; nostrils weakly protuberant, directed dorsolaterally; canthus rostralis moder- ately sharp, straight to weakly concave; loreal region weakly concave, sloping abruptly to lips; lips not flared; inter- orbital space flat (lacking cranial crests in most individuals, weakly evident in adult females); upper eyelid lacking prominent tubercles; temporal region sloping; supratympanic fold present, ob- scuring upper edge of tympanum; tym- panum visible, not distinct, round in males, higher than long in females, sepa- rated from eye by 1-14 tympanum length; postrictal tubercles non-conical; choanae round, small to moderate-sized, not concealed by palatal shelf of maxil- lary arch; vomerine odontophores usually present, moderately prominent, median and posterior to choanae, oblique (slant- ed), widely separated medially (1-2 choanal widths), bearing 0-4 teeth on posterior edge, each odontophore smaller (to equal) than a choana; odontophores less elevated in males (frequently not visible in males); tongue 1-2 times as long as wide, posterior edge weakly notched, posterior 4-7 not adherent to floor of mouth; males with vocal slits LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 33 lateral to tongue; external, subgular vocal sac. Skin of dorsum shagreened with scat- tered flat warts, that on flanks bearing more numerous warts, that on venter and underside of thighs coarsely areo- late; no dorsolateral folds; anal opening not ornamented with tubercles; discoidal folds prominent, well anteriad of groin; upper surfaces of limbs coarsely sha- greened; ulnar tubercles not evident ex- cept for small, non-conical antebrachial; palmar tubercle bifid (or divided with large inner and small outer), 2 times size of oval thenar tubercle; supernu- merary palmar tubercles low, indistinct; subarticular tubercles round, elevated; fingers bearing indistinct lateral fringes; all fingers bearing discs (broader than long) on dilated pads (pads of outer fingers progressively larger than those on inner digits); pads apically rounded to truncate; those of fingers III and IV as large as tympanum;; first finger shorter than second. Knee, heel, and outer edge of tarsus lacking tubercles; inner edge of tarsus bearing indistinct fold on distal 4 (may appear as a tubercle); inner metatarsal tubercle twice as long as wide, not com- pressed, 3-4 times size of elongate, low outer metatarsal tubercle; supernumer- ary plantar tubercles numerous, low and indistinct, those at bases of toes II-IV moderately distinct; subarticular tuber- cles subconical, smaller than those of fingers; toes bearing narrow lateral fringes; toes bearing discs on pads; pads of toes about as large as those of outer fingers; heels of flexed hind limbs touch; heel of adpressed hind limb reaches to tympanum. The most common coloration and pattern (spotted morph, Pattern A, Table 7) is as follows: pale brown above with indefinite darker interorbital triangle, series of brown spots forming dorsal chevrons (extending onto flanks)—scap- ular, sacral, and suprainguinal (these spots sometimes nearly black); limb bars nearly transverse, about as broad as paler interspaces; canthal and supratym- panic stripes dark brown; labial bars in- distinct; posterior thigh finely stippled with brown, groin finely stippled with brown or not pigmented; venter cream (very finely stippled with brown), throat of large females sometimes dusky. A variant (mid-dorsal line morph, Pattern B, Table 7) of the above pattern differs only in having a narrow cream vertebral stripe running the length of the body. Another color pattern (striped morph, Pattern C, Table 7) consists of tan, medium-brown, and dark brown stripes on the back (Fig. 17). In this variant, a thin cream mid-dorsal stripe is usually visible. Specimens from the Quito-Otavalo and Latacunga areas never have the un- pigmented area in the groin. Frogs in the Pasto-Tulcan region usually have the unpigmented areas. These five northern populations are structurally homogeneous but differ in the frequency of striped morph pattern, size, and testes coloration. Specimens from the Quito-Machachi area uniformly have white testes without gray to brown reticulation (testis color #1, Table 7). Specimens from Otavalo-Cayambe have gray to brown reticulation on white testes (testis color #2, Table 7) or have white testes. Those from _ Provincia Carchi, Ecuador, and Departamento Narifio, Colombia, usually have brown testes with dark reticulation (or uni- formly black testes) (testis color #3, Table 7). Specimens from the vicinity of Latacunga resemble those from Ota- valo. Females from Prov. Carchi, Ecua- dor, and Depto. Narifio, Colombia, fre- quently have pigmentation of the mesen- tary covering the ovaries. The striped morph is less common in collections from Quito than from else- where in northern Ecuador and southern Colombia (Table 7). The highest fre- quency of striped frogs is in extreme northern Ecuador but this may reflect small samples. Two subsamples from Carchi Province have 28.6 and 53.3% striped individuals respectively (N of 14 and 15). Four subsamples from De- 34 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY partamento de Narifio include 8.7, 12.0, 12.9, and 53.8% striped frogs. Four sub- samples from Otavalo include 9.1, 12.9, 14.7, and 17.5% striped frogs. These con- trast with three subsamples from Quito of 3.0, 4.3, and 14.0% striped frogs. The striped morph of E. unistrigatus was confused with E. buckleyi by Boulenger (1882: pl. 14, fig. 5) who unfortunately illustrated this cotype (BMNH 78.1.25.- 42), an action which led Peracca (1904) to report at least two species having a similar morph as Hylodes buckleyi. In life, E. unistrigatus is pale yellow- brown to medium brown or terra cotta (if brown, often with terra cotta blotches dorsally ); dorsum spotted with darker brown and/or black; flanks creamy brown flecked with brown; limbs same color as dorsum, sometimes slightly more pale; some specimens have creamy yellow flecks superimposed on pattern; throat of male bright yellow, venter yellow-white; in females, throat flesh- colored and/or flecked with brown, ven- ter white; posterior surfaces of thighs a pale version of dorsum, yellowish in males; iris gray-bronze to silvery, with many black flecks, dark brown horizon- tal streak. Variation—In addition to the vari- ation in color patterns and testes color, E. unistrigatus exhibits geographic vari- ation in body size (Table 7). Males from extreme northern Ecuador and ad- jacent Colombia are markedly smaller than males from other areas in Ecuador. The trend is not so evident among fe- males because female E. unistrigatus from Quito are small. In general, the largest individuals come from those pop- ulations in the Ambato-Latacunga hoya. Specimens from 5-10 km W Bajos, Prov. Tungurahua, are distinctive in their smaller tympana (Table 8) and because the vomerine odontophores are very indistinct (scarcely elevated). These frogs have odontophores (at least on one side, if not both, but very few teeth [0-2/odontophore]. The posterior sur- faces of the thighs are cream marbled with brown. The skin is more pustulose than that seen in most populations of E. unistrigatus. Geographic variation in E. unistri- gatus is more gradual than in E. cur- tipes. Adjacent populations do not ex- hibit markedly different proportions or coloration (Tables 7-8) This can be ex- plained in that populations of E. unistri- gatus are more contiguous than are populations of E. curtipes. Natural History—As mentioned in the account for E. modipeplus, E. uni- strigatus breeds between December and February. Calling males can be heard from Tulcén to Riobamba below eleva- tions of 3200 m. In January, 1978, I drove from Quito to Otavalo and in mesic areas heard an unbroken chorus of E. unistrigatus, occasionally joined by enclaves of Gastrotheca riobambae. De- velopment of E. unistrigatus was de- scribed by Nina and del Pino (1977). In spite of having collected thousands of individuals of this frog, I have found only a single egg mass. KU 180302 was found 18 January 1978 under a clod of dirt 6.9 km SE San Rafael, Prov. Imba- bura, 2900 m. The embryos, in an ad- vanced stage of development (large fan- like tail, limbs well developed), were in a small cavity 2 x 2 & 1 cm with no avenue of access into the clutch. The 17 eggs were not adherent. Young frogs are abundant in June in Quito but only adults are encountered in December. Presumably the frogs ma- ture in a single year. The seasonality of reproduction is marked. I heard no calling males in June through August, even in pastures in cloud forests (e.g., Pilal6 or El Car- melo). This seasonality makes sense in the relatively arid interandean valleys but is striking when duplicated in cloud forests. The population of E. unistrigatus at Pilalé, Provincia Cotopaxi, Ecuador, 2400 m, is isolated from the contiguous populations in the interandean valleys of Ecuador by the paramos de Apaqua but in mid-January 1978, E. unistrigatus was calling throughout the pastures and cleared areas in the vicinity of Pilald. LEPTODACTYLID FROGS OF THE CENUS ELEUTHERODACTYLUS 35 TaBLe 7. Geographic variation in body size, color pattern, and color of testes in Eleutherodactylus unistrigatus. Locality Depto. Narifo Prov. Carchi Otavalo Cayambe Quito Machachi Latacunga Pilalo Ambato W of Banos Riobamba Pungala Pattern Males Females A B C 17.7-23.6(17)* 24.0-31.1(26) 76.8 3.6 19.6 20.2+0.8 28.1+0.7 14.9-23.2(25) 23.3-30.8 (46) 55.2 3.4 41.4 18.8+0.8 26.3+0.4 20.5-25.6( 26) 26.5-34.7 (32) 77.6 ee 15.2 22.7+0.6 30.4+0.7 Re PEL ee en 78.8 1.9 19.2 17.7-28.8(39) 22.5-32.7(61 ) 92.2 0.0 Hel 22.1+0.8 27.7+0.6 SIT Gaye, 81.8 0.0 18.2 18.0-28.1(12) 24.8-35.2(23) 85.7 0.0 14.3 24.8+1.4 30.7+1.2 14.5-24.0( 46) 23.0-31.0( 22) 65.8 0.0 34.2 19.1+0.7 26.8+0.9 19.3-28.5( 49) 26.8-38.5( 49) 92.7 0.0 3 24.8+0.6 31.9+0.8 20.2-23.9(9) 27.4-33.9(18) 100.0 0.0 0.0 22.6+1.0 30.2+0.9 20.2-28.2 (20) 24.7-35.7(20) 100.0 0.0 0.0 22,.6+0.8 29.8+1.4 tee, 100.0 0.0 0.0 Testes Tile Spas Sie OLN GNO Sie low Te ee ad) 0 ken 30) 0°) AG NT os ge 2idte< 9 15) 0) 9 ’ First line, range (N); second line +2SE. TABLE 8. Proportions of Eleutherodactylus unistrigatus. First line, range (N); second line, mean +2SE. Locality Shank/SVL HW/SVL Eyelid/IOD Tympanum/Eye E-N/Eye NGTNO: ge 5 ss @ 40.3-46.5(15) 34.9-38.6(15) 75.8-100.0(14) 29.0-44.1(14) 76.5-91.2(15) 42.4+1.0 37.20.5 91.0+3.7 Slals=2o 84.7+2.5 Wtavalo™.. 6 4.< » é 43.4-44.7(5) 35.9-39.0(5) 74.1-96.0(5) 37.5-44.4(5) 84.6-91.3(5) 44.0 37.8 83.0 40.5 86.9 sy ieee 9 40.1-47.3(8) 36.0-38.7(8) 83.3-97.1(8) 36.4-45.9(8) 76.3-88.6(8) 44,0+1.6 37.8+0.8 89.5+3.4 41.1+2.3 83.5+2.8 @uiton. 3 = ss @ 40.0-45.7(13) 30.8-39.1(13) 67.3-100.0(13) 33.3-52.0(13) 76.9-92.0(13) 42.7+1.0 36.8+1.1 88.5+5.1 42.2+3.0 82.4+2.4 Sha earn Q 39.9-44.2(17) 35.1-39.5(17) 71.0-90.3(16) 32.7-51.7(17) 71.0-100.0(17) 41.6+0.6 37.0+0.6 Sie7 E27 43.7+2.8 88.1+3.2 Ambato-Riobamba ¢ 42.2-47.2(20) 35.6-39.4(20) 65.7-90.5(20) 34.0-50.0(20) 76.7-88.5(10) 44.3+0.6 37.3+0.4 78.9+3.1 43a les 82.7+2.4 Brees caus 9 41.6-48.5(20) 34.8-38.7(20) 60.0-88.0(20) 43.0-58.3(20) 77.8-93.3(11) 44.7+0.8 36.7+0.5 74.4+3.1 50.0+1.9 85.3+2.8 5-10 km W Banos ¢ 42.4-48.3(9) 33.2-37.2(9) 69.2-92.0(9) 23.1-38.7(9) 71.0-84.6(9) 44,.5+1.2 35.0+0.8 83.0+4.3 30.1+3.5 18.33.3 Aeon @ 41.7-49.1(18) 32.8-38.2(18) 80.0-116.7(17) 24.2-41.4(18) 73.0-100.0(18) 44,9+1.0 35.8+0.6 91.2+4.4 34.5+2.3 85.0+3.0 36 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY The call is a hollow click. To my ear, it sounds like the noise produced by rapping two hollow bamboo tubes to- gether and is very much unlike the whistle of E. modipeplus. During the day, E. unistrigatus is concealed, usually beneath objects on the ground (rocks, logs, boards, trash, grass), but individuals have also been found in terrestrial bromeliads, and in arboreal bromeliads (up to 3 m high in trees at Lago Cuicocha). At night, in- dividuals are found on vegetation as well as on the ground. Calling males perch on vegetation to at least 1 m above the ground. Lynch (1969, 1970) described microhabitat differences between sexes (at least during breeding activity ). In interandean Ecuador (Fig. 18), E. unistrigatus is most abundant between 2500 and 3200 m, but has been collected as high as 3490 m (Nudo de Tiopullo, Prov. Cotopaxi, Eduador). The contig- uous interandean populations enter the upper Pastaza Trench (as low as Banos, 1800 m). At least three, presumably iso- lated populations occur on the Andean slopes. The Pilaléd (2400 m) population was cited above. A second population (Mindo-Nono area, 1500-2530 m)_ is found in Provincia Pichincha. The third occurs in extreme northern Ecuador (E] Carmelo 2710 m, Prov. Napo). This population may be contiguous with in- terandean populations across the low (3200) frailejon (Espeletia) paramo de las Juntas SE of Tulcan. Remarks.—Barbour and Noble (1920) pointed out that Phyllobates equatorialis Barbour is a synonym of E. unistrigatus. Hylodes lehmanni Boettger and Syr- thophus coeruleus Andersson are also synonyms of E. unistrigatus. Hylodes lehmanni is founded on a striped morph of E. unistrigatus. Except for the color pattern, this morph is identical to the more common spotted morph. Cochran and Goin (1970) emphasized the striped Fic. 18. Eleutherodactylus unistrigatus habitats. pasture habitat, 3 km SSW San Miguelito, Prov. Tungurahua, 2620 m; (C) pasture, 1 km N Otavalo, Prov. Imbabura, 2560 m; (D) brickyard in west Quito—E. unistrigatus was abundant around the pond and in other mesic microhabitats at this site in 1967. (A, B) Aspect and detail of Eucalyptus- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 37 pattern in their description of E. leh- manni but in no subsample of specimens from Pasto region does the frequency of the striped morph exceed 54%. The larg- est specimen I have seen from Colombia is USNM 146232 (31.1 mm). Boettger (1892) reported the larger cotype to be 33 mm SVL and to have a spotted belly. The specimens reported by Cochran and Goin (1970) from Departamento de Caldas, Colombia (including the exam- ple 37 mm long) are striped specimens of an ally of E. vertebralis (Boulenger), E. supernatis Lynch. Syrrhophus coeruleus Andersson is based on five individuals I believe to be conspecific with the material from 5-10 km W Banos, Prov. Tungurahua, Ecua- dor. In these frogs vomerine teeth and odontophores are poorly developed and the posterior surfaces of the thighs are marbled with brown. Based on brief color notes by the late James A. Peters and my study of the preserved speci- mens, I tentatively assign them to E. unistrigatus. However, study of living specimens may reveal this to be a dis- tinct species, in which case Andersson’s name is available. The situation in interandean Ecuador in the Bafios—Ambato region and on the surrounding andean spurs is suggestive of confounding of two or more species. In addition to E. modipeplus and E. uni- strigatus (essentially identical to speci- mens from Quito), the coeruleus-like frogs, and E. pastazensis Andersson (see Lynch and Duellman, 1980, for rede- scription) occur in this restricted region. Specimens from 7-10 km W Banos in- clude frogs with small white testes and frogs with large black testes. Correlated (partially ) with the differences in testes color, are relative distinctness of the tympana, skin texture, distinctness of marbling on the posterior surfaces of the thighs, and duskiness of the venter. I am reluctant to propose additional names until the localities have been recollected, colors in life recorded from fresh mate- erial, and acoustical data are available. Distribution—The Nudo de Pasto in Colombia south to Riobamba in Ecuador through the interandean valleys. Isolated and semi-isolated populations on the andean flanks in northern and central Ecuador (Fig. 9). DISCUSSION The elutherodactyline fauna of high altitude habitats of southern Colombia and northern Ecuador is impoverished compared with that of southern Ecuador (Lynch, 1979). The northern fauna es- sentially consists of three species—E. buckleyi, E. curtipes, and E. unistrigatus. The other six species (E. modipeplus, E. myersi, E. ocreatus, E. orcesi, E. thy- melensis, and Phrynopus brunneus) are rare species having small distribution areas. All eight Eleutherodactylus are species of the wnistrigatus group as de- fined by Lynch (1976) and represent four assemblies within that group: curtipes assembly—E. buckleyi, E. cryophilius, and E. curtipes are very similar frogs whose distributions tend to replace one another (Fig. 19). myersi assembly—E. ginesi (Rivero), E. myersi, E. nicefori Cochran and Goin, E. ocreatus Lynch, E. orestes Lynch, E. trepidotus Lynch, and E. vidua Lynch form a series of species characterized by their minute digital pads. This assembly is distributed primarily on the eastern Cordillera (Fig. 20). orcesi assembly—E. obmutescens Lynch, E. orcesi Lynch, E. racemus Lynch, E. simoterus Lynch, and E. thy- melensis Lynch form a series of five allopatric populations on the Cordillera Central of Colombia and Cordillera Oc- cidental of Ecuador (Fig. 21). unistrigatus assembly—E. cajamar- censis Barbour and Noble and E. versi- color Lynch in southern Ecuador, E. modipeplus Lynch, E. pastazensis An- 38 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY buckley/ Ne : Fic. 19. Distribution of the curtipes Assembly (buckleyi—curtipes—cryophilius). dersson, and E. unistrigatus (Ginther) in southern Colombia and northern Ec- uador, and E. bogotensis (Peters) and E. lynchi Duellman and Simmons on the Cordillera Oriental in Colombia (Fig. 22). Two distributional patterns are ap- parent. The first, involving the species of the curtipes and orcesi assemblies, in- volves distribution along the Cordillera Occidental of Ecuador and the Cordil- lera Central of Colombia, whereas the myersi and unistrigatus assemblies are displaced eastward. The second is a pattern of allopatric and disjunct distri- bution of pairs of relatives (E. bogoten- sis - E. lynchi in Colombia; E. ocreatus - E. trepidotus in northern Ecuador; E. cryophilius - E. curtipes in central Ec- uador; E. ginesi - E. niceforoi in Colom- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 39 Fic. 20. Distribution of the myersi Assembly in Colombia and Ecuador (nicefori—myersi—ocre- atus—trepidotus—orestes—vidua). bia and Venezuela; the five species of the orcesi assembly) and is contrasted to the less common partial ecologic-geo- graphic separations as seen in E. buck- leyi - E. curtipes, E. cajamarcensis - E. versicolor, E. modipeplus - E. unistri- gatus. Most of the species of these four assemblies are inhabitants of cool, moist environments. The two exceptions, E. pastazensis and E. unistrigatus, are re- lated. The former is poorly known (Lynch and Duellman, 1980). The dis- tribution of E. wnistrigatus is most prof- itable contrasted with that of E. curtipes (Figs. 8-9). The two are sympatric at only 10 localities, ranging in altitude from 2750 to 3450 m. The relatively lower sites (Tulcan, Machachi, Lata- cunga) are very wet sites and the mois- 40 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY orces! Fic. 21. Distribution of the orcesi Assembly (simoterus—racemus—obmutescens—thymelensis— orcesi). ture seemingly allows the less common E. curtipes to survive there. At the higher sites (13 km SSE Tulcan, Intag, San Juan, Péramo de Tiopullo, 10 km W Cotalé), E. unistrigatus was the less common. Higher altitudes may provide too cool an environment for reproduction for E. unistrigatus. The geographic variation in E. uni- strigatus (relatively uniform cline) con- trasts with that seen in E. curtipes (punctuated clines). These discordant patterns of geographic variation seem consistent with the notion that the dis- tribution of E. unistrigatus was recently contiguous and is only now fragmenting whereas that of E. curtipes is now ex- panding towards contiguity. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS Al unistriga I< SS modipeplus ~ ae ‘ \ \ LY J cajamarcensis versicolor : _—sy | 80 76 oS bogotensis X - / ad wii a 72 Fic. 22. Distribution of the unistrigatus Assembly (lynchi—bogotensis—unistrigatus—modipeplus —cajamarcensis and versicolor). SUMMARY The paramos and subparamos of northern Ecuador and southern Colom- bia are populated by eight species of Eleutherodactylus. The most widely dis- tributed species are E. buckleyi, E. cur- tipes, and E. unistrigatus. Eleutherodac- tylus modipeplus new species, E. myersi, E. ocreatus new species, E. orcesi, and E. thymelensis have smaller distribution areas. Two new species are named, E. modipeplus from montane regions near Ambato, Ecuador, and E. ocreatus from the paramo del Angél west of Tulcan, Ecuador. Hylodes whymperi and Hyla 42, MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY chimboe are considered synonyms of Eleutherodactylus curtipes. Hylodes lehmanni and Syrrhophus coeruleus are considered synonyms of Eleutherodac- tylus unistrigatus. The eight species of Eleutherodac- tylus are members of the dominant cur- tipes group and are sorted into four assemblies. The curtipes assembly (E. buckleyi, E. cryophilius, and E. curtipes) and the orcesi assembly (E. obmute- scens, E. orcesi, E. racemus, E. simo- terus, and E. thymelensis) are distrib- uted on the Cordillera Central of Co- lombia and in the northern Andes of Ecuador whereas the myersi assembly (E. ginesi, E. myersi, E. nicefori, E. ocreatus, E. orestes, E. trepidotus, and E. vidua) and the unistrigatus assembly (E. bogotensis, E. cajamarcensis, E. lynchi, E. modipeplus, E. pastazensis, E. unistrigatus, and E. versicolor) are dis- tributed on the Cordillera Oriental of Colombia (and one assembly extends onto the Venezuelan Andes) and the Cordillera Real of Ecuador (to the Huancabamba depression). The two distribution patterns for as- semblies may reflect the same dynamic interaction seen between two widely dis- tributed Ecuadorian species (E. curtipes and E. unistrigatus). Eleutherodactylus curtipes is distributed at higher altitudes than E. unistrigatus and appears to be in the process of range expansion whereas E. unistrigatus appears to be experiencing range fragmentation. RESUMEN Ocho especies de Eleutherodactylus se encuentran en los paramos y sub- paramos del norte de Ecuador y de Colombia meridional. Las especies con distribuciones amplias son: E. buckleyi, E. curtipes, y E. unistrigatus. Eleuthero- dactylus modipeplus (especie nueva), E. myersi, E. ocreatus (especie nueva), E. orcesi, y E. thymelensis tienen distri- buciones menores. Se nombran dos es- pecies nuevas, E. modipeplus, de las regiones altas cerca Ambato, Ecuador, y E. ocreatus, del paramo del Angél al oeste de Tulcdn, Ecuador. Hylodes whymperi y Hyla chimboe son consid- erados sindnimos de Eleutherodactylus curtipes. Hylodes lehmanni y Syrrhophus coeruleus son sinénimizados con Eleu- therodactylus unistrigatus. Las ocho especies de Eleutherodactyl- us son miembros del grupo unistrigatus (el grupo mas dominante en esta parte de América del Sur) y se asignan a cuatro asambleas (= subgrupos del grupo unistrigatus). La asamblea cur- tipes (E. buckleyi, E. cryophilius, y E. curtipes) y la asamblea orcesi (E. ob- mutescens, E. orcesi, E. racemus, E. simoterus, y E. thymelensis) se encuen- tran en la Cordillera Central de Colom- bia y los Andes al norte de Ecuador, mientras que la asamblea myersi (E. ginesi, E. myersi, E. nicefori, E. ocreatus, E. orestes, E. trepidotus, y E. vidua) y la asamblea unistrigatus (E. bogotensis, E. cajamarcensis, E. lynchi, E. modi- peplus, E. pastazensis, E. unistrigatus, y E. versicolor) se encuentran en la Cor- dillera Orential de Colombia (y una es- pecie también se encuentra en los Andes venezolanos) y la Cordillera Real de Ecuador (hasta la depresién de Huan- cabamba). Los dos modelos de distribucién para las asambleas posiblemente reflejan la misma accion reciproca dindmica que se ve en las dos especies ecuatorianas con amplias distribuciénes (E. curtipes y E. unistrigatus). Eleutherodactylus curtipes se encuentra en alturas mayores que E. unistrigatus y parece estar en proceso de expandir su distribucién mientras que E. unistrigatus se encuentra en sitios mas bajos y su distribucién aparentemente esta contrayéndose. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 43 LITERATURE CITED Acosta-Souis, M. 1968. Divisiones fitogeogra- ficas y formaciones geobotanicas del Ecuador. Casa de Ja Cultura Ecuadori- ana, Quito, Ecuador. 309 pp. ANDERSSON, L. G. 1945. Batrachians from east Ecuador collected 1937, 1938 by Wm. Clarke-Macintyre and Rolf Blomberg. Arkiv for Zoologi, 37A(2):1-88. Barsour, T. 1908. Some new reptiles and amphibians. Bull. Mus. Comp. Zool. 51:315-325. Barsour, T., Nose, G. K. 1920. Some am- phibians from northwestern Peru, with a revision of the genera Phyllobates and Telmatobius. Bull. Mus. Comp. Zool. 63:395-427. BorttcErR, O. 1892. Katalog der Batrachier- Sammlung im Museum der Sencken- bergischen Naturforschenden Gesell- schaft in Frankfurt am Main. pp. 1-73. BouLENGER, G A. 1882. Catalogue of the Batrachia Salientia s. Ecuadata in the collection of the British Museum. 2nd ed. 503 pp. Cocuran, D. M., Gorn, C. J. 1970. Frogs of Colombia. Bull. U.S. Natl. Mus. (288): 1-655. Fow.er, H. W. 1913. Amphibians and rep- tiles from Ecuador, Venezuela and Yu- catan. Proc. Acad. Nat. Sci. Philadelphia 65:153-176. Gorn, C. J., Cocuran, D. M. 1963. Two new genera of leptodactylid frogs from Co- lombia. Proc. California Acad. Sci. 31: 499-505. Guntuer, A. C. L. G. 1859. Second list of cold-blooded Vertebrata collected by Mr. Fraser in the Andes of western Ecuador. Proc. Zool. Soc. London 1859: 402-422 (issued between Oct. 1859 and Feb. 1860). Lyncu, J. D. 1968. Systematic status of some Andean leptodactylid frogs with a de- scription of a new species of Eleuthero- dactylus. Herpetologica 24:289-300. Lyncu, J. D. 1969. Taxonomic notes on Ecua- dorian frogs (Leptodactylidae: Eleu- therodactylus). Herpetologica 25:262- QTA. Lyncu, J. D. 1970. Identity of two Andean Eleutherodactylus with the description of a new species (Amphibia: Leptodac- tylidae). J. Herpetol. 3:135-143. Lyncu, J. D. 1971. Evolutionary relationships, osteology, and zoogeography of lepto- dactyloid frogs. Misc. Publ. Mus. Nat. Hist. Univ. Kansas, no. 53: 1-238. Lyncu, J. D. 1972a. Systematics and ecology of robberfrogs in western Ecuador. Year- book Amer. Philos. Soc. 1971:332-333. Lyncu, J. D. 1972b. Two new species of frogs (Eleutherodactylus: | Leptodactylidae ) from the paramos of northern Ecuador. Herpetologica 28:141-147. Lyncu, J. D. 1976. The species groups of the South American frogs of the genus Eleu- therodactylus (Leptodactylidae). Occas. Pap. Mus. Nat. Hist. Univ. Kansas, no. 61:1-24. Lyncu, J. D. 1979. The frogs of the genus Eleutherodactylus in the Andes of south- ern Ecuador. Misc. Publ. Mus. Nat. Hist. Univ. Kansas, no. 66:1-62. Lyncu, J. D. 1980. A taxonomic and dis- tributional synopsis of the Amazonian frogs of the genus Eleutherodactylus. Amer. Mus. Novitates (2696) :1-24. Lyncu, J. D., DuELLMaAN, W. E. 1980. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes. Misc. Publs. Mus. Nat. Hist. Univ. Kansas, no. 69:1-86. Nina Hey i Pino” Ve eho Me per. 197% Estructura histologica del ovario del sapo Eleutherodactylus wunistrigatus y observaciones sobre el desarrollo em- brionario. Revista Universidad Catdlica 5(16):31-41. Peracca, M. G. 1904. Viaggio del Dr. Enrico Festa nell’Ecuador e regioni vicine. Rettili ed Amphibii. Boll. Mus. Zool. Anat. Comp. 19( 465) :1-41. APPENDIX: SPECIMENS EXAMINED Eleutherodactylus buckleyi (255 spec.) COLOMBIA, Depto. Cauca: Paramo de las Papas, MLS 144; Paramo de Puracé, 3400 m, KU 145018; Paramo de Puracé, Laguna de San Rafael, 3200 m, ICN 1539, 1549-55, 2082- 84, 2086, 2088, 2090, 2599, KU 143896-933, UMMZ 166582, UVMP 794-800; 17 km E Puracé, 3300 m, KU 167904-16, 168431, WED 42427-28: 18 km E Puracé, Rio Cocuy, 3320 m, KU 167903; 19 km E Puracé, 3420 m, KU 167918-23, 170167; 23 km E Puracé, 3275 m, KU 167924-30, 170125-26 (CS); 26 km E Puracé, 3130 m, KU 167931-44; 30 km E Puracé, 3030 m, KU 167945-48. Depto. Huila: Hda. Meremberg, Santa Leticia, 2400 m, ICN 2576. Depto. Narino: ca. 4 km N Laguna de la Cocha, 3200 m, AMNH 86755-60; Paramo 44 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY El Tabano, 3200 m, ICN 2498-99, 2502, 2504; Municipio de Pasto, lado norte del Volcan Galeras, 3310 m, UVMP 2867, 2870; 8 km NE Pasto, 3020 m, KU 167949-50; 12 km E Pasto, 3050 m, KU 167951-56, WED 42816-19; 4 kn S Pasto, UVMP 2881; Tuquerres, 3000 m, KU 138724. Intend. Putumayo: 13 km W Santiago, 2900 m, KU 167957-58; 16 km W Santiago, 3230 m, KU 167959-60. Depto. Valle (de Cauca): Paramo de las Hermosas, 3570 m, KU 167965; 7 km NE Tenerife, 2850 m, KU 167961; 13 km NE Tenerife, 3310 m, KU 167962-63; 15 km NE Tenerife, 3460 m, KU 167964. ECUADOR, Prov. Carchi: El Carmelo, 2713 m, USNM 204745-47; 10 km (airline) WNW EI Carmelo (=10 km SSE Tulcan), 3201 m, USNM 204742-44, 204748-53; 5.7 km NW EI Carmelo, 2910 m, KU 177235-41; 10.8 km NW EI Carmelo, 3050 m, KU 177231-34; 14.6 km NW EI Carmelo, 3130 m, KU 177226- 30; Tulcan, 2770 m, KU 177225; 13 km SSE Tulcan, 3018 m, USNM 204733-41; 14 km SW Tulcan, 3340 m, KU 165150-58, 23 km SW Tulcin, 3700 m, KU 117558-66. Prov. Imba- bura: Intag, BM 78.1.25.40-41, 78.1.25.43; Urcusiqui, 3300-3500 m, KU 117492-515. Prov. Napo: Laguna San Marcos, + 10000 feet, CAS-SU_ 8288-99; Santa Barbara, 2591 m, MCZ 92500-01, USNM GOV 7168-70, 7218, 8547, 8551-52, 8554. Eleutherodactylus curtipes (1388 spec. + 7 lots of eggs) ECUADOR, no other locality, MNHNP 06- 276. Prov. Bolivar: Arenal, 3 km SW frontier on Ambato-Guaranda road, 4000 m, KU 130521-25; Arenal, 6 km SW frontier on Am- bato-Guaranda road, 3800 m, KU 130526-43, 177800-01; 16-19 km E Guaranda, 3500-3800 m, KU 130466 (lot of eggs), 130467-519; Guanujo, 2923 m, KU 177802; mountains above San Jose del Chimbo, 3049-3293 m, ANSP 18137 (holotype of Hyla chimboe); Sinche (= Sinchig), 7 leagues E Guaranda, 4000 m, BM 99.10.30.38-40. Prov. Carchi: 10 km WNW El Carmelo, 3201 m, USNM 211575-80; 14.6 km NW EI Carmelo, 3130 m, KU 177748-54; Tulcan, 2770 m, KU 177736-41, MZS 8653-54; 14 km SW Tulcan, 3340 m, KU 165270-92; 20 km SW Tulcan, 3350 m, KU 165243-69; W slope Volcan Chiles, 10 km W Tufino, 3500- 3600 m, KU 117556-57, 118107 (lot of eggs). Prov. Chimborazo: 9 km S Cajabamba, 3260 m, KU 165241-42; 12 km SW Cajabamba, 3800 m, KU 119633-56; Desierto de Palmira, near Palmira, 3110 m, KU 165239-40; 20 km E Guaranda, 3811 m, USNM 211581-82; Gual- labamba, 3500 m, BM 99.10.30.35-37; 15 km SE Pungala, Hda. Alao, 3100 m, KU 130591- 94; near Rosario, 3500 m, BM 1932.10.2.54; 10 km W San Juan, 3400 m, KU 119573-632; Tortorillas, 4024 m, BM 82.7.13.4-5/RR 1947. 2.17.22-23 (cotypes of Hylodes whymperi); Urbina, 3609 m, KU _ 130596-640, UMMZ 132902(8), USNM 211583-609. Prov. Coto- paxi: Guilo, Paramo de Apagua, 8 km E Pilald, 3500 m, KU 131357-402; Latacunga, 2750 m, KU 177743; Paramo de Apagua, 3800-3860 m, KU 131334-56, 131696; Paramo de Milin, 3900 m, KU 119682-715; Paramo de Tigua, 3450 m, KU 130465; Paramo de Tiopullo, 3490 m, KU 130753-830; slopes of Volcan Cotopaxi, 3620- 3750 m, KU 123492-94. Prov. Imbabura: Coto- cachi, 3963 m, BM 82.7.13.3/RR 1947.2.17.24 (cotype of Hylodes whymperi); Intag, BM 78.1.25.29-36 (cotypes of Hylodes curtipes); Lago Cuicocha, 3010 m, KU 177742; Nudo de Mojanda, 3400-3440 m, KU 130726-51; Nudo de Mojanda, 3650-3680 m, KU 130648-724. Prov. Leén: Satacunga, UMMZ 55522 (this locality is probably Latacunga, Prov. Cotopaxi; Leon was the older name for Prov. Cotopaxi). Prov. Napo: east slope, Paso Guamani, 3950 m, KU 165376-400, 3650 m, KU 109009-58; 10 km W Papallacta, 3610 m, KU 110804-44; 9 km W Papallacta, 3800 m, MCZ 90018; 5 km W Papallacta, 3232-3400 m, KU 110845- 90, 177746-47, 180286-89 (lots of eggs), USNM 211717-800; Laguna Papallacta, 3330- 3350 m, KU 108990-9008, 165293-375, WED 47707-11, MCZ 95556-58; 1 km W Papallacta, 3155-3200 m, KU 110891-95, USNM 211695- 716 + 2 without tags; 2.7 km NW Papallacta, 3300 m, MCZ 91913-30; 18.9 km NW Papal- lacta, road to Pifo, 3880 m, MCZ 91931-37; Papallacta, 3110 m, UIMNH 55898-915, 55918- 21, USNM 211693-94; 3 km E Papallacta, 2900 m, KU 117567-69. Prov. Pichincha: MCZ 3021-22; 6-7 km W Chillogallo, ca. 3000 m, UMMZ 132905 (21); Machachi (Univer- sidad Catdélica Quito); 17 km E Pifo, Paso Guamani, 3800 m, KU 110794-803; trib. Rio Pita, N base Cerro Ingaloma, 3780 m, KU 123495-500; 3 km SW Quito, 3200 m, USNM 211688-92; San Juan, 3400 m, KU 177744-45; Volcan Pichincha, 3400-3800 m, KU 130831- 69; W slope Volcan Pichincha, 4400 m, CAS- SU 10628. Prov. Tungurahua: Arenal, 2 km NE frontier on Ambato-Guaranda road, 3900 m, KU 130520; 7 km W Banos, 2805 m, USNM 211661-82; 8 km W Banos, 2805-2927 m, USNM 211657-60; 10 km W Banos, 2927-3049 m, USNM 211610-56; 10 km W Cotald, 3300 m, KU 130641-47; 10-10.5 km SSW Mocha, 3450-3480 m, KU 177755, 177757-99, 180290 (lot of eggs), 180291 (4); 10 km SW Mocha, 3700 m, KU 119657-81; 12 km SW Santa Rosa, 3400 m, KU 130566-90; 18 km SW Santa Rosa, 3600 m, KU 130556-65; 20 km SW Santa Rosa, 3700 m, KU 130544-55; 10 km N_ Urbina, Puente de Mallo, 2974 m, USNM 211683-85, JAP 6432; 12 km N Urbina, USNM 211686-87. The following are surely the product of LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 45 data confusion: FMNH 196028-29, 196035, 196037-38, 196041-42, 196044-45, 196047, 196050-51, 196053-54, 196056-59, purportedly from Bogota, Depto. Cundinamarca, Colombia, but probably from the vicinity of San Jose (Chillogallo to Chiriboga road), Prov. Pichincha, Ecuador [the Field Museum has a series of E. bogotensis catalogued as having originated from Ecuador; both series of specimens were purchased from E. H. Taylor]. Eleutherodactylus modipeplus (43 spec.) ECUADOR, Prov. Chimborazo: Gualla- bamba, 3500 m, BM 99.10.30.41-43; 1 km S Ur- bina RR station, 3650 m, KU 131283-85. Prov. Tungurahua: 8.1 km S on Banos-Riobamba road, 2560 m, KU 141296: 10.9 km S on Banos-Riobamba road, 2750 m, KU 141297; 10 km N Mocha, Puente de Mallo, 3079 m, UMMZ 132896(3), USNM 212036-38; 2 km S Mocha, 3000 m, USNM 212031-35; 10 km SW Mocha, 3700 m, KU 120018-21; 10-10.5 km SSW Mocha, 3450-3480 m, KU _ 179361-64; Patate, 16 km SSE, 2940 m, KU 141306, 16.5 km SSE, 2950 m, KU 141303-05, 17.1 km SSE, 2970 m, KU 141307-08, 20.6 km SSE, 3120 m, KU 141298-302; 3 km SSW San Miguelito, 2620 m, KU 131276-80; NW slope Volcan Tungurahua, 2730 m, KU 142132-34. Eleutherodactylus myersi (20 spec.) COLOMBIA, Depto. Cauca: Puracé high- way, eastern slopes, 2900-3000 m, KU 143956; Puracé highway at San Juan Agua Terminales, 3000 m, KU 143954; 23 km E Puracé, 3275 m, KU 168432-35; 26 km E Puracé, 3130 m, KU 168436-38; 30 km E Puracé, 3030 m, KU 168439. Depto Narino: ca. 4 km N Laguna de la Cocha, 3200 m, AMNH 86751-53; Paramo El Tabano, 3200 m, ICN 2503; 12 km E. Pasto, 3050 m, KU 168440-44; “20 miles east of Pasto,” CAS 85177 (holotype). Eleutherodactylus ocreatus (10 spec.) ECUADOR, Prov. Carchi: E] Pelado, 4150 m, MNHNP 03-217; W slope Volcan Chiles, 10 km W Tufino, 3500-3800 m, KU 117573-81. Eleutherodactylus orcesi (17 spec.) ECUADOR, Prov. Bolivar: 5 km SW Boli- var-Chimborazo frontier, SW slope Nevado Chimborazo (= 24.8 km by road N Guaranda), 3800 m, KU 130314-16, 177814-16; 3 km W Bolivar-Chimborazo frontier, 3500 m, KU 130303; 21.7 km E Guaranda, 3410 m, KU 140003-04; 27.3 km E Guaranda, 3800 m, KU 140002. Prov. Chimborazo: 7.2 km WNW San Juan, 3160 m, KU 140001. Prov. Pichincha: 6-7 km W Chillogallo, ca. 3000 m, UMMZ 132903(4); Km 26 Quito-Chiriboga road, USNM 212039; 2 km E San Juan, UMMZ 132904. Eleutherodactylus thymelensis (72 spec.) COLOMBIA, Depto. Narino: Mpio Pasto, N slope Volcan Galeras, 3310 m, UVMP 2865-66. ECUADOR, Prov. Carchi: E] Pelado, 4150 m, MNHNP 03-218 4 220; Péramo del Angél, 20 km SW Tulcan, 3350 m, KU 165944, ca. 23 km SW Tulcan, 3700 m, KU 117719-70; W slope Volcan Chiles, 10 km W Tufifio, 3500- 3600 m, KU 117706-18. Prov. Pichincha: Km 924, Lago Agrio-Batan road (2.4 km W of crest), 3860 m, KU 177860. Eleutherodactylus unistrigatus (1346 spec. + 1 lot eggs) COLOMBIA, Depto. Narifio: El Encano, 2630 m, ICN 1325-29, 4805-11; 7 km NE Guachucal, 3000 m, AMNH 86763-64, 86779; Guaitarilla, 2000 m, FMNH 54372, 54393-411, KU 138726-28; Laguna de la Cocha, 2800 m, AMNH 87874; Laguna de la Cocha, N shore, 2790-2850 m, AMNH 84853, 86765-77, KU 168587-617, 170148-49 (cleared and _ stained skeletons ); 4 km N Laguna de la Cocha, 3200 m, AMNH 86762; Paramo El] Tabano, 3200 m, ICN 2500; Municipio de Pasto, lado norte de Volcan Galeras, 3040 m, UVMP 2862; Pasto, Barrio San Fernando, LACM 50206-37; 8 km NE Pasto, 3020 m, KU 168567-83, 170147 (cleared and stained skeleton); 12 km E Pasto, 3050 m, KU 168584-86; west of Pasto, USNM 146220-35, 146238-47; Tangua, UVMP 2872; Tuqueres, FMNH 54373. Intend. Putumayo: Colon, 2220 m, KU 168618-24; 4 km SE San Francisco, 2320 m, AMNH 84837-38; Sibun- doy, 2200 m, MCZ 56207-35, 58015-23. ECUADOR, no other locality, MCZ 2261, 85082 (cotypes of Phyllobates equatorialis) ; “west Ecuador,’ BM _ 60.6.16.97/RR 1947.2. 17.7-8, 60.6.15.105/RR 1947.2.17.9 (cotypes of Hylodes unistrigatus). Prov. Carchi: Atal, near San Gabriel, 2700 m, UMMZ 83675(25), 83676(27), 83677(26), 92153(6), 92154(5); El Carmelo, 2710 m, KU 177521-28; 5.7 km NW El Carmelo, 2910 m, KU 177529-32; 10 km WNW EI Carmelo, 3201 m, USNM 212064- 67; Tulcan, 2770 m, KU 117623-36, 177472- 512: 13 km SSE Tulcan, 3018 m, USNM 212051-52; 3 km S Tulcdn, 2851 m, USNM 212053-63. Prov. Chimborazo: 1 km SW Guallabamba, 2860 m, KU 119750-92; 1 km S Guano, 2600 m, KU 131076; 8 km SE Licto, road to Alao, 3000 m, KU 141382-96; below Pungala, at Rio Alao, 2800 m, KU 141397-403; 1.6 km SE Pungala, 3055 m, KU 141404-10; 8.2 km SE Pungalé, 3085 m, KU 141411-14; 8.5 km SE Pungalé, 3090 m, KU 141415-17; 11 km SE Pungala, 3050 m, KU 131075; Rio- bamba, 2780 m, KU 119793; 15 km NW Rio- bamba, 3040 m, KU 165590. Prov. Cotopaxi: 46 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Chillo (0°51’S, 78°34’W), 2745 m, MCZ 3023- 27, 3034(3), UMMZ 47217; 2 km SE Lasso at Rio Cotuchi, 2957 m, USNM 212068-69; Lat- acunga, 2750 m, KU 177515-17; 7 km E, 6 km S Latacunga, 2750 m, KU 127014; 1 km W Latacunga, 2750 m, KU 127015, 127024-30; Mulalé, 2990 m, KU 146124-35; 0.3 km N Mulald, 3005 m, KU 141368; 1 km NE Mulald, 3050 m, KU 127016-23, 127035-36 (skel.); 1 km S Mulalé, 3005 m, KU 141367; 0.3 km W Mulalé, 2980 m, KU 141366; Nudo de Tio- pullo, El] Chasqui, 3490 m, KU 130978; Pilald, 2400 m, KU 143557, 177888-969; 11.3 km W Pujili, 3500 m, KU 141369-71. Prov. Imba- bura: Intag, BM 78.1.25.42 (cotype of Hylodes buckleyi); La Floresta, 2995 m, CAS-SU 9486- 90; Laguna Cuicocha, 10 km W Quiroga, 2890-3000 m, KU 135351-92, 138788, 138125, 138832-51, 177513-14, MCZ 95546-54; Ota- valo, 2550 m, KU 135397-427; near Otavalo, 2600 m, UMMZ 92152(11), 92155(8); 1 km N Otavalo, Quebrada San Miguel, 2650 m, KU 117638-47, 117666-67; 8 km NW _ Otavalo, 3100 m, KU 117698-705; 10-12 km NW Ota- valo, 3300 m, KU 117674-97; 13-15 km NW Otavalo, 3300-3500 m, KU 117668-73; Quiroga, 2500 m, KU 135343-46; 7 km W. Quiroga, 2950 m, KU 135347-50; 4 km S San Pablo, NW slope Nudo de Mojanda, 3050 m, KU 131489-532; 6.9 km SE San Rafael, 2900 m, KU 180302 (lot of eggs); Urcusiqui, 12 km NW Otavalo, 3300 m, KU 117648-65. Prov. Napo: ca. 25-30 km SE Cayambe, 3190 m, CAS-SU 8280; Cotapino River (? locality), CAS-SU_ 10338(2); Santa Barbara, 2591 m, USNM 212070-72. Prov. Pichincha: 16 km W Aloag, 2810 m, KU 111132-35; 5 km W Aloag, N slope Cerro Corazon, 2945 m, KU 109066- 67; 7 km W Aloag, 2810 m, KU 109068-69; 0.5 km N Cayambe, 2820 m, KU 130931-77; 6-7 km W Chillogallo, 3000 m, UMMZ 132905(4); Machachi, MCZ 3010, USNM 212138-48; region of Mindo, USNM 212150- 51; Molino Ureu, AMNH 20139; 1 km E Nanegal Grande, 1524 m, UMMZ 132895(3); 2 km W Nono, USNM 212136-37; 9.5 km NW Nono, 2530 m, KU 165591-92; Pacto, USNM 212149; Quito, 2850 m, CAS-SU 2273, 2275- 79, 10622-27, KU 94391, 111085-91, 138852- 75, 152010-12, 177469-71, USNM 212073-79; 1.6 km ENE Quito, 2896 m, USNM 212132- 33; 3.2 km ENE Quito, 2896 m, USNM 212134-35; 1.6 km SSE Quito, Rio Machangara, 2744 m, USNM 212122-26; 2.4 km SSE Quito, 2744 m, USNM 212127-31; 2.4 km W Quito, SE slope Volcan Pichincha, 3500 m, USNM 212083-95; 1.6 km NW Quito, SE slope Vol- can Pichincha, 3000 m, USNM 212080-82; 3.2 km NW Quito, 2988-3049 m, USNM 212096- 121; San Juan, 3400 m, KU 177518-20; Volcan Pichincha, E. slope, 2900 m, KU 111092-131, 111136-37 (C&S skel.), 3480 m, KU 152013- 25, 3390 m, KU 152026-31, S slope, 3480 m, KU 130979-80. Prov. Tungurahua: Ambato, 2700 m, KU 119794-800; Banos, 1900 m, CAS- SU 5083, UIMNH 55722, KU 141379-80; 5 km W Banos, 2439 m, USNM 212152-73; 7 km W Banos, 2805 m, USNM 212174-88, JAP 5938; 10 km W Banos, 2927-3049 m, USNM 212190-210; Chambo Grande, 7.6 km SE Peli- leo, 2340 m, KU 141372, 141975-76, 146121- 23; 10 km W Cotalé, 3300 m, KU 130897-930; 1 km W Juan Benigo Vela, 3080 m, KU 130981-89; Hda. Leito, Cerro Llanganti, 2439- 2744 m, USNM 212189, 212211; 4 km N Mocha, 3140 m, KU 119801-04; 3 km SSW San Miguelito, 2620 m, KU _ 130990-1074, 131272-75; W slope Volcan Tungurahua (6.1 km SW jct. Rio Pastaza and Banos-Riobamba road), 2470 m, KU 141373-74; W slope Vol- can Tungurahua (13.2 km SW jet. Rio Pastaza and Banos-Riobamba road), 2760 m, KU 141375-78; NW _ slope Volcan Tungurahua, 2730 m, KU 141977-85. L668.E257 L93 Harvard MCZ Librar ailing 3 2044 062 RECENT MISCELLANEOUS PUBLICATIONS UNIVERSITY OF KANSAS MUSEUM OF NATURAL HISTORY 52. Reproductive cycles in lizards and snakes. By Henry S. Fitch. Pp. 1-247, 16 fig- 53. 55. 57. 59. 61. 62. 65. 66. 67. 68. 69. 70. 71. ures in text. June 19, 1970. Paper bound. Evolutionary relationships, osteology, and zoogeography of leptodactyloid frogs. By John D. Lynch. Pp. 1-238, 131 figures in text. June 30, 1971. Paper bound. Middle American lizards of the genus Ameiva (Teiidae) with emphasis on geo- graphic variation. By Arthur C. Echternacht. Pp. 1-86, 28 figures in text. De- cember 14, 1971. Paper bound. A systematic review of the Teiid lizards, genus Bachia, with remarks on Heter- odactylus and Anotosaura. By James R. Dixon. Pp. 1-47, 15 figures in text. February 2, 1973. Paper bound. Systematics and evolution of the Andean lizard genus Pholidobolus (Sauria: Teiidae). By Richard R. Montanucci. Pp. 1-52, 8 figures in text. May 14, 1973. Paper bound. Reproductive strategies in a tropical anuran community. By Martha L. Crump. Pp. 1-68, 13 figures in text. November 15, 1974. Paper bound. A demographic study of the ringneck snake (Diadophis punctatus) in Kansas. By Henry S. Fitch. Pp. 1-53, 19 figures in text. April 3, 1975. Paper bound. The biology of an equatorial herpetofauna in Amazonian Ecuador. By William E. Duellman. Pp. 1-352, 198 figures in text. August 30, 1978. Paper bound. Leptodactylid frogs of the genus Eleutherodactylus from the Andes of southern Ecuador. By John D. Lynch. Pp. 1-62, 23 figures in text. February 28, 1979. Paper bound. An ecogeographic analysis of the herpetofauna of the Yucatan Peninsula. By ‘i C. Lee. Pp. 1-75, 27 plates, 22 figures in text. February 29, 1980. Paper ound. Internal oral features of larvae from eight anuran families: Functional, systematic, evolutionary and ecological considerations. By Richard Wassersug. Pp. 1-146, 37 figures in text. June 24, 1980. Paper bound. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). By John D. Lynch and William E. Duellman. Pp. 1-86, 8 fig- ures in text. August 29, 1980. Paper bound. Sexual size differences in reptiles. By Henry S. Fitch. Pp. 1-72, 9 figures in text. February 27, 1981. Paper bound. Late Pleistocene herpetofaunas from Puerto Rico. By Gregory Pregill. Pp. 1-72, 26 figures in text. May 8, 1981. Paper bound.