Lara Luncly . eSy:. Herp. QL 668 MAR 1 »E257 NIVERSITY OF KANSAS re ARVAI MISCELLANEOUS L92. \USEUM OF NATURAL HISTORY UNIVERSIT gk cert ia Leptodactylid Frogs of the Genus Eleutherodactylus from the Andes of Southern Ecuador By John D. Lynch UNIVERSITY OF KANSAS LAWRENCE 1979 February 28, 1979 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. Tue UNIVERSITY OF KANSAS MusEuUM OF NATURAL HiIsToRY MISCELLANEOUS PUBLICATION No. 66 February 28, 1979 Leptodactylid Frogs of the Genus Eleutherodactylus From the Andes of Southern Ecuador By Joun D. Lyncu Associate Professor of Zoology, School of Life Sciences The University of Nebraska, Lincoln, Nebraska 68588 and Associate in Herpetology, Museum of Natural History The University of Kansas, Lawrence, Kansas 66045 THe UNIVERSITY OF KANSAS LAWRENCE 1979 UNIVERSITY OF KANSAS PUBLICATIONS, MUSEUM OF NATURAL HISTORY Editor: E. O. Wiley Miscellaneous Publication No. 66 pp. 1-62; 23 figures; 4 tables Published February 28, 1979 MusEuM oF NATURAL History Tue UNIVERSITY OF KANSAS LAWRENCE, Kansas 66045 Wi saAe PRINTED BY UNIVERSITY OF KANSAS PRINTING SERVICE LAWRENCE, KANSAS CONTENTS lake © 18) Lo) CO) Ngee ee sy 8 6 Pe nee ie, Ae eee J BN GERSON ED NAUTINER ee ae ee 2 Key To ELEUTHERODACTYLINE FROGS OF SOUTHERN ANDEAN ECUADOR _- + RECOUNTS OF “SEE CINS 22 ae). iS Ls Se SE ea 5 Elenthnerodactylus atratus mew Species: =. 2 5 mieusnerodactylus: balionotus mew Species 21 7 Eleutherodactylus baryecuus new species ____---------- 10 Eleuthenodactylus bromeliaceus new species: — 2 2 12 Eleutherodactylus cajamarcensis Barbour and Noble _------------------- 14 Eleutherodactylus colodactylus new species ____.--------------------------------- 15 Hlemnerodactylus cryopihilius mew species 22.2) 19 Eleutherodactylus cryptomelas new species = 21 Hleutherodactylus lymani Barbour and Noble _...22 = 23 iguiherodactylus orestes mew. Species, = 24 Eleutnerodactylus percultus new species. 26 Eleutherodactylus phoxocephalus new species ___..---- 29 Eleutherodactylus proserpems new species = = 32 Eleutherodactylus pycnodernnis new species 35 Hicuinenodactylusemociio Despax)) £2 o.xs2e a ee eee 37 Eleutherodactylus muidus new species 2 40 Elenthenodactylus: spinosusmew, SPCCleS =.= se 43 Elcutherodactylus versicolor mew. species =) 22 45 inieutherodactylus tidud mew species == 8 ee ee 49 Ficutnerodactylus w-merum (Boettger) 2 51 DES GS STO IN pg can ee eee 51 Retoushipssor tae southern fauna 2 aes eee ee eee 51 eo locicalmseO reed Oi) eats oi Se eee Zen ee eee ee ee ee 52 Correlates with species densities and diversities __... 53 Comparisons with distributions of other Andean genera 56 SHON ABNAUAUG oN hese ae ee Mere 2 eR as Sees OEE Le, Joe de ee Gees 57 Pie Uy ULE IN escaeactcnee rene Pe Oe eS Re a Oe 58 TE PTE Ra SVAN TB OE Re. DY Se) el ee teenie de cae 58 BPE NID] SPECIMENS EXAMINED e223 60 INTRODUCTION The genus Eleutherodactylus is a prominent component in most faunas from northwestern South America. The genus is well represented in the faunas of the lowlands as well as the high An- dean pdramo and pajonal (of Acosta- Solis, 1968) habitats. Such habitats gen- erally occur at elevations above 3000 m. The Andes of northern Ecuador have considerable contiguous areas at eleva- tions above 3000 m, whereas in southern Ecuador (south of the Nudo de Azuay ) areas above the 3000 m contour are dis- continuous (Fig. 1). This patchiness of high montane territory continues into northern Pert, where south of the Hu- os ~ B x ry 5 - y i & ancabamba pass the Andes again have vast areas of contiguous high altitude habitats which grade into the puna in southern Pert and Bolivia. The genus Eleutherodactylus is found in the high montane localities in Ecuador and ex- treme northern Pert, but south of Huan- cabamba, suitable habitat is meager or other frogs occupy the habitats occupied by Eleutherodactylus at more northern localities. The eleutherodactyline fauna of northern Andean Ecuador is now rela- tively well known (at least taxonomical- ly). In contrast, that of the Andes of southern Ecuador is poorly known. | \ Wy A \ AREA ABOVE 3000 METERS ELEVATION 76 74 Fic. 1—Map of Ecuador illustrating the distribution of Eleutherodactylus phoxocephalus (circles). The area in southern Ecuador enclosed by the box is shown in greater detail in Figure 2. 2 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Lynch (1969a) reported E. cajamarcen- sis Barbour and Noble from the moun- tain ranges to the north and east of the Loja hoya and E. lymani Barbour and Noble from the Loja hoya. Phrynopus flavomaculatus (Parker) is found at high altitudes on the mountain ranges to the north and east of the Loja hoya and is distributed along the Cordillera Ori- ental to east of Cuenca (Lynch, 1975). The type-locality of Eleutherodactylus w-nigrum (Boettger) is on the interior slopes of the Cordillera Occidental just west of Cuenca. In June 1968 I visited a mountain range approximately 15 km E of Cuidad Loja, Ecuador (Abra de Zamora), the type-locality for Phrynopus flavomacu- latus. Five species of eleutherodactyline frogs were collected on the crest and the western slopes in 1968. William E. Duellman collected there in 1971 and 1975 and obtained five other species. In 1968, 1970, and 1971, several collectors made collections of eleutherodactyline frogs in pdramos and pajonales at many localities in the Andes south of Cuenca. In addition to these collections, those made by the late James A. Peters were made available for study; Peters’ collec- tions were made along two transects crossing the Cordillera Oriental to the northeast and east of Cuenca. Within the area I term southern An- dean Ecuador (Fig. 2), 21 species of eleutherodactyline frogs are found. These include one species of Phrynopus (P. flavomaculatus) and 20 species of Eleutherodactylus. Two of the Eleu- therodactylus belong to the predomi- nantly lowland fitzingeri group: 1) E. lymani, found in the interandean valleys from the Rio Jubones south to the val- leys of the Rio Catamayo and Rio Zaru- milla (draining to the Pacific) as well as in the valleys of the Rio Huancabamba and Rio Mayo or Chinchipe (draining to the Amazon), and 2) E. w-nigrum, which was found at one pajonal (the type-locality); E. w-nigrum also occurs on both the Amazonian and the Pacific slopes peripheral to the high Andean regions. The other 18 species belong to what I called the unistrigatus group (Lynch, 1976a). All are termed “southern” even though one ranges considerably north- ward as a Pacific versant frog (Fig. 1). Only two of the 18 southern species of the unistrigatus group have been named. Lynch (1969a) redescribed E. cajamar- censis on the basis of specimens from the Loja hoya and also reported speci- mens from north-central Andean Ecua- dor. At present I do not consider the frogs reported from Provincia Tungura- hua by Lynch to be conspecific with the frog found in southern Ecuador. The status of those northern specimens will be discussed in a paper treating E. uni- strigatus. A species commonly found on the mountains surrounding the Cuenca hoya is E. riveti (Despax). This name had not been previously associated with any specific Andean population. In the species accounts the following abbreviations are used: SVL (snout- vent length), E-N (eye-nostril dis- tance), and IOD (interorbital dis- tance). Adult females include only those females having extensively convu- luted oviducts and/or large, yellow, Ovarian eggs. ACKNOWLEDGMENTS Field work in Ecuador was sup- ported by grants from the Watkins Fund of the Museum of Natural History, The University of Kansas, the Penrose Fund of the American Philosophical Society, and by the University of Nebraska Re- search Council. Robert W. Henderson accompanied me during the summer of 1968. William E. Duellman, Thomas H. Fritts, Richard Montanucci, and Linda Trueb have contributed substantially by making collections and recording notes on habitat and colors in life at the ex- pense of their own field-oriented studies on other groups. The late James A. Peters made his collections and those of Gustavo Orces- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 3 MOO \ \ \ MACHALA R Jubones Bre. 2 a CUENCA N Picacho : % x ° & % N x Map of southern Ecuador showing localities at which Eleutherodactylus discussed in the text were collected. Land masses above 3000 m are hatched; dotted line traces the 2000 m contour. V. available for study. Material and/or provision of working space were made available by Werner C. A. Bokermann, William E. Duellman, Alice G. C. Gran- dison, Jean Guibé, W. Ronald Heyer, Ronald A. Nussbaum, Douglas Ross- man, Richard Thomas, Charles F. Walk- er, Ernest E. Williams, and George R. Zug. I am especially grateful to Alice Grandison, Jean Guibé, Konrad Klem- mer, and Ernest Williams for loan of type-specimens or photographs of type specimens. Jaime Péfaur kindly _pro- vided the Spanish summary. Lastly, to those in agreement with Barbour’s (1921:25) dictum, my apol- ogies. MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Abbreviations for collections used throughout the text are: . Tympana absent American Museum of Natural History BMNH British Museum ( Natural History ) AMNH KU University of Kansas Museum of Natural History LSUMZ Louisiana State University Museum of Zoology MCZ Museum of Comparative MHNP UMMZ USNM WCAB Zoology, Harvard University Muséum National d Histoire Naturelle ( Paris ) University of Michigan Museum of Zoology National Museum of Natural History ( Washington ) Private collection of Werner Bokermann (Sao Paulo) KEY TO ELEUTHERODACTYLINE FROGS OF SOUTHERN ANDEAN ECUADOR . Skin of venter smooth; thumb longer than: second: fimeer 2 Skin of venter areolate; thumb shorter than second finger + PDicits lackimGqdises. .2ts2 2 Sa eee Phrynopus flavomaculatus All digits bearing discs on pads 3 Inner tarsal fold present ___-._-.____- Eleutherodactylus lymani Novimmner tarsal’ fold = 2. = = 2 ee Eleutherodactylus w-nigrum . Digits bearing narrow pads (Fig. 7) Rt As nee 9 Ree nk Oe ea, 5 Digits bearing moderate to broad pads} (Rigs: 4:20). ge . Large frogs (29-50 mm SVL), cra- Mialcrestsupresene a= eee = = = eee Pde Eleutherodactylus cryophilius Small frogs (18-27 mm SVL) lack- MapeCraiiall rests = 6 , Groin. dot pattermed 2 22 5 Eleutherodactylus vidua Groin black with white spots — Eleutherodactylus orestes Tympana clearly visible or con- CC Ale Clah SERS A os MINN o =. 27 Ate 10 . Small frogs (< 22 mm SVL) with- out vomerine odontophores; digits short and thick 10. EE Larger frogs (25-43 mm SVL), vo- merine odontophores evident 9 . Skin of dorsum smooth with scat- tered low warts; tip of snout round- ed; fingers not fringed _____________ Eleutherodactylus baryecuus Skin of dorsum warty; tip of snout pointed; fingers bearing _ lateral fringes __ Eleutherodactylus ruidus Prominent conical tubercles on eye- lids. and heels: =. 1 No tubercles on eyelid and heel, or if tubercles present, their length = basal width (not elongate) — 13 Skin of dorsum tuberculate; males lacking vocal slits and sac; venter usually reticulated with brown _ 12 Skin of dorsum smooth with longi- tudinal ridges; males with vocal sac and slits; venter white, occasionally with brown spots (not reticulate) — Eleutherodactylus atratus . Adults lacking cranial crests; skin of dorsum tuberculate; posterior sur- face of thigh: black _ .: === (itses Eleutherodactylus cryptomelas Adults bearing low cranial ridges; skin of dorsum finely tuberculate (numerous pustules on lower back); posterior surface of thigh black en- closingacreammspots’ == eae Eleutherodactylus spinosus LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 5 13. Outer edge of tarsus bearing row of conical ‘tubercles! 22 14 Outer edge of tarsus lacking conical tubercles SNS «ey 6 Se 18 14. Skin of dorsum smooth or bearing flattened warts posteriorly — 15 Skin of dorsum finely to coarsely UD ERCUIAtC sa eee ee 16 15. Small frogs (adults less than 30 mm SVL), skin of dorsum smooth —- am Eleutherodactylus bromeliaceus Large frogs (adults 30-40 mm SVL), skin of dorsum shagreened anterior- ly, bearing flat warts posteriorly —__ LAs Eleutherodactylus percultus 16. Posterior surface of thigh brown with cream flecks, pattern of dor- SUN Agee) (6 aan ee cranes Be ens Eleutherodactylus versicolor Posterior surface of thigh black without cream flecks ___.. 17 17. Groin, posterior thigh, and con- cealed shank black = __. Eleutherodactylus cryptomelas Concealed surfaces of hind limb dull gray — Eleutherodactylus balionotus 18. Fingers bearing prominent lateral fringes ___ Eleutherodactylus riveti Fingers lacking lateral fringes or having narrow, keel-like lateral hgh eee ee ee oe we 19 19. Skin of body areolate, digits short _ ssp Eleutherodactylus proserpens Skin of dorsum finely shagreened _ fpr ct tant Se Rae ech DS shah iS hd 20 20. Large black blotches on flanks and concealed surfaces of limbs nal Eleutherodactylus pycnodermis Flanks not black, posterior thigh sur- face marbled or spotted oi 21. Tip of snout bearing vertical keel; posterior surface of thigh yellow, marbled with brown or gray __-..- __ Eleutherodactylus phoxocephalus Tip of snout rounded; posterior surface of thigh unicolor or dark Spotted wate winite serene Eleutherodactylus cajamarcensis ACCOUNTS OF SPECIES In the following accounts I attempt to make comparable statements in the diagnoses and descriptions. The diagno- ses consist of a series of 14 numbered statements (or set of statements ) and fol- low the format used in several other papers on Ecuadorian species of Eleu- therodactylus (Lynch, 1968a, 1969a, 1970a, b, 1971, 1972a, 1974a, b, and 1976b). The sequence was used (but without numbers) in Lynch (1968b, 1969b) and Lynch and Schwartz (1972). Following the numbered statements is a paragraph distinguishing the taxon from those species it most nearly resembles (in my opinion). Measurements and proportions are given for each sex when the means differ significantly (p = 0.05); otherwise these data are com- bined. Complete accounts are provided for each of sixteen species. The accounts for E. cajamarcensis, E. lymani, and E. w-nigrum include only updating infor- mation [for more detail on these species see Lynch (1969a) and Cochran and Goin (1970) ]. Eleutherodactylus atratus new species Fig. 3A Holotype —USNM 199675, an adult female, obtained at Suro Rancho, Pro- vincia Morona-Santiago, Ecuador, 2683 m, by James A. Peters on 23 August 1962. Paratypes.—USNM_ 199679-82, topo- types; between Sapote and Suro Rancho, 2604-22 m, USNM 199683-89. Diagnosis.—(1) skin of dorsum bear- ing low ridges, venter areolate; (2) tym- panum round, 4-% diameter of eye; (3) snout subacuminate in dorsal view, short, E-N less than length of eye; (4) interorbital space broader than upper eyelid; cranial crests low; (3) vomerine odontophores oval, oblique; (6) males 6 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY with subgular vocal sac and vocal slits; (7) first finger shorter than second; all digits bearing broad discs on dilated pads; (8) fingers lacking lateral fringes; (9) ulnar tubercles indistinct; (10) heel and outer edges of tarsus bearing conical tubercles; short inner tarsal fold present; (11) two metatarsal tubercles, outer round, % size of non-compressed, elon- gate inner; numerous supernumerary plantar tubercles; (12) toes bearing nar- row lateral fringes, all bearing discs and pads as large as those of fingers; (13) tan above with brown stripes; flanks, concealed surfaces of thigh, shank, and tarsus densely spotted with black; venter ‘creamy-yellow, sometimes bearing small brown spots; (14) adults small, ¢ ¢ 17.4-24.0 mm, 9 2 24.9-29.2 mm SVL. Eleutherodactylus atratus is a mem- ber of the wnistrigatus group and might be confused with only E. cryptomelas. It is smaller than E. cryptomelas, has vocal sac and slits, is less tuberculate, and colored differently. They are simi- lar in having heel, tarsal, and eyelid tubercles, and in having black areas on the concealed surfaces of the hind leg. Description—Head narrower than to as wide as body, wider than long; head width 34.6-38.6 percent SVL (x = 36.8, N = 29); snout subacuminate in dorsal view, rounded in lateral profile, short, E-N 64.7-96.4 percent eye length in males (Xx = 82.6, N = 18), 78.9-96.7 percent in females (x = 88.2, N = 10); nostrils weakly protuberant, directed dorsolaterally; canthus rostralis sharp, straight; loreal region weakly concave, sloping abruptly to lips; lips not flared; interorbital space broad, edges of fron- toparietals upturned (low cranial crests); upper eyelid bearing 1-2 conical warts; width of upper eyelid 64.3-91.7 percent IOD in males (x = 74.1, N = 18), 59.4-78.1 in females (x = 67.4, N = 10); supratympanic fold present, con- cealing upper edge of tympanum; tym- panum distinct, round, directed postero- laterally, separated from eye by two- thirds its diameter, its length 23.5-44.4 percent eye length (x = 33.9, N = 28); no enlarged postrictal tubercles; choanae small, round, not concealed by palatal Fic. 3.—(A) Eleutherodactylus atratus, KU 165236; (B) E. balionotus, KU 142136; (C) E. caja- marcensis, KU 120007; (D) E. colodactylus, KU 165219. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS he shelf of maxillary arch; vomerine odon- tophores prominent, median and_ pos- terior to choanae, oval in outline, slanted, bearing 4-5 teeth in a slanted row, sep- arated by distance equal to one-half odontophore width, each 3-5 times size of a choana; tongue as long as wide, pos- terior % not adherent to floor of mouth, posterior border not notched; males with vocal slits and subgular vocal sac. Skin of dorsum bearing numerous low ridges; on lower back, ridges break up forming rows of small warts; flanks smooth; venter and posteroventral sur- faces of thighs coarsely areolate; discoi- dal folds prominent; anal opening not extended in sheath; upper limb surfaces coarsely shagreened; ulnar tubercles present, small and indistinct; palmar tubercle bifid, larger than oval thenar tubercle; supernumerary palmar tuber- cles round, pungent, smaller than round, non-conical subarticular tubercles; fin- gers lacking lateral fringes, keeled later- ally; all fingers bearing discs (broader than long) on dilated pads; pad of thumb small, round; those on outer fin- gers broad, rounded apically, slightly larger than ear; thumb shorter than sec- ond finger. Knee lacking tubercles; heel bearing small, conical tubercle; outer edge of tarsus bearing indistinct, subconical tubercles; inner edge of tarsus bearing indistinct fold; inner metatarsal tubercle twice as long as wide, not compressed, 24-3 times size of round outer metatarsal tubercle; numerous plantar supernumer- ary tubercles; subarticular tubercles round, non-conical; toes bearing narrow lateral fringes; outer edge of sole bear- ing keel; all toes bearing discs (broader than long) on dilated pads; pads of toes as large as those of outer fingers; heels overlapping when legs flexed at right angles to sagittal plane; heel of ad- pressed hind limb reaching to eye; shank 44.5-52.9 percent SVL (x = 48.8, N = 29). In preservative, tan to pale brown above with pale brown stripes; flanks and limbs gray-brown; canthal and su- pratympanic stripes pale brown; no la- bial bars; limbs not or only faintly barred, if present, bars oblique; flanks, axillae, groin, anterior thigh, posterior thigh, concealed shank, and concealed tarsus spotted with black; spotting on thighs and shank sometimes so dense as to appear solid black with cream spots or lines; extent of black increases onto- genetically; venter creamy-yellow with or without small brown spots. In life, E. atratus is yellowish brown above with pale brown stripes; flanks more yellowish than dorsum; groin, con- cealed thigh and shank surfaces deep black with shiny white spots; venter cream. Measurements of holotype in mm.— SVL. 27.3, shank 13.7, head width 10.0, head length 9.6, width of upper eyelid 2.0, IOD 3.2, tympanum 1.2, length of eye 3.2, E-N 2.7. The holotype is a gravid female. Etymology.—Latin, meaning dressed in black, in reference to the black areas on the concealed surfaces. Natural history—James Peters and his associates collected calling males at night on bushes and by day caught in- dividuals of both sexes beneath rocks and logs along the trail between Gua- laceo and General Plaza. The species was found only on the more southerly of the two transects made by Peters and his associates across the Paramos de Matanga. All specimens were found on the Amazonian slopes. William E. Duellman found 4 individuals in terres- trial bromeliads on the Abra de Zamora. Distribution.—Eleutherodactylus _at- ratus occurs at elevations of 2195-2850 m on the Amazonian slopes of the eastern Andean Cordillera in southern Ecuador. Eleutherodactylus balionotus new species Fig. 3B Holotype—KU 142136, an adult fe- male from 13.5 km E Loja, at the crest of the cordillera (Abra de Zamora) be- tween Provincia Loja and_ Provincia Zamora-Chinchipe, Ecuador, 2800 m, 8 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY obtained 22 July 1971 by William E. Duellman. Paratypes—KU 142135, 142137-44, collected syntopically with the holotype. Diagnosis. —(1) skin of dorsum tu- berculate, that of venter areolate; no dorsolateral folds; (2) tympanum visi- ble, round to slightly higher than long, its length two-fifths to three-fifths that of eye; (3) snout subacuminate in dorsal view, rounded in lateral profile; E-N slightly greater than eye length; (4) in- terorbital space flat, no cranial crests; upper eyelid width subequal to IOD; (5) vomerine teeth and odontophores present, round; (6) male with vocal slits and subgular vocal sac; (7) first finger shorter than second, all digits bearing discs on dilated pads, dilation ratios I: 4a 6 156, TV. 62 (8) “ingers bearing lateral fringes; (9) ulnar tuber- cles present, obscure; (10) inner tarsal surface bearing indefinite tubercles, out- er surface bearing a row of tubercles; heel bearing non-conical, but distinct, tubercles; (11) two metatarsal tuber- cles, outer round, conical, one-fourth to one-half size of oval, non-compressed in- ner; numerous indefinite supernumerary plantar tubercles; (12) toes bearing lat- eral fringes, not webbed; toes bearing discs on pads, pads as large as those on outer fingers; (13) dorsum gray with small black spots on center of back; venter pale gray without markings; no markings in groin or on _ concealed thighs; (14) adults small, males 21.8- 22.2, females 27.1-29.1 mm SVL. Eleutherodactylus balionotus is most easily recognized by its distinctive color pattern. This species is most similar to E. riveti (Despax) from which it differs in width of the digital pads (pads slight- ly wider in riveti), snout length (slightly shorter in riveti), width of upper eyelid (narrower in riveti), and skin texture (warts on dorsal surfaces flatter in riveti, more tuberculate in balionotus). Description.—Head as wide as body, wider than long; head width 37.5-40.8 percent SVL (x = 39.5, N = 10); snout subacuminate in dorsal view, rounded in lateral profile, tip pointed; canthus ros- tralis moderately sharp, concave; loreal region concave, sloping abruptly to lips; lips not flared; snout of moderate length (length of eye slightly less than E-N distance); nostrils directed dorsolateral- ly, weakly protuberant; interorbital space flat, no cranial crests; frontopari- etals complete (no fontanelle); nasals in contact; eyes large, eyelid width 88.2- 110.4 percent IOD (x = 103.2, N = 10); tympanum distinct, round or slightly higher than long, its length 39.6-55.9 percent length of eye (x = 47.8, N = 10); tympanum separated from eye by distance equal to tympanic diameter; supratympanic fold present, thick, con- cealing upper edge of tympanum, warty; tongue slightly longer than wide, not notched posteriorly, posterior one- third not adherent to floor of mouth; choanae and vomerine odontophores small and equal in size; choanae widely separated, completely visible when roof of mouth is viewed from directly above; vomerine odontophores situated well posterior and median to choanae, sepa- rated by twice width of one odonto- phore; each odontophore rounded, ele- vated, bearing a clump of 4-6 teeth; males with vocal slits and a subgular vocal sac. Skin on top of head shagreened, that on eyelids bearing a few large warts (as large as those on back), that on face smooth; skin of body and limbs covered with various-size warts (not flattened); no dorsolateral folds; some females hav- ing a thin sagittal fold; skin of venter coarsely areolate, discoidal folds promi- nent; skin of throat weakly areolate. Forearm areolate but some enlarged tubercles present along ulnar edge (in an indefinite row); two palmar tuber- cles, median larger, both smaller than oval thenar tubercle; numerous super- numerary palmar tubercles, not extend- ing onto digits; subarticular tubercles prominent, round, subconical, simple; fingers bearing discs on pads; pad on thumb round, those on other fingers wid- er than long, not emarginate; dilation LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS ratios I: 1.2-1.7 (x = 1.4), II: 1.4-1.8 (x = 16), III: 1.4-1.8 (x = 16), IV: 1.4- 18 (x = 1.6); fingers bearing lateral fringes (Fig. 4); first finger shorter than second. Hind limbs short, shank 44.8-52.7 per- cent SVL (x 47.0, N 10); heel bearing three or four rounded tubercles (not conical); tarsus bearing a row of similar tubercles along its outer edge and a less definite row along its inner edge; two metatarsal tubercles, both longer than wide, inner two to four times size of outer; numerous, indefinite plantar supernumerary tubercles; sub- articular tubercles round, somewhat flat- tened, simple, smaller than those of fin- gers; toes bearing lateral fringes but no basal webbing; all toes bearing discs on pads which are wider than long. In preservative, the dorsal surfaces are gray, flecked and spotted with black. The flecking is most intense on the back but usually some flecking is found on 9 the limbs. The venter is dull grayish- white with occasional brown or black spots. The tympanum is brown. Can- thal and supratympanic stripes are brown to black and the lips are not barred. The posterior surfaces of the thighs are pale rusty-brown with or without indefinite faint brown marbling. The limbs are not barred. In life, E. balionotus was described as follows: “When collected dark brown; dorsum later changed to pinkish or grayish tan with black flecks; venter dirty white with or without black flecks: iris bronze with median horizontal red streak.” (William E. Duellman field notes, 22 July 1971). Measurements of holotype in mm.— SVL, 28:4, shank 12:8, head width 11.5, head length 10.7, upper eyelid width 3.1, IOD 2.8, tympanum length 1.7, eye length 3.3, eye-nostril 3.6. The holotype has heavily convoluted oviducts and large yellow ovarian eggs. Fic. 4—(A) Eleutherodactylus balionotus, KU 142136; (B) E. riveti, KU 131172. Lines equal 5 mm. 10 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Etymology—Greek, meaning speck- led back, in reference to the predomi- nant color pattern. Natural history—E. balionotus is known only from the type-series which was collected by day from terrestrial bromeliads in the subparamo habitats atop the windswept ridge east of Ciu- dad Loja. The type-series consists of two males, one immature female (KU 142143), and seven gravid females with strongly convoluted oviducts and large yellow eggs. Remarks.—The lateral fringes on the digits, large inner metatarsal tubercle, and large nasal bones (in median con- tact) suggest that E. balionotus is allied to the squat-bodied species found in central and northern Andean Ecuador (E. orcesi, E. riveti, E. thymelensis, and E. unistrigatus). Eleutherodactylus rui- dus differs markedly in lacking the mid- dle ear and vocal apparatus; in propor- tions, skin texture, and most features of the hand and foot it resembles E. balio- notus and E. riveti. Eleutherodactylus balionotus is most like E. riveti; more fieldwork on the isolated and semi-iso- lated mountain ridges of southern An- dean Ecuador (Cordilleras de Chilla and Condorcillo) is needed to establish whether E. balionotus is a distinct spe- cies from E. riveti or a geographic variant. Eleutherodactylus baryecuus new species Fig. 5 Holotype -—USNM 199714, an adult female, obtained at Suro Rancho, Pro- vincia Morona-Santiago, Ecuador, 2683 m, by James A. Peters on 24 August 1962. Paratypes.—All Provincia Morona- Santiago: USNM 199715, Cerro Negro, 2927 m; USNM 199717-22, between San Juan Bosco and El] Cruzado, 2226 m; USNM 199723-24, El] Cruzado, 2195 m; USNM 199725, San Juan Bosco, 2195 m; USNM_ 199726-28, Sapote, 2470 m; USNM 199729, San Vicente, 2805-2835 m; USNM 199730, 3 km W San Vicente, 2988 m; USNM 199716, between Sevilla de Oro and Méndez (probably between the crest and Pailas). Diagnosis—(1) skin of dorsum smooth with scattered, low warts, venter finely areolate; (2) tympanum absent; (3) snout round in dorsal view, truncate in lateral profile, short; (4) interorbital space as wide as upper eyelid in males, broader in females; low cranial crests present; (5) vomerine odontophores elevated, triangular in outline; (6) males lacking vocal sac and slits; (7) first finger shorter than second; all digits bearing broad discs on dilated pads; distal subarticular tubercles grooved; (8) fingers lacking lateral fringes; (9) ulnar tubercles indistinct except ante- brachial; (10) non-conical tubercles on heel, none on tarsus; (11) two metatar- sal tubercles, inner oval, non-com- pressed, 1.5-2 times size of subconical outer; no supernumerary plantar tuber- cles; (12) toes bearing lateral fringes, broad discs on dilated pads; toe pads as large as those of fingers; (13) brown a Ng pees Fic. 5.—Eleutherodactylus baryecuus, USNM 199727; line equals 10 mm. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS i above with darker brown spots, limb bars narrow, oblique; venter cream re- ticulated with brown; (14) adults mod- erate-size, males 27.2-30.4 mm, females 38.2-43.5 mm SVL. Eleutherodactylus baryecuus is a member of the wnistrigatus group most similar to E. surdus (Boulenger) from the Pacific slopes of Andean Ecuador. Both lack ears, have relatively smooth skin of the dorsum, moderate-sized digi- tal pads, and low cranial crests. The two frogs are identical in size (6 ¢ sur- dus have a mean SVL of 28.0 + 1.9 mm and 16 9 have a mean SVL of 40.5 + 1.3 mm). Eleutherodactylus surdus does not have so rounded a snout as does E. baryecuus in dorsal view [compare Fig. 5 with Boulenger’s (1882) illustration of surdus], has larger eyes (and hence a lower upper eyelid width/IOD ratio, see Lynch, 1970a: 140), and longer legs than E. baryecuus; E. surdus also has poorly developed, but evident, dorso- lateral folds. Description—Head narrower than body, wider than long; head width 35.1- 40.8 percent SVL (x = 36.9, N = 13); snout rounded in dorsal view, angularly rounded to truncate in lateral profile, short; E-N 64.7-77.4 percent eye length in males (x = 70.0, N = 5), 67.3-90.0 percent in females (x = 78.5, N = 8); nostrils not protuberant, directed dorso- laterally; canthus rostralis moderately sharp, weakly concave; loreal region concave, sloping gradually to lips, lips not flared; upper eyelid width 92.8-113.0 percent IOD in males (x = 104.6, N = 5), 70.0-97.1 percent in females (x = 86.7, N = 8); edges of frontoparietals upturned forming low cranial crests; up- per eyelid bearing flattened tubercles; supratympanic fold thick; tympanic an- nulus, cavum tympanicum absent; pos- trictal tubercles prominent; choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odon- tophores median and posterior to choa- nae, elevated, triangular in outline, each 3-4 times as wide as a choana, separated by a choanal width, bearing a transverse row of 4-6 teeth along posterior edge; tongue as broad as long, posterior % not adherent to floor of mouth, posterior border not notched; males lacking vocal sac and slits. Skin of dorsum smooth but bearing low flat warts, those on eyelids and be- hind eye pungent; no dorsolateral folds evident in adult females, males have weakly evident glandular dorsolateral folds; anal opening extended onto back of thighs by sheath; venter finely areo- late; discoidal folds obscure; 1-2 ulnar tubercles evident on distal quarter of forearm; palmar tubercle bifid, much larger than oval thenar tubercle; super- numerary palmar tubercles small, round; subarticular tubercles larger than super- numeraries, round to slightly broader than long; distal subarticular tubercles on fingers III and IV grooved (in some males, nearly bifid); fingers lacking lat- eral fringes; all fingers bearing discs (broader than long) on __ apically rounded, dilated pads; pads on thumb smallest; males with swollen, non-spi- nous nuptial pad on thumb; thumb shorter than second finger. Heel bearing non-conical tubercles; no tubercles on knee or tarsus; inner metatarsal tubercle twice as long as wide, non-compressed; outer metatarsal tubercle }-% size of inner, low, subconi- cal, round; no supernumerary plantar tubercles; basal subarticular tubercles longer than wide, distal subarticular tubercles round on toes I, II, V, grooved on IIT and IV; toes bearing narrow lat- eral fringes but not webbed; all toes bearing discs on dilated pads; toe pads as large as those of outer fingers; when legs are flexed at right angles to body, heels just overlap; heel of adpressed hind limb reaches area just posterior to eye; shank 43.4-48.9 percent SVL (x = 45.8, N = 13). In preservative, brown above with darker brown markings (viz., canthal- supratympanic stripes, labial bars, inter- orbital bar, spots on back sometimes co- alescing forming scapular W,_ sacral chevron); flanks tinged with yellow and 12 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY spotted with dark brown; anal triangle faint; brown bar across thigh coalescing on posterior thigh forming brown back- ground which may be spotted with yel- low; shank bars, if complete, oblique, narrower than interspaces; venter cream, spotted and reticulated with brown; spotting on venter most prominent in large females; males and small females have cream venters with faint, if any, spotting. In life, E. baryecuus is olive to brown above with black markings. The venter is gray with dull yellow spots. The throat is dull yellow with some gray re- ticulation. Between the bands on the thigh and to a lesser extent on the pos- terior flanks the ground color is stippled with yellow. In some individuals a yel- low wash is seen adjacent to the black markings. Measurements of holotype in mm.— SVL 40.2; tibia 19.5; head width 14.1; head length 12.3; upper eyelid width 3.1; IOD 3.5; eye length 4.1; E-N 3.2. The holotype is an adult female with heavily convoluted oviducts and large yellow ovarian eggs. Etymology—Greek, baryecoos, meaning hard of hearing; in reference to the loss of the middle ear in this frog. Natural history—Most of the speci- mens were found beneath logs and rocks; James Peters found one clasping pair (USNM_ 199718-19). The eight adult females contain large ovarian eggs, and the five adult males have swollen thumbs and large testes. These obser- vations as well as the capture of an am- plectant pair suggest the species was ready to breed when collected in August 1962. Remarks.—I once (Lynch, 1970a) included E. chloronotus, E. devillei (Boulenger), and E. surdus (Boulen- ger) in the surdus group within what I then referred to as the unistrigatus com- plex. With the recognition of the wni- strigatus group as a_ species group (Lynch, 1976a), I now view my earlier surdus group as an Artenkreis but am no longer convinced that E. chloronotus is a member. I do include, as vicariants, E. baryecuus, E. devillei, and E. surdus. The differences between the three are adequate to insist on species level rec- ognition for each and in spite of the re- tention of a complete middle ear in E. devillei and the loss of the ear in E. bar- yecuus and E. surdus, I do not view the latter two as more closely related to one another than either is to E. devillei. The three species are separated by formid- able barriers (the valley of the Rio Pas- taza and the interandean valley and paramos of the Cordillera Occidental). All three species occur in forested habi- tats near tree-line. At present I am aware of no members of this Artenkreis among the faunas of Colombia or Pert. Distribution—Known only from the eastern face of the Cordillera de Oriental east of Cuenca at elevations of 2195- 2988 m. Eleutherodactylus bromeliaceus new species Fig. 6 Holotype —USNM 199731, an adult female, obtained in bromeliads between Sapote and Suro Rancho, Provincia Morona-Santiago, Ecuador, 2622 m, by Manuel Olalla on 24 August 1962. Paratypes——USNM_ 199732-34, Pai- las, 2195 m; USNM 199735, Mirador, 1982 m; USNM 199736-37, Plan de Mila- gro, 1707 m; USNM 199738-39, El Cru- zado, 2195 m; USNM 199740, % km E Sapote, 2393 m; USNM 199742, moun- tain above Sapote to south, 2500 m; USNM 199741, 1 km E Sapote, 2332 m; all in Provincia Morona-Santiago, Ecua- dor. Diagnosis —(1) skin of dorsum smooth (eyelid bearing tubercles), that of venter coarsely areolate; no dorso- lateral folds; (2) tympanum visible, round, its length 4-% that of eye; (3) snout subacuminate in dorsal view (tip pointed), pointed in lateral profile; (4) interorbital space broad, no cranial crests; (5) vomerine odontophores ob- lique, prominent; (6) males with vocal slits and external subgular vocal sac; LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 13 (7) first finger shorter than second; all digits bearing broad discs on dilated pads; numerous supernumerary palmar tubercles; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) heel and outer edge of tarsus bearing low tubercles; inner tarsal tubercle pres- ent; (11) two metatarsal tubercles, in- ner oval, three times size of outer; nu- merous supernumerary plantar tuber- cles; (12) toes bearing lateral fringes; pads and discs of toes slightly smaller than those of fingers; (13) tan above be- coming brown on flanks; dorsum flecked or blotched with brown; venter cream (flecked with brown in adult females); limb bars transverse, narrower than in- terspaces; concealed thigh brown with diffuse yellow spots; (14) adults small, males 16.7-23.2 mm, females 22.9-28.1 mm SVL. Eleutherodactylus bromeliaceus _ is most similar to E. lacrimosus (Jiménez de la Espada) and perhaps E. chalceus (Peters). The latter purportedly lacks vomerine odontophores; E. lacrimosus has a more rounded snout (see illustra- tion in Lynch and Schwartz, 1972) and is a smaller frog lacking supernumerary palmar and plantar tubercles, lateral fringes on the digits, and tubercles on the eyelid and tarsus. Description—Head as_ broad as body, wider than long; head width 34.4- 39.8 percent SVL (x = 37.1, N = 12); snout subacuminate in dorsal view, tip pointed, pointed in lateral profile (acute- ly rounded); snout relatively long, E-N 75.0-97.0 percent eye length (x = 84.4, N = 12); nostrils weakly protuberant, directed anterodorsolaterally; canthus rostralis obtuse; loreal region weakly concave, sloping rapidly to lips; lips not flared; interorbital space broad, flat, no cranial crests; upper eyelid width 66.7- 95.6 percent IOD (x = 80.3, N = 12); upper eyelid bearing 2-3 non-conical tubercles; temporal region sloping (tym- panum directed dorsolaterally); supra- tympanic fold thin, distinct, concealing upper edge of tympanic annulus; tym- panum prominent, round, its length 28.6- 42.3 percent eye length (x = 38.0, N = 12), separated from eye by % tympanic length; postrictal tubercles not promi- nent; choanae moderate-sized, oval, not concealed by palatal shelf of maxillary arch; vomerine odontophores median and posterior to choanae, oblique, each % size of a choana, bearing a clump of 4-5 teeth, separated by 1% choanal widths; tongue much longer than wide, posterior §& not adherent to floor of mouth, posterior edge not notched; males with vocal slits and large external subgular vocal sac. Skin of dorsum (except eyelid) smooth, no dorsolateral folds, that of Fic. 6.—Eleutherodactylus bromeliaceus, USNM 199731; line equals 5 mm. 14 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY flanks areolate, that of venter coarsely areolate; discoidal folds prominent; no anal sheath; limbs finely areolate; ulnar tubercles not evident; palmar tubercle bifid, twice as large as oval thenar tu- bercle; many supernumerary palmar tubercles; subarticular tubercles round, non-conical, elevated; fingers bearing prominent fringes, discs (broader than long), and dilated pads; pad on thumb smallest, those on fingers III and IV as large as tympanum; first finger shorter than second. Knee and heel bearing small warts; tarsus bearing row of non-conical tuber- cles along outer edge, single tubercle along inner edge; inner metatarsal tuber- cle twice as long as wide, non-com- pressed; outer metatarsal tubercle elon- gate, non-conical, % size of inner; many supernumerary plantar tubercles; sub- articular tubercles round, non-conical; toes bearing lateral fringes; toes bearing broad discs on dilated pads (slightly smaller than those of outer fingers); heels of flexed legs overlap, heel of ad- pressed hind limb reaches eye; shank 47.4-52.6 percent SVL (x = 50.0, N = PANS In preservative, males are pale cream-tan with few brown spots on the head, back, and lower limbs; adult fe- males are tan above becoming brown on the flanks, dorsum bearing brown chev- ron, interorbital bar, suprainguinal spots, and marbling; no canthal stripe; labial bars, supratympanic stripe brown; ven- ter cream in males, cream heavily flecked with brown in females; posterior surface of thigh brown with diffuse cream spots. In life, E. bromeliaceus is pale green to olive above with brown to black markings. The venter is pale bronze- yellow and the throat dull yellow. The iris is brown flecked with gold or bronze. One adult female (USNM 199740) had a dull orange ground color; the dark markings resulted in a large orange blotch on the center of the back. Measurements of the holotype in mm.—SVL 27.2; tibia 13.5; head width 9.9; head length 9.3; upper eyelid width 2.1; IOD 3.1; tympanum length 1.3; eye length 3.1; E-N 3.0. The holotype is an adult female with extensively con- voluted oviducts and large yellow ovar- ian eggs. Etymology.—Latin, meaning living in bromeliads; most of the specimens se- cured by Peters and his associates were collected in bromeliads on trees. Natural history—Some__ specimens were collected at night on leaves of ele- phant ear plants but most specimens were collected during the day in the axils of bromeliads taken from trees. Two specimens were collected by day in the axils of elephant ear plants. Many of the males have distended vocal sacs suggesting that they may have been call- ing. All five adult females contain large Ovarian eggs. Remarks.—In addition to the low- land E. lacrimosus ranging from Colom- bia and Ecuador east to Belém in Brasil and E. bromeliaceus from the Amazon- ian versant of the Cordillera de Matanga in southern Ecuador, I am aware of two allied species in Ecuador (one of the Amazonian Andean versant of northern Ecuador and another in the valley of the Rio Pastaza). The species in the Pas- taza valley is possibly E. chalceus (Pe- ters). Another allied apparently unde- scribed species is found in the Andean foothills in Bolivia. Distribution—The Amazonian ver- sant of the Cordillera de Matanga, in Provincia Morona-Santiago, Ecuador, at elevations of 1707-2622 m. Eleutherodactylus cajamarcensis Barbour and Noble Fig. 3C Lynch (1969a) presented a_ rede- scription of this species based largely on series collected between Loja and Sara- guro, Ecuador; he also identified some specimens from central Andean Ecuador as possibly conspecific. At that time I anticipated that the specimens from Provincia Tungurahua would prove to be a distinct subspecies and that E. ca- jamarcensis would be collected in the LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 15 intervening 200 km of interandean Ec- uador. The Gustavo Orces-V and James A. Peters collections at the National Mu- seum of Natural History and the con- tinued field work of William E. Duell- man and his associates at the University of Kansas yielded more specimens of each population but none from the in- tervening area. The northern popula- tions represent a related but distinct spe- cies of frog having closer relationships with a complex of species including E. lehmanni (Boettger) and E. unistrigatus (Giinther). Eleutherodactylus cajamarcensis re- mains known from the Loja basin and transandean Pert at elevations between 1870 and 3000 m. The additional speci- mens examined do not appreciably en- large the known distribution given by Lynch (1969a). Eleutherodactylus colodactylus new species Fig. 3D Holotype —KU 142151, an adult fe- male from 13.5 km E Loja, at the crest (Abra de Zamora) on the frontier be- tween Loja and Zamora-Chinchipe provinces, Ecuador, 2800 m, obtained on 22 July 1971 by William E. Duellman. Paratunes.—KU 149159.59— tano- types; KU 142160-61, 14 km E Loja, Provincia Zamvura-Coincuipe, Ziéu at; KU 142162-64, 15 km E Loja, Provincia Zamora-Chinchipe, 2710 m. Diagnosis.—(1) skin of dorsum and venter areolate; (2) tympanic annulus, cavum tympanicum, and columella ab- sent; (3) snout rounded or obtuse in dorsal view, rounded or truncate in lat- eral profile; E-N greater than eye length; (4) interorbital space flat, no cranial crests; upper eyelid width one- half to three-fourths IOD; (5) vomerine teeth and odontophores present, con- cealed beneath tissue of palate; (6) male lacking vocal sac and slits; (7) first finger shorter than second, all digits short; distal subarticular tubercles tend to be bifid; all digits bearing discs on pads: dilation’ ratios 1: 11, Tle 12.1: 1.4, IV: 1.4; (8) fingers bearing broad lateral fringes (reducing dilation ratio values ); (9) no ulnar tubercles; (10) no tarsal tubercles; prominent tubercle on heel (rounded); (11) two metatarsal tu- bercles, outer rounded, one-fifth to one- fourth size of oval (length twice width) inner; plantar surface areolate, numer- ous supernumerary tubercles continue onto toes; (12) toes bearing prominent lateral fringes, basal webbing, discs and pads; toe pads as large as those of fin- gers; (13) dorsum brown (rarely gray ) with pale dorsolateral stripes or pale in- terorbital bar; dark brown canthal and postocular stripes present; limbs not barred; venter cream with varying in- tensity of brown peppering; (14) adults minute, 6 6 14:0:20'7, 9-0. 16:5295:8 mm SVL. Eleutherodactylus colodactylus _ is readily distinguished from all other eleu- therodactyline frogs except E. anotis, E. baryecuus, E. pugnax, E. ruidus, and E. surdus by virtue of having discs on digital pads (and T-shaped terminal phalanges) and lacking the tympanic annulus, cavum tympanicum, and _ col- umella. Eleutherodactylus colodactylus has short, stubby fingers (Fig. 7) and toes and does not have readily visible vomerine odontophores in contrast to E. anotis, E. baryecuus, E. pugnax, E. rui- dus, and E. surdus all of which are con- siderably larger frogs (adult females measure—E.. anotis 9° 2 39.0-47.0 mm; E. pugnax, 1 @, 30.8 mm; E. surdus 2 2 35.5-43.8 mm). Eleutherodactylus colodactylus is included in Cochran and Goin’s (1970) group II but has no close relatives among the described species of that group. Description—Head as broad as body; head as broad as long; head width 33.1-40.2 percent SVL (x = 36.0, N = 14); snout obtuse in dorsal view, rounded or truncate in lateral profile; canthus rostralis rounded; loreal region flat, sloping gradually to lip; lips not flared; nostrils weakly protuberant, di- rected dorsolaterally; snout short, E-N 16 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY slightly greater than eye length; eyes small; width of upper eyelid 56.3-71.8 percent IOD (x = 63.3, N = 14); inter- orbital region flat, no cranial crests; no frontoparietal fontanelle; tympanum ab- sent; tongue large, round, not notched posteriorly, posterior two-fifths free; choanae moderate-sized, round, com- pletely visible when roof of mouth is viewed from directly above, separated by a distance equal to four times width of a choana; no vomerine dentigerous processes, vomerine teeth present (2-4) in a clump beneath skin of palate; male lacking vocal sac and vocal slits. Skin of body finely areolate; no dor- solateral folds or discoidal folds; upper eyelid bearing one or two small but prominent tubercles; skin of posterior surfaces of thigh and concealed shank smooth; areolation less pronounced on dorsal surfaces; shank 42.0-47.8 percent SVL (x = 44.1, N = 14); areolation on forearm not enlarged to form ulnar tu- bercles; two palmar tubercles, median largest, larger than oval thenar tubercle, outer palmar tubercle not always distin- guishable from supernumerary thenar tubercles; palmar surface areolate; su- pernumerary tubercles present on ven- tral surfaces of digits; subarticular tu- bercles broader than long, non-conical, tending to bifurcate distally; digits short, broad, flat; edges of digits coa- lesce proximally to form thick, basal web (Fig. 7); all digits bear pads, pads broader than long; dilation ratios I: 0.9- 1.2 (x = 11), Ul: 1.1-L3)Ga=aeae III: 1.3-1.5 (x = 14), IV: 13-16 (x = 1.4); first finger shorter than second. Fic. 7.—Hands of Eleutherodactylus having narrow digital pads. (A) E. colodactylus, KU 142154; (B) E. orestes, KU 141998; (C) E. vidua, KU 120089; line equals 5 mm. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS ey, Heel bearing moderate-sized conical tubercle; inner and outer edges of tarsus lacking tubercles or folds; two metatar- sal tubercles, not compressed, non-coni- cal, inner larger than outer, inner one and one-half to two times as long as wide, four to five times as large as rounded outer metatarsal tubercle; plan- tar surface areolate, supernumerary tu- bercles continue onto toes; subarticular tubercles bifid or wider than long and tending to be bifid; toes short, broad, and flat, lateral edges coalescing to form basal web; all toes bear pads, pads wider than long. In preservative, the ground color is usually brown (pale to reddish brown) although some individuals have a gray ground color. The venter is speckled with brown and thus appears dark; in- dividuals with pale dorsal coloration ap- pear dirty cream ventrally. The limbs are not barred and the posterior surface of the thigh is uniform brown. All in- dividuals have a dark brown canthal stripe and postocular stripe. Five indi- viduals have silvery or cream flecking in a line above the canthal stripe, across the upper eyelid, and to above the groin as a dorsolateral stripe. In five speci- mens, the dorsum is flecked with silver or white; the flecks are edged in brown. In life, E. colodactylus was described as “dorsum greenish tan; venter pale yellow; iris reddish bronze” (William E. Duellman field notes 21 July 1971); “dorsum varying from brown to red to tan with or without orange or tan dor- solateral stripes; venter gray; iris red- dish-bronze” (W. E. Duellman field notes 22 July 1971). Measurements of the holotype in mm.—SVL, 20.7, shank 8.7, head width 7.9, eyelid width 1.8, interorbital dis- tance 2.8. The holotype is an adult fe- male with convoluted oviducts and yel- low ovarian eggs (2.0-2.3 mm in diam- eter). Etymology.—Greek, kolos + dakty- los, meaning stunted toes, in reference to the stubby fingers and toes. Natural history—All specimens for which data are available were collected in bromeliads by day. Those from the Abra de Zamora and those from Depto. Piura (Pert) were collected in terrestri- al bromeliads (see Duellman, 1974, for a habitat photograph). James A. Peters and his associates collected many (most ?) in arboreal bromeliads at low- er elevations (2200-2550 m). The north- ern-most samples (eastern slopes of Cer- ro El Picacho, Paramos de Matanga) were taken at lower elevations than the more southern samples and_ contain smaller frogs (Table 1). There appears to be a cline in size among males; fe- males from the three areas and altitudes do not differ significantly in size. Remarks.—The specimens from the vicinity of the type-locality exhibit only two pattern morphs (dorsolateral stripes vs a pattern lacking such stripes). Five of 14 specimens have dorsolateral stripes (35.7 percent). Three morphs are evi- dent among the specimens from EI Pi- cacho. The dorsolateral stripe morph occurs in 4 of 27 specimens (14.8 per- cent). An equally abundant morph has a brown hour-glass on the dorsum. The most common morph is gray or brown with or without brown spots and/or cream flecks and may exhibit darker flanks than the center of the back; this TABLE 1. Size of Eleutherodactylus colodactylus. Locality Altitude Males Females E] Picacho 2200-2550 m IS) S= (O48) (CUS) 20.4 + 1.4 (18) UE 2X0 16.5 — 25.8 Abra de Zamora 2850 m 17.0 + 0.6 (8) 20.6 + 0.3 (6) UGG) — IS, 201 — 21-9 Piura, Pert 2745-3100 m 18.3 + 0.4 (19) DE =t=5 01969) 16.8 — 19.6 19.1 — 23.2 1 mean + 2 standard errors (N) 2 Range 18 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY morph accounts for 70.4 percent of the specimens and is not more abundant in the El] Picacho population than in the populations from the type-locality (64.3 percent). Specimens from Depto. Piura, Peru, exhibit continuous variation in color pattern from having a few dark flecks to having a well-defined hour- glass pattern. The absence of the middle ear is rare among Eleutherodactylus—I_ am aware of only five other earless species —E. anotis Walker and Test ( Venezue- la), E. baryecuus Lynch, E. pugnax Lynch, E. ruidus Lynch, and E. surdus (Boulenger) (all Ecuadorian); all are members of the wunistrigatus group as currently defined (Lynch 1976a). In these five species, unlike E. colodacty- lus, the fingers and toes are compara- tively long and slender and the vome- rine odontophores are prominent struc- tures on the palate. The bifid (or near bifid) distal subarticular tubercles of colodactylus are unlike the “normal” subarticular tubercles found in the other five earless species. Bifid distal subartic- ular tubercles occur in E. areolatus, a small species found on the Pacific low- lands in Colombia and Ecuador, but the similarity is not convincing evidence of relationship (E. areolatus is a close rela- tive of E. gularis, a member of the di- astema group). In dorsal view, the skull of E. colo- dactylus is not especially noteworthy; the nasals are comparatively large but not in median contact and are separated from the frontoparietals (Fig. 8). The sphenethmoid is large and extends well anteriad beneath the nasals. The fron- toparietals are complete and appear to be fused to the prootics. In ventral view, the vomers are small but not great- ly reduced in size. The palatines are short (compare with Fig. 11). The me- dian rami of the pterygoids are short and are widely separated from the para- sphenoid alae. The alae of the para- sphenoid are weakly deflected posterior- ly. An unusual condition is the greatly elongate anterior ramus of the parasphe- noid, extending anteriorly to a point anterior to the palatines and between the dentigerous rami of the vomers. This condition is similar to that seen in some of the species of the Phrynopus flavomaculatus group (Lynch, 1975) but E. colodactylus differs markedly in hav- ing very short palatines (as well as in digital morphology). The skull of E. colodactylus exhibits the character-states of the Alpha group of Eleutherodacty- lus. Distribution —E. colodactylus _ is known from the high Amazonian slopes of the Andes in southern Ecuador be- tween 2200 and 2850 m and from the Fic. 8.—Dorsal (A) and ventral (B) views of skull of Eleutherodactylus colodactylus, KU 142155. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 19 crest and Pacific versant of the Cordil- lera between Chanchaque and Huanca- bamba, Depto. Piura, Peru, at elevations of 2745-3100 m. Eleutherodactylus cryophilius new species Fig. 9A Holotype —USNM 199993, an adult male, obtained 6 km W San Vicente, Provincia Morona-Santiago, Ecuador, 3110 m, on 27 August 1962 by Robert Mullen, Manuel Olalla, James A. Peters, and Peter Spoecker. Paratypes—USNM 200391-93, topo- types; USNM 200390, crest between Se- villa de Oro and Méndez, Provincia Mo- rona-Santiago, 3384 m; KU 120091, La- guna de Zurucuchu, Provincia Azuay, 3200 m; AMNH_ 13970. Rest'4n on southwestern slopes Cerro Bestién, 3079 m; all from Ecuador. Diagnosis—(1) skin of dorsum bearing flattened warts; dorsolateral folds present, indistinct; skin of venter coarsely areolate; (2) tympanum con- cealed beneath skin; (3) snout subacu- minate in dorsal view, pointed in lateral profile; (4) interorbital space broader than upper eyelid; cranial crests promi- nent; (5) vomerine odontophores. tri- angular in outline, prominent; (6) males lacking vocal sac and slits; (7) first fin- ger shorter than second; all digits bear- ing discs on narrowly dilated pads; (8) fingers bearing indefinite lateral fringes; (9) no ulnar tubercles; (10) heel and tarsus lacking tubercles; (11) two meta- tarsal tubercles, outer % size of oval, non- compressed inner; supernumerary plan- tar tubercles indistinct; (12) toes keeled laterally, no lateral fringes; toes bearing discs on narrow pads; (13) brown with darker brown reticulation above; venter pale brown; concealed limb surfaces brown, sometimes with cream spots; up- per lip bears cream stripe; (14) adults relatively large, ¢ ¢ 28.9-36.8 mm, 2 9 43.1-49.8 mm SVL. Eleutherodactylus cryophilius is most similar to E. buckleyi (Boulenger) and E. curtipes (Boulenger); it differs from buckleyi in not having visible tympana and having a dark venter instead of a white to cream venter with brown retic- ulation. Eleutherodactylus cryophilius resembles E. curtipes in having the tym- pana concealed but differs in that the tympanic annulus is reduced in size compared to that of E. curtipes. Eleu- therodactylus curtipes has lateral fringes on the toes whereas E. cryophilius does not. Although there is considerable geo- graphic variation in size and ventral pig- mentation in E. curtipes, no population reaches the body size of E. cryophilius. The largest E. curtipes I have seen in- clude a male 32.55 mm (USNM JAP 5883, 10 km W Bafios, Provincia Tungu- rahua) and a female 42.9 mm (KU 130867, Volcan Pichincha, Provincia Pi- chincha). The populations (with range, mean and + 2 standard errors) from which these extremes are drawn have the following sizes—Volcan Pichincha: 12 ¢ 6, range 21.6-30.6 mm SVL (x = 26.3 mm + 16mm), 8 ¢ 9, range 35.3- 42.9 mm SVL (x = 40.4 mm + 15 mm); 7-10 km W Banos: 26 ¢ ¢ 20.6- 32.5 mm SVL (x = 266 mm + 1.0 mm), 15 @ 9, range 32.4-41.8 mm SVL (x = 37.4 mm + 1.4 mm). Eleuthero- dactylus buckleyi is about the same size as E. cryophilius [E. buckleyi—31 ¢ @ 24.5-38.7 mm SVL (x = 33.1 mm + L7 mm); 10 ¢@ ¢ 37.1-48.8 mm SVL (x = 45.3 mm + 2.4 mm)]. Description—Head narrower than body, wider than long, its width 34.6- 37.5 percent SVL (x = 33.6, N = 9); snout subacuminate in dorsal view, pointed in lateral profile; snout short, E-N 68.2-87.2 percent eye length (x = 80.0, N = 9); nostrils not or weakly pro- tuberant, directed laterally; canthus ros- tralis sharp, straight; loreal region weak- ly concave, sloping abruptly to lips; lips not flared; interorbital space deeply fur- rowed, furrow extends onto snout (edges of nasals higher than midline; edges of frontoparietals upturned); upper eyelid width 72.5-95.4 percent IOD (x = 81.6, N = 6); upper eyelid lacking tubercles; 20 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 9.—(A) Eleutherodactylus cryophilius, KU 120097; (B) E. cryptomelas, KU 120096; (C) E. lymani, KU 119503; (D) E. orestes, KU 120094. temporal region swollen, tympanic re- gion vertical; supratympanic fold prom- inent; tympanic annulus small, reduced in size, concealed beneath skin; choanae small, round, not concealed by palatal shelf of maxillary arch; vomerine odon- tophores large, median and posterior to choanae, triangular in outline, bearing a transverse row of 4-5 teeth along pos- terior margin, each 4-5 times width of a choana, separated by distance equal choanal width; tongue longer than wide, posterior #% not adherent to floor of mouth, posterior edge feebly notched; males lacking vocal sac and slits. Skin of dorsum and flanks bearing low, flat warts; indefinite dorsolateral folds present; skin of venter coarsely areolate; discoidal folds indistinct; anal opening not extended in sheath; no ul- nar tubercles or fold; palmar tubercle bifid, larger than oval thenar tubercle: supernumerary palmar tubercles pres- ent, non-conical, slightly smaller than subarticular tubercles which are round, elevated, simple; fingers bearing indefi- nite lateral fringes; all fingers bearing discs (broader than long) on pads; pad of thumb not wider than digit below pad, those on fingers II-IV_ slightly wider than digit below pad; first finger shorter than second; base of thumb swollen in males. Knee, heel, and tarsus lacking tuber- cles; inner metatarsal tubercle 2% times as long as wide, not compressed, twice as large as round, non-elevated outer metatarsal tubercle; supernumerary plantar tubercles obscure, low, flattened, smaller than subarticular tubercles which are round to slightly longer than wide, non-conical, elevated, simple; toes not bearing lateral fringes (edges of toes weakly keeled); all toes bearing discs (wider than long) on pads; pads slightly broader than width of digit be- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 21 low pad; heels of flexed legs touch; heel of adpressed hind limb reaches eye; hind legs short, shank 42.4-50.3 percent SVL (x = 46.8, N = 9). Brown with darker brown reticula- tion on dorsum, limbs, and flanks; venter paler than dorsum; groin, concealed limb surfaces brown with or without cream spots; canthal and supratympanic stripes indefinite; upper lip bears cream stripe. In life, E. cryophilius is medium brown with a network of darker brown on the body and limbs; some yellow flecking on flanks; posterior surface of thigh and venter rose-brown, latter flecked with cream. The iris is golden yellow flecked with brown. Measurements of holotype in mm.— SVL 36.8; tibia 17.3; head width 13.2; head length 11.7; upper eyelid width 2.9; IOD 4.0; eye length 4.4; E-N 3.0. The holotype is an adult male with swollen thumbs and large white testes. Etymology.—Greek, kryos, cold and philios, loving; in reference to a habitat preference for the general climate where the species is found. Natural history—E. cryophilius was found beneath rocks and logs in sub- paramo and pdramo habitats on the mountain ranges surrounding the Cuen- ca hoya. All five adult females are ma- ture and contain large ovarian eggs. The five males have swollen thumbs and large testes. Remarks.—All available evidence points to E. cryophilius as the southern vicariant of the E. curtipes Artenkreis. The known distribution of E. cryophil- ius is small compared to those of E. buckleyi and E. curtipes but I am con- fident E. cryophilius does not range fur- ther south. I likewise doubt that E. cryophilius and E. curtipes will prove to be sympatric. The distribution of E. curtipes is totally within Ecuador from the Colombian border south along both Cordilleras to the Palmira desert at the southern end of the Riobamba hoya. Eleutherodactylus buckleyi is found from northern Ecuador to Cauca and Valle departamentos of Colombia. The vocal apparatus progressively declines and the ear is progressively reduced as one proceeds southward through the series of three species: E. buckleyi (vo- cal slits present, vocal sac subgular; tym- panum visible externally )—E. curtipes (vocal sac and slits absent; tympanum concealed, annulus large)—E. cryophil- ius (vocal sac and slits absent; tympa- num concealed, annulus reduced in size). Distribution.—Known from subpdra- mo and pdramo habitats east and west of Cuenca at elevations of 2835-3384 m. Eleutherodactylus cryptomelas new species Fig. 9B Holotype.—KU 141992, an immature female, taken 15 km E Loja, Provincia Zamora-Chinchipe, Ecuador, 2710 m, May 1971 by William E. Duellman. Paratypes—KU _ 141993, taken syn- topically with holotype; KU 120095-96, 8-9 km N San Lucas, Provincia Loja, Ecuador, 3000 m; USNM 198480-82, Sa- pote, Provincia Morona-Santiago, Ecua- dor, 2470 m; USNM 198483, 2 km W Sapote, Provincia Morona-Santiago, Ecuador, 2560 m. Diagnosis —(1) dorsum shagreened and bearing conical warts and _ ridges, skin of venter areolate; no dorsolateral folds; (2) tympanum visible, round, one-third to two-fifths eye length; (3) snout subacuminate in dorsal view, tip pointed, rounded in lateral profile; (4) no cranial crests, interorbital space flat, as wide as or slightly wider than upper eyelid; (5) vomerine teeth and odonto- phores present, odontophores round to oval; (6) males lacking vocal sac and slits; (7) first finger shorter than second; all digits bearing discs on dilated pads; mean dilation ratios I: 1.3; II: 2.1; III: 2.6; IV: 2.5; (8) fingers bearing weakly defined lateral fringes; (9) ulnar tuber- cles prominent; no enlarged tubercle on elbow; (10) inner edge of tarsus bear- ing indistinct tubercle, outer edge of 22 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY tarsus bearing a series of conical tuber- cles continuing onto heel; (11) two metatarsal tubercles, outer conical, one- sixth to one-fourth size of oval inner; numerous supernumerary plantar tuber- cles; (12) toes fringed, no basal web- bing; digits bearing discs on pads, toe pads smaller than those of fingers; (13) dorsum gray to brown sparsely marked with brown; limb bars oblique; venter white to cream reticulated with brown; anterior and posterior thigh surfaces, posterior lower flank, and concealed shank dark black (in life and in pre- servative); (14) adults of moderate size, 4 males 28.2-30.2 mm, one adult female 38.6 mm SVL. Eleutherodactylus cryptomelas is a long-legged species with broad digital pads and unlike the other high Andean Eleutherodactylus (short limbs, stocky habitus, comparatively narrow digital pads). The black areas on the con- cealed thigh and groin together with the tuberculate skin texture (with eyelid tubercles, ulnar tubercles, and heel and outer tarsal tubercles) recall the condi- tions seen in E. crucifer (Boulenger ) from intermediate elevations on the Pa- cific slopes of Ecuador. Eleutherodac- tylus crucifer is green in life with red eyes and upon preservation, loses the black areas in the groin and on the con- cealed thigh surfaces. The eyelid tuber- cles and the heel and tarsal tubercles of E. crucifer are considerably more promi- nent than are those of E. cryptomelas. Description—Head as wide as or slightly wider than body, head as long as or slightly longer than wide in males, slightly wider than long in females; head width 36.6-41.9 percent SVL (x = 39.9, N = 8); snout subacuminate in dorsal view, tip tends to be pointed, rounded in lateral profile; canthus _ rostralis rounded but distinct, slightly concave; loreal region concave, sloping gradually to lips; lips not flared; nostrils near tip of snout, not protuberant, directed dorso- laterally; snout of moderate length, eye length equal to or slightly greater than E-N distance; upper eyelid width 85.7- 112.9 percent IOD (x = 99.2, N = 8); interorbital space flat, frontoparietal bones complete (no fontanelle), not forming lateral crests (although crests may occur in larger individuals); tym- panum prominent, round, its length 30.0- 39.1 percent eye length (x = 34.8, N = 8) (slightly larger in males than in fe- males); upper edge of tympanum par- tially concealed by thick supratympanic fold; tongue round, weakly notched posteriorly, posterior one-third to two- fifths not adherent to floor of mouth; choanae moderate-sized, round, com- pletely visible when palate is viewed from directly above; vomerine teeth and odontophores present, round to oval in outline, each about twice the size of a choana, lying medial and posterior to choanae, separated medially by a dis- tance equal to choanal width, each bear- ing 3-5 teeth in a transverse row across posterior edge of process; males lacking vocal sac and vocal slits. Skin of dorsum finely tuberculate and bearing tuberculate ridges begin- ning at posterolateral corners of upper eyelids and converging medially onto scapular region; upper eyelids tubercu- late, no tubercle greatly enlarged; flanks weakly tuberculate; throat and venter coarsely areolate, skin on undersurfaces of thighs areolate, that on undersurfaces of forearms, shanks, and tarsi smooth; no dorsolateral folds; discoidal folds promi- nent, disc ending posteriorly well an- terior of groin; forearm bearing promi- nent ulnar tubercles, that on elbow not greatly elongate; two palmar tubercles, median largest; thenar tubercle oval, larger than either palmar tubercle; thenar surface bearing numerous, low supernumerary tubercles; subarticular tubercles large, round, flattened, simple; fingers bearing ill-defined lateral fringes; digits bearing discs on large pads, pads on thumb smallest; dilation ratios I: 1.2- 1.4 (x = 13), I: 1.9-2.2. (x = 22) ie 2.4-3.0 (x = 2.6), IV. 2.2-2.7 (x = 2.5); pad on thumb nearly round, those on other fingers broader than long; first fin- ger shorter than second. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 23 Hind limbs of moderate length, shank 49.2-55.4 percent SVL (x = 52.7, N = 8); inner edge of tarsus bearing a short tubercle-like fold just proximal to inner metatarsal tubercle; outer edge of tarsus bearing a series of enlarged spine- like tubercles beginning on the heel and extending distally to base of fifth toe; inner metatarsal tubercle large, not com- pressed, twice as long as wide, four to six times size of conical outer metatarsal tubercle; plantar surface bearing numer- ous weakly-defined supernumerary tu- bercles; toes bearing lateral fringes and basal webbing (web not extending be- yond proximal edge of basal subarticu- lar tubercles except between toes IV and V; subarticular tubercles like those of fingers but smaller; digits bearing discs on rounded pads, pads smaller than those of fingers; heel of adpressed limb reaches to between eye and nostril. In preservative, E. cryptomelas is gray to reddish-brown above’ with brown markings (interorbital bar, scap- ular W, sacral chevron, limb bars). Canthal stripes are absent and _ labial bars and supratympanic stripes are ob- scure. The limb bars are oblique and slightly narrower than the interspaces. The venter is off-white or cream with slight to prominent brown reticulation. The dorsal color extends onto the upper flanks where it is abruptly replaced by cream. The posterior lower flanks, an- terior and posterior surfaces of the thighs, concealed surfaces of the shank, and axilla are deep black. The black areas in the groin are not continuous across the venter. In life, E. cryptomelas was as fol- lows: “Dorsum tan, reticulated with brown markings; cream lines on upper flank; flanks cream reticulated and spotted with black; black patch in groin; limbs colored as dorsum; posterior sur- faces of thighs black; hidden shank cream bronze reticulated with black; venter bronze-cream with less definite black reticulations; iris coppery with broad reddish-copper median stripe.” (J. D. Lynch field notes, 16 VI 1968). Measurements of holotype in mm.— SVL 28.5, shank 15.8, head width 11.3, head length 10.5, upper eyelid width 3.2, IOD 3.2, tympanum length 1.2, eye length 3.5, E-N distance 3.5. The holo- type is an immature female with straight oviducts and small white ovarian eggs. Etymology.—Greek, in reference to the black areas on the concealed limb surfaces and in the groin and axilla. Natural history—The only speci- mens of this species available were found beneath rocks or in terrestrial bromeliads in pdramo and subpdramo habitats. Four females (19.7-29.1 mm SVL) are immature and show no be- ginnings of oviducial convolutions or enlargement of eggs. The males lack vocal slits but are mature (enlarged testes, swollen thumbs); hence the spe- cies is probably mute. One adult female (JDL 10396) was found by day under a rock ledge beside a stream in a pas- ture (7 January 1978) 7 km N San Lucas, Provincia Loja, Ecuador, 2840 m. Remarks.—Too few specimens are available to warrent preparation of a skeleton, thus restricting the number of characteristics with which to compare E. cryptomelas with other species. The appearance of the frog suggests to me that this species is not closely allied to the other high Andean species but is a member of one of the complexes of spe- cies found primarily on the Andean slopes. The black areas in the groin and on the concealed thigh surfaces, large digital pads, and row of tubercles along the outer edge of the tarsus serve to distinguish this species from all other congeners. Distribution.—Known only from ele- vations between 2470 and 3100 m on the eastern Andean cordillera of southern Ecuador. Eleutherodactylus lymani Barbour and Noble Fig. 9C Descriptions of this species are avail- able in Barbour and Noble (1920) and Parker (1932), the latter under the name 24 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY E. carrioni (Lynch, 1969a). Several more specimens of E. lymani have be- come available establishing the occur- rence of the species in the semi-arid val- ley of the Rio Catamayo in Provincia Loja, Ecuador, as well as in the semi- arid valley of the Rio Jubones in Azuay Province, Ecuador. The _ altitudinal range is 690-2500 m. Eleutherodactylus orestes new species Fig. 9D Holotype.—KU 141998, an adult fe- male, obtained 11 km NE Urdaneta, Provincia Loja, Ecuador, 2970 m, 24 July 1971 by William E. Duellman and Bruce MacBryde. Paratypes—KU_ 141999-142003, col- lected syntopically with the holotype. Diagnosis—(1) skin of dorsum fine- ly tuberculate, that of venter areolate; no dorsolateral folds; (2) tympanum obscure, round, its length one-third to one-half that of eye; (3) snout rounded to subacuminate in dorsal view, rounded in lateral profile; E-N less than eye length; (4) interorbital space flat, no cranial crests; upper eyelid width three- fourths IOD; (5) vomerine teeth and odontophores visible, odontophores flat- tened, oval; (6) males with vocal slits and median, subgular vocal sac; (7) first finger shorter than second, all digits bearing discs on moderately dilated pads: dilation ratios I: 1:2) I. 13, 0T: 1.6, IV: 1.5; (8) fingers lacking lateral fringes; (9) ulnar tubercles obscure; (10) inner and outer tarsal tubercles present, outer tarsal tubercles promi- nent; tubercles on heel small, non-coni- cal; (11) two metatarsal tubercles, outer round, conical, one-half to three-fourths size of non-compressed, slightly longer than wide inner metatarsal tubercle; plantar surface areolate; (12) toes not fringed, basal web present, all bearing discs on pads as large as those of outer fingers; (13) dorsum gray to brown with darker interorbital bar and dorsal chevrons; flanks darker than dorsum; venter gray to pale. brown spotted with brown; groin, anterior and_ posterior surfaces of thigh, concealed shank black enclosing large white spots; (14) adults small, one 6 19.8, @ 2 18.1-27.2 mm SVE: The small size, color pattern, and comparatively narrow digital pads of E. orestes prevent its confusion with any of the other named species of Eleuthero- dactylus. It is probably most closely allied to a sympatrant, E. vidua sp. nov., from which it differs in having a visible, although obscure, tympanic disk, larger digital pads (see Fig. 7B), and the bold black and white areas in the groin and on the concealed limb surfaces. Description—Head narrower than body, slightly wider than long; head width 33.9-37.7 percent SVL (xX = 35.4, N = 12); snout obtuse to subacuminate in dorsal view, rounded in lateral pro- file; canthus rostralis moderately sharp, weakly concave; loreal region weakly concave, sloping abruptly to lips; lips not flared; nostrils weakly protuberant, directed dorsolaterally, near tip of snout; snout short, E-N much less than eve length; upper eyelid width 68.0-84.2 percent IOD (x = 76.8, N = 11); inter- orbital space flat, no cranial crests; fron- toparietal bones in median contact, no fontanelle; tympanum present, visible externally although obscure in some specimens, its horizontal diameter 33.3- 49.6 percent eye length (x = 41.4,N = 11); tongue longer than wide, posterior one-fourth to one-third not adherent to floor of mouth, not notched posteriorly; choanae small, situated near edge of palate, partially concealed by palatal shelf of maxillae when roof of mouth is viewed from directly above; vomerine odontophores present, median and _ pos- terior to choanae, each process approxi- mately twice the size of a choana, and bearing 4-6 teeth in a patch; odonto- phores low and flattened; males with median subgular vocal sac and_ short vocal slits. Skin of dorsal surfaces finely tuber- culate, that of venter areolate; discoidal folds present, poorly defined, ending anterior to groin; vertebral ridge pres- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 25 ent; no dorsolateral or paravertebral folds present, except as rows of pustules; skin around anus not modified; ulnar tubercles poorly defined; two palmar tubercles, inner much larger than outer (Fig. 7), inner larger than oval thenar tubercle; palm usually bearing numer- ous supernumerary tubercles, all low, flat; subarticular tubercles of fingers moderate-sized, round, non-conical, sim- ple; fingers not bearing lateral fringes; digit tips dilated into pads broader than long, bearing discs on ventral surfaces; pads of inner fingers smaller than those of outer fingers; dilation ratios I: 1.1-1.2 (eee 2) Il: 10-14 (x = 13), WI: 1.5-1.7 (x = 16), IV: 1.4-1.6 (x = 1.5); first finger shorter than second. Hind limbs short, shank 38.3-49.8 per- cent SVL (x = 42.7, N = 16); tarsus bearing ill-defined tubercles on inner and outer edges, those on outer edge better defined than inner tarsal tubercle; several small, non-conical tubercles on heel; two metatarsal tubercles, inner larger, one and one-half to two times size of outer (Fig. 10); inner metatarsal tubercle slightly longer than wide, not compressed, outer longer than wide, conical; plantar surface areolate, none of the supernumerary tubercles especially prominent; subarticular tubercles round, not conical, simple; toes not bearing lat- eral fringes but basally webbed; fifth toe fused to fourth for nearly one-third its (fifth) length (Fig. 10); tips of toes bearing discs on pads, all pads equally large, slightly broader than long. In preservative, the dorsal surfaces vary from gray to dark brown with darker markings. The dorsal markings include an indefinite interorbital bar and chevrons on the back. The lower flanks are darker than the dorsum and suffused with dark brown marbling. The limbs are barred. The face is gray to dark brown with markings (labial bars, canthal stripe) visible in the more pale individuals but not in the more darkly pigmented individuals. The venter var- ies from dusky gray to pale brown usual- ly marbled and spotted with brown. Fic. 10.—(A) Foot of Eleutherodactylus orestes, KU 141995; (B) foot of E. phoxo- cephalus, KU 142063; line equals 5 mm. The throat tends to be paler than the spots on the venter. The groin, anterior thigh, posterior thigh, concealed shank, and concealed tarsus are dark brown to black with large white spots. This pat- tern is also evident in the axillae of a few individuals. In life, E. orestes is brown (rarely black) above with brown markings. The venter is gray with cream and/or dark brown to black spots. The posterior sur- face of the thigh and groin are black bearing ivory white, bluish-white, or pale pink spots. The iris is gray-bronze with a brown median horizontal streak. Measurements of the holotype in mm.—SVL 23.7, tibia 9.4, head width 8.4, head length 7.8, upper eyelid width 2.4, IOD 2.8, tympanum length 0.75, eye length 2.9. The holotype is a gravid fe- male with heavily convoluted oviducts and large, yellow, ovarian eggs. Etymology.—Greek, Orestes, a moun- taineer. Natural history—Adult females col- lected in May, June, and July have large eggs but no egg masses have been found. The frogs have been taken from terres- trial bromeliads but most individuals 26 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY were found beneath stones, in dirt banks, or in mulch in subpdramo habitats. A single male has been collected—it has vocal slits but was not observed or heard calling. No amplectant pairs have been found. Remarks.—The skull of E. orestes differs from those of most Andean Eleu- therodactylus in that the nasals are per- ceptibly separated (Fig. 11). The sym- patric E. vidua has a similar arrange- ment of the nasal bones. The fronto- parietals appear to be fused to the pro- otics and have a narrow, anterior fonta- nelle. The sphenethmoid is relatively large and extends anteriorly about one- half way beneath the comparatively small nasals. In ventral view, the vo- mers are relatively small and separated medially. The palatines are moderately long (compare with those of E. colo- dactylus). The anterior ramus of the parasphenoid is neither greatly short- ened nor elongate. The parasphenoid alae are oriented at right angles to the anterior ramus and in tenuous contact with the median rami of the pterygoids. The zygomatic ramus of the squamosal is greatly shortened (Fig. 12), recalling the condition usually seen in West In- dian species of Eleutherodactylus. The comparatively narrow digital pads and the color pattern in the groin caused me _ initially to suspect that this species was Paludicola simonsii (=Phrynopus simonsii) named by Bou- lenger (1900) from Cajamarca, Pert. Certain discrepancies in the description of simonsii prompted reservations which were borne out following examination of the cotypes of that species. However, the large sphenethmoid and separated nasal bones are features usually associ- ated with frogs of the genus Phrynopus (although not of the group to which simonsii belongs). The skulls of E. orestes and E. vidua do resemble those of the Phrynopus wettsteini group but the digital structure (external and inter- nal) of these narrow-toed Eleutherodac- tylus precludes assignment of these spe- cies to Phrynopus. Distribution.—Eleutherodactylus_ or- estes occurs at elevations between 2720 and 3120 m on the eastern Andean cor- dillera from the Cuenca hoya to the Loja hoya in southern Ecuador. Eleutherodactylus percultus new species Fig. 13A Holotype —KU_ 166058, an adult male, obtained at the Abra de Zamora, Provincia Zamora-Chinchipe, Ecuador, 2850 m, on 8 March 1975 by Dana K. Duellman. Paratype —KU 166057, an adult fe- male topotype, obtained on 7 March 1975. Diagnosis.—(1) skin of dorsum sha- greened anteriorly, covered with large flat warts posteriorly, that of venter are- olate; no dorsolateral folds; (2) tym- panum prominent, its length %-% eye length; (3) snout subacuminate in dor- sal view, tip pointed, acutely rounded in lateral profile, long; (4) upper eyelid narrower than IOD; low cranial crests; (5) vomerine odontophores prominent, triangular in outline; (6) males lacking vocal sac and slits; (7) first finger short- er than second; all fingers bearing broad discs on dilated pads; (8) fingers bear- ing lateral fringes; (9) ulnar tubercles non-conical; (10) heel tubercles sub- conical; outer edge of tarsus and foot bearing large conical tubercles; ventral surface of tarsus bearing non-conical warts; (11) two metatarsal tubercles, inner oval, non-compressed, 4 times size of outer; numerous supernumerary plan- tar tubercles; (12) toes bearing lateral fringes and broad discs on dilated pads; toe pads as large as those of fingers; (13) brown with gray and dark brown flecking and marbling; venter cream; flanks spotted with brown; upper lip bearing broad cream stripe from tip of snout to tympanum; in life, upper lip red; (14) adults moderate-sized, one ¢ 29.8 mm, one ? 38.2 mm SVL. On the basis of its coloration E. per- cultus is readily distinguished from all other Eleutherodactylus. Several spe- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 27 Fic. 11.—Skulls of Andean Eleutherodactylus. (A-C) lateral, dorsal, and ventral views of E. nicefori, KU 150724; (D-E) dorsal and ventral views of E. orestes, KU 151052; (F-G) dorsal and ventral views of E. vidua, KU 120093; line equals 5 mm. 28 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Fic. 12.—(A) Right squamosal of Eleutherodactylus unistrigatus (JDL S-263) and posterolateral corners of skulls of (B) E. orestes, KU 151052, and (C) E. vidua, KU 120093. Fic. 13.—(A) Eleutherodactylus percultus, KU 166058; (B) E. phoxocephalus, KU 131408; (C) E. versicolor, KU 119858; (D) E. vidua, KU 120089. cies have color variations including a pale lip stripe (E. achatinus, E. fitzing- eri, E. lanthanites, E. pygmaeus, E. rho- dopis) but in all of these the stripe is whitish or bronze-yellow in life and the labial pigmentation is all that distin- guishes the morph from other individ- uals of the species. Eleutherodactylus percultus resembles E. balionotus in tu- berculation of the tarsus, size of the digital pads, and proportions but is near- ly twice as large, males lack vocal sac and slits, and the skin on the anterior one-half of the body is shagreened rather than tuberculate. Description—Head as broad as body, broader than long; snout rounded with pointed tip to subacuminate in LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 29 dorsal view, acutely rounded in lateral profile; snout long; nostrils weakly pro- tuberant, directed dorsolaterally; can- thus rostralis rounded but evident; loreal region flat, sloping gradually to lips; lips weakly flared; interorbital space broader than upper eyelid; edges of frontopari- etals upturned slightly; upper eyelid bearing small, non-conical tubercles; temporal region sloping; supratympanic fold thick, concealing upper edge of tym- panum; tympanum distinct, round, sep- arated from eye by 1s times its diameter; postrictal tubercles prominent, non-con- ical: choanae moderate-sized to small, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores median and posterior to choanae, ele- vated, triangular in outline, each 1-4 times size of a choana, separated by dis- tance nearly equal odontophore width, each bearing 4-5 teeth in transverse row across posterior border; tongue slightly longer than broad, posterior % not ad- herent to floor of mouth, posterior edge notched; male lacking vocal sac and slits. Skin of dorsum shagreened anterior- ly, covered with large flat warts pos- teriorly; that of flanks and limbs like that of lower back; no dorsolateral folds; skin of venter coarsely areolate; discoidal folds not prominent; ulnar tubercles non- conical; palmar tubercle bifid, as large as oval thenar tubercle; supernumerary palmar tubercles indistinct; subarticular tubercles round, non-conical, simple; fingers bearing lateral fringes; all fingers bearing discs (broader than long) on pads; pads rounded apically; dilation rations» Te W518: TM: 1.822:7: I: 2:15 2.7; IV: 1.8-2.4; first finger shorter than second; thumb swollen in male. Knee and heel tuberculate; tubercles on heel subconical; outer edge of tarsus and sole bearing large conical tubercles; inner edge of tarsus bearing row of low warts; ventral surface of tarsus bearing non-conical warts; inner metatarsal tu- bercle twice as long as wide, non-com- pressed, four times size of elongate, non- conical outer; plantar supernumerary tu- bercles numerous, obscure; subarticular tubercles round, low, simple; toes bear- ing lateral fringes coalescing at base of toes forming basal web (enclosing prox- imal one-half of proximal subarticular tubercle on toes I-IV); all toes bearing discs (broader than long) on dilated pads; pads of toes as large as those of fingers; hind limbs long; heels of flexed hind limbs overlap; heel of adpressed hind limb reaches between eye and nostril. Brown above flecked with gray and darker brown; limb bars not evident: upper lip bearing cream stripe from tip of snout to tympanum; flanks spotted brown; venter cream with or without brown spots; undersides of limbs mot- tled brown; posterior surface of thigh brown with indefinite cream reticulation or flecking; anal triangle indistinct. In life, E. percultus was “Dull olive- brown becoming reddish brown posteri- orly and on limbs. Flanks bluish gray mottled with black. Upper lip bright orange. Throat creamy white with orange flecks. Belly bluish gray. Iris black with greenish gold flecks” (W. E. Duellman field notes, 7 March 1975). Measurements in mm (holotype, par- atype).—SVL 29.8, 38.2; tibia 15.5, 21.2: head width 12.1, 15.0; head length 11.0, 13.2; upper eyelid width 3.4, 3.2; IOD 3.9, 4.3; tympanum length 1.5, 2.0; eye length 3.5, 4.4; E-N 3.4, 4.3. The holo- type is an adult male with swollen thumb and large, non-pigmented, gran- ular testes. The venter bears few spots. Etymology.—Latin, meaning highly adorned, in reference to the orange lip stripes. Natural history——The holotype was found in a terrestrial bromeliad during the day. The paratype, a gravid female, was found on a branch of a bush at night. Eleutherodactylus phoxocephalus new species Fig. 13B Holotype—KU_ 142075, an adult male, obtained at Pilal6, Provincia Coto- paxi, Ecuador, 2340 m, on 7 July 1971 30 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY by William E. Duellman and Linda Trueb. Paratypes—KU_ 142076-103, collect- ed with the holotype. Diagnosis——(1) skin of dorsum sha- greened, that of venter areolate; no dor- solateral folds; (2) tympanum visible, round, its length one-third to one-half that of eye; (3) snout round in dorsal view except for pointed tip (a vertical keel), pointed in lateral profile; E-N slightly greater than eye length; (4) in- terorbital space flat, no cranial crests, upper eyelid width narrower than IOD; (5) vomerine teeth and odontophores present, odontophores slanted and tear- drop-shaped; (6) males with vocal slits and subgular vocal sac; (7) first finger shorter than second; all digits bearing discs on dilated pads, dilation ratios: PekG ele es We Vi 2 3F WV 2.2)(8)) fin= gers bearing indistinct lateral fringes; (9) ulnar tubercles inconspicuous or ab- sent; (10) tarsal surfaces lacking tuber- cles or folds (occasional individuals have indistinct outer tarsal tubercles) ; (11) two metatarsal tubercles, outer conical, round, one-sixth to one-fourth size of oval inner; plantar surfaces areo- late; (12) toes bearing lateral fringes and basal webbing; all toes bearing discs on pads, pads on outer toes larger than those on inner toes, toe pads smaller than those of fingers; (13) dorsum gray to brown, brown most evident posterior- ly; little or no dorsal color pattern evi- dent; groin, lower flanks, axilla, con- cealed surfaces of hind legs bearing white field bordered by and reticulated with dark brown or black; venter dirty cream, usually without brown flecking; (14) adults small to moderate-sized, males 22.3-29.9 mm, females 29.6-38.4 mm SVL: Eleutherodactylus phoxocephalus is unlikely to be confused with other spe- cies of Eleutherodactylus because it has a vertical keel at the tip of the snout; E. phoxocephalus is superficially similar to a frog called E. cruentus by Ruthven (1922) and Cochran and Goin (1970). That frog lacks a keel on the snout (it has a pointed snout), has slightly longer legs and a slightly broader head than E. phoxocephalus, and has a_ shorter snout (E-N three-fourths to four-fifths eye length compared to the nearly equal E-N and eye length of E. phoxocepha- lus). Description—Head narrower than body, wider than long; head width 33.4- 38.0 percent SVL (x = 36.1, N = 40) (females have slightly broader heads than do males); outline of head semi- circular in dorsal view except for verti- cal keel at tip of snout, in lateral profile snout pointed, tip extending beyond edge of lower jaw; canthus_ rostralis obtuse or rounded, concave; loreal re- gion flat, sloping gradually to non-flared lips; nostrils weakly protuberant, di- rected dorsolaterally; snout of moderate length, E-N 78.4-100.0 percent eye length in males (x = 93.7, N = 20), 94.7-113.3 percent in females (x = 102.6, N = 20); upper eyelid width 72.5-106.7 percent IOD (x = 92.5, N = 40); interorbital space flat, no cranial crests; frontoparietals complete, no fon- tanelle; supratympanic fold not promi- nent, but concealing upper edge of tym- panum; tympanum round, distinct, ob- lique in orientation (directed dorsolat- erally and somewhat posteriorly), sep- arated from eye by distance equal to one-half tympanic diameter; tympanum of male smaller than that of female, tym- panum length 32.4-54.5 percent eye length in males (x = 40.3, N = 20), 38.5-54.7 percent in females (x = 46.1, N = 20); tongue longer than wide, pos- terior one-half to two-fifths not adherent to floor of mouth, notched posteriorly; choanae small, round, completely visible when palate is viewed from directly above; vomerine odontophores lie medi- an and posterior to choanae, each is larger than a choana, separated medially by distance less than width of a choana, bearing 1-7 teeth; odontophores strongly slanted (oblique) and not greatly ele- vated above palatal surface; males have short vocal slits and a large external, subgular vocal sac. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 31 Skin of dorsum finely shagreened without folds or large warts except for thin sagittal ridge in larger females; flanks and skin posterior to ear areolate; skin of venter coarsely areolate, that of throat shagreened; concealed surfaces of limbs smooth except about vent; discoi- dal folds prominent. Most specimens lack ulnar tubercles or folds other than the antebrachial tu- bercle; palmar tubercle bifid, larger than oval thenar tubercle; palm bearing scat- tered supernumerary tubercles; subartic- ular tubercles simple, round, non-coni- cal; fingers bear weak, keel-like lateral fringes; all digits bearing discs on di- lated pads, pad of thumb slightly wider than long, those on other fingers distinct- ly wider than long; dilation ratios I: 1.5- Peo = 1,6), IW: 1.7-2.5 (x =,2.1), III: 1.9-2.6 (x = 2.3), IV: 1.9-2.6 (x = 2.2): first finger shorter than second; outer edge of hand bearing a cutaneous fold. Hind limbs of moderate length, shank 41.7-52.8 percent SVL (x = 47.5, N = 40) (shank of males slightly longer than those of females); limbs lacking tuber- cles except for faint inner tarsal tubercle in most individuals and a trace of outer tarsal tubercles in some individuals; in- ner metatarsal tubercle twice as long as wide, not compressed, four to six times as large as rounded, weakly elevated outer metatarsal tubercle (Fig. 10); toes bearing lateral fringes and a brief basal web (web not reaching base of basal subarticular tubercles except that be- tween toes IV and V); plantar surface bearing numerous supernumerary tuber- cles; subarticular tubercles like those of fingers but smaller; toes bearing discs on dilated pads, pads of toes wider than long, about same size as those of fingers. In preservative, E. phoxocephalus is gray-brown above with the lower back more brown than gray. The body usually lacks any trace of a color pattern. The posterior surfaces of the thighs are white with black edging and black reticula- tions on the white field. A similar pat- tern is evident in the groin and on the anterior surface of the thighs. The ven- ter is dirty cream with indefinite brown flecking (weak reticulation). The throat and breast lack reticulation and are yel- lowish-cream. A few adults do exhibit diffuse color patterns. Some individuals have brown and black flecking forming a loose pattern of chevrons, circles, and lines on the back. A frequent pattern consists of a dirty-brown spot on the snout and another above the sacrum. No individual has distinct limb bars al- though a few have faint transverse bars on the shanks. An individual from the slopes of Volcan El] Corazon (KU 109137) has a canthal stripe and a series of short black interrupted stripes on the dorsum (similar to the color pattern exhibited by E. ornatissimus). In life, E. phoxocephalus was de- scribed as follows: “Iris pale bronze with fine black reticulations and a brown horizontal streak; ground color highly variable—pale greenish-brown, pale yel- low, rusty brown, and light to dark brown, mottled and finely reticulated with brown to black (black toward cen- ter of back); axilla, groin, posterior thigh and concealed shank yellow, reticulated with dark brown (reticulations on flash colors developed ontogenetically); ven- ter cream, faintly washed with green, finely reticulated with brown; throat of female dusky, of male pale yellow, in- dividuals having pale dorsal spots have enamel-cream or cream spots edged in black” (J. D. Lynch field notes, 3 July 1970). Measurements of holotype in mm.— SVL 27.9, shank 13.6, head length 10.6, head width 10.4, upper eyelid width 2.9, IOD 3.5, tympanum length 1.3, eye length 3.2, E-N 3.2. The holotype is an adult male with vocal slits and a vocal sac. The testes are small and granular. Etymology.—Greek, in reference to the keel at the tip of the snout which gives the frog a “pointed head” appear- ance. Natural history—In subpdramo hab- itats south of the type-locality, E. phoxo- cephalus has been found beneath rocks 32 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY as well as in terrestrial bromeliads. At the type-locality, this species was the most abundant eleutherodactyline frog encountered in early July but nearly every specimen found was taken from bromeliads. The bulk of the bromeliads checked were the large terrestrial plants where other animals (Gastrotheca and Proctoporus as well as two other Eleu- therodactylus) were found, but E. phox- ocephalus was also taken in bromeliads in trees up to 3 meters above the ground. Sporadic calling at night from the large terrestrial bromeliads was heard in early July 1970, and late June 1977, but could not be identified to a particular frog. In January 1978, male E. phoxo- cephalus were actively calling at night and during afternoon rains. The call is a single sharp whistle. Males were call- ing on bushes and trees up to 5 m above the ground. Gravid females were taken in number during June and July. Indi- viduals placed together frequently en- gaged in axillary amplexus, and one am- plectant pair was found in a bromeliad (June 1970). The smallest gravid female collected is 29.6 mm SVL—the largest juvenile female (small ovarian eggs, non- convoluted oviducts) is 30.1 mm SVL. The smallest “adult” male (with vocal slits) found is 22.3 mm SVL. A subadult male (no vocal slits) is 21.0 mm SVL. Remarks.—The southern samples of E. phoxocephalus (Azuay, Cafiar, Loja, and Zamora-Chinchipe provinces) were taken in subpdramo habitats at eleva- tions between 2060 and 2960 m. The northern samples (Cotopaxi and Pichin- cha provinces) were taken in cloud for- est habitats at elevations between 2030 and 2580 m. In spite of the striking ecologic differences among localities, the frogs all seem to be conspecific. Collec- tions made at the appropriate elevations in Provincia Bolivar failed to yield spec- imens of this species, but the absence may have been due to the locally arid environments around Guaranda. I was unable to collect at what seemed to be ideal habitat between Guaranda and Balzapamba (elevation of 2000-2200 m). In attempting to discover if this spe- cies had been previously named, I en- countered specimens of a similar species from Colombia identified as E. cruentus. The frog Ruthven (1922) reported from Colombia under this name is not cruen- tus of Peters. That frog was named E. dubitus by Taylor (1952) from Costa Rica and E. marshae by Lynch (1964) from Panama (as pointed out by Savage, 1965) and is similar to, but not identical with, E. latidiscus (Boulenger) of the Pacific lowlands of Colombia and Ecua- dor. The Colombian frog confused with cruentus by Ruthven and Cochran and Goin (1970) apparently is unnamed. Distribution—Intermediate _ eleva- tions (2030-2960 m) on the Pacific slopes of the Ecuadorian Andes from Volcan El] Corazon to the Loja hoya; E. phoxo- cephalus enters the interandean hoja south of Cuenca and occurs along the northern and eastern edges of the Loja hoja. Eleutherodactylus proserpens new species Fig. 14 Holotype -——USNM 198484, an adult female obtained between Sapote and Suro Rancho, Provincia Morona-Santia- go, Ecuador, 2622 m, by M. Olalla on 24 August 1962. Paratypes—USNM _ 198486-91, % km W Sapote, 2546 m; USNM 198492-95, 198502, bet. Sapote and Suro Rancho, 2622 m; USNM_ 198496-98, mountain above Sapote to south, 2500 m; USNM 198499-501, 1 km E Sapote, 2332 m; all Provincia Morona-Santiago, Ecuador. Diagnosis.—(1) skin of body areo- late, no dorsolateral or discoidal folds; anal opening extended by skin sheath onto posterior surface of thighs; (2) tympanum prominent, round, its length 4-4 that of eye; (3) snout subacuminate in dorsal view, round to pointed in lat- eral profile, tip bearing papilla; (4) in- terorbital space flat, broader than upper eyelid width; (5) vomerine odonto- phores low, oval; (6) males lacking vo- cal sac and slits; (7) first finger shorter LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 33 Fic. 14.—(A-B) head of Eleutherodactylus proserpens, USNM 198499; (C) hand of E. proserpens, USNM 198494; lines equal 1 mm. than second; all digits short, stocky, bearing discs on broad, non-emarginate pads; distal subarticular tubercles of III and IV bifid; (8) fingers bearing ridge- like lateral fringes; (9) no ulnar tuber- cles or folds; (10) knee, heel, and tar- sus lacking tubercles; (11) two meta- tarsal tubercles, inner twice size of out- er; plantar surface areolate; (12) toes bearing ridge-like lateral fringes; toes short, stocky, bearing discs on broad pads; (13) color pattern polymorphic, most common morph is: dorsum tan to brown with darker brown limb bars, in- terorbital bar, X-shaped mark on back, postocular bar; venter pale brown; pos- terior thigh cream to brown; other frogs have pale dorsolateral stripes; (14) adults small, males 15.2-21.0 mm, fe- males 20.2-23.5 mm SVL. Eleutherodactylus proserpens is simi- lar to E. celator Lynch and E. colodac- tylus Lynch in size, having rounded canthi, short limbs, and areolate skin. It resembles E. colodactylus in having stubby digits. Some of these features may be adaptations to living in bromeli- ads (where all known examples of all three species have been found). Eleu- therodactylus proserpens differs from E. celator and E. colodactylus in having an anal sheath and differs from E. colo- dactylus in having fully developed ears and prominent vomerine odontophores. Eleutherodactylus proserpens differs from E. celator in having a prominent papilla on the snout. Description.—Head as wide as body (not as wide as body in gravid females), wider than long; head width 31.8-40.6 percent SVL (x = 34.1, N = 15); snout acuminate in dorsal view, tip pointed, round or weakly pointed in lateral pro- file; snout long, E-N 87.8-122.2 percent eye length (x = 105.5, N = 15); can- thus rostralis rounded, straight; nostrils not protuberant, directed dorsolaterally; loreal region weakly concave, sloping 34 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY gradually to lips; lips not flared; inter- orbital space flat, no cranial crests; up- per eyelid width 54.2-78.3 percent IOD (x = 63.0, N = 13) (eyes small); su- pratympanic fold thick, concealing up- per edge of tympanum; tympanum dis- tinct, round, separated from eye by 12-2 times tympanic diameter, its length 30.3- 50.0 percent eye length (x = 38.2, N = 15); one to several non-conical tubercles on upper eyelid; tongue moderate-sized, 1% times as long as wide, posterior edge not notched, posterior % not adherent to floor of mouth; choanae small, round, not concealed by palatal shelf of maxil- lary arch; vomerine odontophores round, low, posterior and median to choanae, separated by %- width of a choana or odontophore, each bearing clump of 2-4 teeth; males lacking vocal sac and slits. Skin of body areolate (smooth skin in axillae, groin, behind knee, concealed shank and tarsi); no dorsolateral folds; anal opening enclosed in sheath extend- ing vent onto posterior surfaces of thighs; no discoidal folds; ulnar tuber- cles not distinct from areolations of skin except for antebrachial tubercle; two palmar tubercles, inner largest, slightly larger than oval thenar tubercle; nu- merous supernumerary palmar tubercles; basal subarticular tubercles broader than long, non-conical; distal subarticu- lar tubercles of fingers III-IV bifid; fin- gers bearing ridge-like lateral fringes; all fingers bearing discs on dilated pads; discs slightly broader than long; all dig- its short; first finger shorter than second. No prominent tubercles on knee or heel (each does have one larger areola- tion but this wart is not conical or elon- gated); tarsus lacking distinct tubercles or folds; outer metatarsal tubercle round, non-conical, %-% size of elongate, non-compressed inner metatarsal tuber- cle; numerous supernumerary plantar tubercles, some as large as subarticular tubercles; subarticular tubercles round, non-conical, simple; toes bearing ridge- like lateral fringes and basal webbing (coalesced lateral fringes); toes bearing discs on round pads; discs as broad as long; all toes short; shank short, shank of males 41.0-47.2 percent SVL (x = 43.3, N = 11), of females 37.8-46.4 percent (x = 40.7, N = 7). In preservative, tan to brown or red- dish-brown above with brown interorbi- tal bar, X-shaped mark on back, postoc- ular bar, and limb bars; or with same ground color—and cream dorsolateral stripes edged medially with dark brown; all have gray to brown venter. Eleven individuals have the first pattern listed above; two others have a single blotch extending from the eyes to the sacrum (dorsal pattern similar to that of Hyla leucophyllata); five have pale dorso- lateral stripes enclosing a dark dorsum; and two have dark dorsolateral stripes enclosing a pale dorsum. In life, E. proserpens was described as “dorsum orange with gray markings, venter gray; dorsum rusty brown with darker brown markings, venter pale yel- low with minute brown flecks; and dor- sum tan, venter light tan” (John E. Sim- mons field notes, 4-7 January 1972). Measurements of holotype in mm.— SVL 22.5; tibia 8.5; head width 7.3; up- per eyelid width 1.6; IOD 2.5; tympa- num length 0.9; eye length 2.0; E-N 2.2. The holotype is a gravid female with convoluted oviducts and large ovarian eggs (2.5-2.7 mm diameter). Etymology.—Latin, one who creeps, in reference to the short limbs and pre- sumed non-hopping gait of this small frog. Natural history——All of the speci- mens secured by Peters and his associ- ates were collected in bromeliads. All males over 15 mm SVL have large gran- ular testes (lacking brown or black peritoneal linings) and swollen thumbs. Peters’ field notes are not precise but apparently E. proserpens and E. colo- dactylus were collected in the same bromeliads. Both species are of similar size and coloration and could be easily confused in life. The specimens col- lected by John Simmons on the Cordil- lera del Condor were found on low vege- tation at night. Gravid females were LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 35 found in January and August. Remarks.—At present, E. proserpens seems to be related to E. colodactylus, and I know of no other species sufficient- ly similar to these two to be considered close relatives. Eleutherodactylus pros- erpens is larger than E. colodactylus (adult females of former xX = 22.0, of latter x = 20.5) and the size difference is significant (p < 0.05); the size dif- ferential (¢ = 1.07) is much lower than that suggested by Hutchinson and Mac- Arthur (1959) as adequate to prevent competition. Distribution—Known at elevations between 1707-2622 m on the Amazonian slopes of the eastern Andean Cordillera in southern Ecuador and from the adja- cent Cordillera del Condor. Eleutherodactylus pycnodermis new species Fig. 15 Holotype-—USNM 199754, adult fe- male taken at San Vicente, Provincia Morona-Santiago, Ecuador, 2805-2835 m, by James A. Peters on 26 August 1962. Paratypes—(Provincia Azuay) USNM 199851-53, 3 km E Sevilla de Oro, 2713 m; USNM 199854, 5 km ESE Sevilla de Oro, 2957 m; USNM 199855- 59, crest at Azuay-Morona-Santiago bor- der, ca. 8 km SE Sevilla de Oro, 3354 m; USNM_ 199758-59, 199860-65, crest at Azuay-Morona-Santiago border, ca. 8 km SE Sevilla de Oro, 3384 m; (Provin- cia Morona-Santiago) USNM 199755-57, between Cerro Negro and Pailas, 2652 m; USNM 199815-50, San Vicente, 2805- 35 m; USNM 199813-14, San Vicente, 2851 m; USNM 199794-812, 3 km W San Vicente, 2988 m; USNM 199787-93, 6 km W San Vicente, 3100 m; USNM 199760- 66, Suro Rancho, 2683 m; USNM 199767-86, 0.5 km W Suro Rancho, 2744 m. Diagnosis —(1) skin of dorsum thick and glandular, surface texture coarsely shagreened; no dorsolateral folds; skin of venter coarsely areolate; Fic. 15.—Eleutherodactylus pycnodermis, USNM 199801; line equals 10 mm. (2) tympanum prominent, its length '-% eye length; (3) snout subacuminate in dorsal view, truncate in lateral profile; (4) interorbital space broader than up- per eyelid; low cranial crests present; (5) vomerine odontophores prominent, oblique in males, triangular in outline in females; (6) males with vocal slits and subgular, external vocal sac; (7) first finger shorter than second; fingers bear- ing discs on broad pads; (8) fingers bearing narrow lateral fringes; (9) no ulnar tubercles except antebrachial; (10) no tubercles on heel or tarsus; (11) two metatarsal tubercles, inner oval, 4 times size of outer; few plantar supernumerary tubercles; (12) toes bearing lateral fringes; pads on toes dilated, smaller than those of outer fingers; (13) brown without limb bars; large black blotches on flanks, concealed surfaces of thigh, shank, and tarsus: venter creamy, fre- quently spotted with brown; (14) adults moderate-sized, males 18.0-32.3 mm, females 32.5-44.4 mm SVL. Two features set E. pycnodermis apart from the other nearly 400 species of the genus—none has the thick glan- dular skin covering the dorsal surfaces of the body and limbs and no other spe- cies has the very distinctive color pattern of this frog. Like most other highland 36 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Ecuadorian species of Eleutherodacty- lus, E. pycnodermis has short legs and some evidence of cranial crests. Description—Head narrower than body, wider than long; head width 34.6- 41.1 percent SVL (x = 37.5, N = 35); snout subacuminate in dorsal view, trun- cate in lateral profile; snout short, E-N 65.8-82.1 percent eye length in males (x = 72.8, N = 16), 73.3-95.1 percent in females (Xx = 82.8, N = 18); nostrils not protuberant, directed dorsolaterally; canthus rostralis sharp, straight; loreal region weakly concave, sloping abruptly to lips; lips not flared; interorbital space flat, edges of frontoparietals weakly up- turned forming low cranial crests; upper eyelid width 67.4-111.1 percent IOD (x = 89.2, N = 25); no tubercles on eyelid; temporal region vertical; supra- tympanic fold present, concealing upper edge of tympanum; tympanum promi- nent, round, its length 22.6-48.7 percent eye length in males (x = 35.4, N = 16), 32.5-50.0 percent eye length in females (x = 40.2, N = 18), separated from eye by tympanic width; choanae moderate sized, round, not concealed by palatal shelf of maxillary arch; vomerine odon- tophores median and posterior to choa- nae, each 1%-2 times size of a choana, oblique in smaller individuals (males and young females), triangular in out- line in large females, elevated, bearing a transverse row of 3-7 teeth along pos- terior edge; tongue longer than wide, posterior % not adherent to floor of mouth, its posterior border not notched; males with vocal slits, large external vocal sac. Skin of dorsum and limbs coarsely shagreened, very thick and glandular; no dorsolateral folds; skin of flanks smooth, that of venter coarsely areolate; discoidal folds prominent; no anal sheath; antebrachial tubercle prominent, no Other ulnar tubercles; palmar tuber- cle bifid, larger than oval thenar tuber- cle; supernumerary palmar tubercles present, indistinct; subarticular tubercles round, elevated, non-conical; fingers bearing weak lateral fringes; all fingers bearing broad discs on dilated, apically rounded pads; pads on fingers II-IV larger than tympanum; first finger short- er than second. No distinct tubercles on knee, heel, or tarsus; inner metatarsal tubercle ele- vated, non-compressed, its length 1% times its width, 4 times size of low, rounded outer metatarsal tubercle; su- pernumerary plantar tubercles obscure, small; subarticular tubercles round, low; toes bearing narrow lateral fringes; toes bearing broad discs on dilated pads; pads of toes smaller than those on outer fingers; when legs are flexed at right angles to body, heels overlap; heel of ad- pressed hind limb reaches to tympanum; shank 39.4-49.2 percent SVL (x = 43.5, N = 35). In preservative dark brown without limb bars, dorsal spotting, labial bars, or canthal and supratympanic stripes; large black blotch behind eye extending onto anterior flank, smaller blotch on upper arm, large blotch on posterior flank ex- tending to above insertion of hind leg with a lateral ramus extending along anterodorsal surface of thigh to knee, another on posterior surface of thigh from vent to knee, another along con- cealed surface of shank, and one or two on dorsal surface of tarsus and foot; black blotches edged above by cream; no anal triangle; inner digits creamy- yellow; venter dull cream or yellowish cream lightly to heavily spotted with brown; ventral to the black areas in the axilla, groin, and posterior thigh, the skin is pale yellow. Unfortunately colors in life were not recorded by the collectors nor were colored photographs taken of this abun- dant species. Measurements of holotype in mm.— SVL. 37.3; tibia 16.6; head width 13.6; upper eyelid width 3.6; IOD 4.3; tym- panum length 1.6; eye length 4.0; E-N Bios Etymology.—Greek, pyknos and der- ma, meaning thick skin, in reference to the distinctive thick glandular skin of this species. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 37 Natural history —Eleutherodactylus pycnodermis was collected in pdramo as well as in meadows within the upper reaches of montane forests 700 meters lower on the Andean slopes. All speci- mens having specific field notes were collected by day beneath rocks and logs. Many males have wrinkled, distended vocal sacs and one adult female (USNM 199841) had ovulated. None of the adult females is spent. The smallest adult female is 32.5 mm SVL. Several young females (weakly convoluted ovi- ducts, small ovarian eggs) are larger than the smallest adults. Twelve young females are 28.1-35.7 mm SVL [x = 32.1 seule 2SE) |: Remarks.—Eleutherodactylus pycno- dermis is a member of the wnistrigatus group (Lynch, 1976a) but its peculiarly thick skin is quite different from all other species of the genus. The frog seems best viewed as a large member of the assemblage including FE. balionotus Lynch, E. coeruleus (Andersson), E. orcesi Lynch, E. riveti (Despax), E. thymelensis Lynch, and E. unistrigatus (Gunther); I do not now consider it closely related to any of these. Distribution—Known only from the Cordillera de Matanga east of Cuenca at elevations of 2652-3384 m. Eleutherodactylus riveti (Despax) Fig. 16 Hylodes riveti Despax, 1911:92 [Holotype.— MHNP_ 1902-357, an adult female from “Equateur’ J. Eleutherodactylus riveti—Barbour and Noble, 1920:404. [Eleutherodactylus curtipes (part )]—Lynch, 1969:266. Eleutherodactylus w-nigrum (misapplication) —Lynch, 1972:144. Fic. 16.—Eleutherodactylus riveti. (A) KU 119814; (B) KU 119813; (C) KU 120028; (D) KU 120026. 38 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Diagnosis ——(1) skin of dorsum bear- ing flattened warts, that of venter coarsely areolate; dorsolateral folds faintly evident; (2) tympanum promi- nent, round, its length one-third to two- thirds that of eye; (3) snout subacumi- nate in dorsal view, rounded in lateral profile; snout short, E-N less than to equal eye length; (4) interorbital space broad, flat, no cranial crests; upper eye- lid width three-fourths IOD; (5) vo- merine odontophores round; (6) males with vocal slits and subgular vocal sac; (7) first finger shorter than second, all digits bearing discs on apically dilated pads; dilation ratios I: 1.3, II: 1.6, HI: 1.7, IV: 1.7; (8) fingers bearing promi- nent lateral fringes; (9) ulnar tubercles present, but ill-defined; (10) tarsus tu- berculate, but distinct tubercles not ap- parent; (11) two metatarsal tubercles, outer conical, one-fifth to one-fourth size of oval (length twice width) inner meta- tarsal tubercle; few supernumerary plan- tar tubercles, at base of toes; (12) toes bearing lateral fringes; only basal web- bing present between toes IV and V; all toes bearing discs on pads, pads as large as those of fingers; (13) dorsum brown with or without dark brown and/or black markings; venter cream, usually with brown reticulations; individuals from most localities have prominent can- thal and supratympanic stripes, broad limb bars and dorsal chevrons and spots or a dorsal pattern of brown and black stripes; the stripes or portions of dorsal chevrons on the upper flanks are black; in populations having little dorsal mark- ings, the upper flanks are spotted with black (Fig. 16); concealed thigh and groin dull brown; (14) adults small, males 20.6-26.8 mm, females 25.6-32.7 mm SVL. Eleutherodactylus riveti is consid- ered to be most closely allied to E. bali- onotus. Both are readily distinguished from E. curtipes in having larger digital pads, prominent tympana, and in lack- ing cranial crests. Eleutherodactylus ri- veti has very short limbs (equally short as those of E. thymelensis of northern Andean Ecuador). These two species have even shorter limbs than E. curtipes, a slightly larger species with cranial crests which is sympatric with E. thy- melensis. Eleutherodactylus balionotus has longer limbs although they are still appropriately described as short. E. ri- veti has weakly developed dorsolateral folds and thereby resembles E. curtipes (prominent dorsolateral folds). Description—Head narrower than body, wider than long; head width 35.5- 39.0 percent SVL (x = 37.1, N = 32); snout subacuminate in dorsal view, rounded in lateral profile; canthus ros- tralis sharp, straight; loreal region weak- ly concave, sloping abruptly to lips; lips not flared; snout of moderate length, eye length equal to or slightly greater than E-N; interorbital space flat, no cranial crests; frontoparietals complete, no fon- tanelle; upper eyelid width 69.1-93.3 percent IOD (x = 77.0, N = 31); tympanum prominent, round, its hori- zontal diameter 35.5-69.8 percent eye length (x = 44.6, N = 32), its upper edge concealed by thick supratympanic fold; tongue large, longer than wide, posterior one-third not adherent to floor of mouth, weakly notched posteriorly; choanae small, round, completely visible when roof of mouth is viewed from di- rectly above or partially concealed by edge of jaw; vomerine odontophores round, situated between and well pos- terior to choanae, nearly in median con- tact, bearing teeth arranged in a trans- verse row; each odontophore about twice size of a choana; male with vocal slits and subgular vocal sac. Skin of dorsum appears smooth but bears low, flattened warts, that of flanks, venter and posterior surfaces of thighs areolate, warts flat; discoidal folds prom- inent; dorsolateral and postocular folds faintly evident in many specimens; up- per eyelid not tuberculate. Ulnar tubercles present but not prominent; two palmar tubercles, medi- an largest, larger than oval thenar tu- bercle; thenar surface bearing a few low, flattened supernumerary tubercles; sub- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 39 articular tubercles round, flattened, mod- erate-sized, simple; fingers bearing thin lateral fringes; all fingers bearing discs on dilated pads, pads on outer two fin- gers larger than those on inner two fingers; pad on first finger round, those on other fingers wider than long; dilation manos i: VI-16 (x = 1.3), I: 1.2-2.0 Ga 9h6), TI: 15-2.0 (x = 17), IV: 14-19 (x = 1.7); first finger shorter than second. Hind limbs short, heel of adpressed limb reaches to shoulder, vicinity of tympanum, or in very young specimens, to posterior edge of eye; shank 37.2-45.9 percent SVL (xX = 41.6, N = 32); skin of heel and tarsus tuberculate, none on heel enlarged; ill-defined row of outer tarsal tubercles present, one or two tu- bercles on inner edge of tarsus; inner metatarsal tubercle oval, not com- pressed, twice as long as wide, four to five times as large as conical outer meta- tarsal tubercle; plantar surface bearing two to four supernumerary tubercles at bases of toes II, III, IV, and V; sub- articular tubercles of toes like those of fingers; toes bearing lateral fringes; basal web between toes IV and V; toes bearing discs on pads, pads wider than long, about as large as those of outer fingers. In preservative, E. riveti from those populations with a distinct color pattern (Fig. 16A, B) is brown to rusty-brown (rarely gray) with dark brown and/or black spots or stripes. The flanks are cream reticulated with brown or black. The canthal and supratympanic stripes are dark brown. The labial bars are ob- scure and the lips are not striped with cream. The limbs are barred with brown, the bars are wider than the inter- spaces and edged with gray-brown lines. The dorsal markings may also be edged with gray-brown or cream. The venter is cream reticulated with dark brown or black. The concealed thigh, shanks, and groin are colorless and speckled with cream. Specimens from “pattern-less” populations (Paramos de Matanga) have a bronze or brown ground color. The venter is cream and may or may not be reticulated with brown. The upper flanks may bear spots (Fig. 16, C, D) but the limbs, center of the back, and face lack markings. In life, E. riveti (patterned popula- tions) is ‘colored as follows: “Dorsum pale to medium brown, some yellow- brown with dark brown or black stripes; some rust above, all with some brown markings; snout tan in some individuals; some individuals with a green wash on markings; limb bars brown; flanks tan to light brown with brown to black bars and spots; vocal sac of male pale yellow; venter pearl white to pale rose with brown reticulations; posterior surfaces of thighs pale maroon brown to rust reticu- lated with cream; iris pale copper-tan above, brown below with black reticu- lations” (J. D. Lynch field notes, 19 June 1968). The “patternless” popula- tions are gray, gray-brown, reddish, or orange in life with black spots on the up- per flanks; the lower flanks are colored as the dorsum but a paler hue; venter off-white to cream, sometimes flecked with black; posterior surface of thigh same color as dorsum, sometimes more cream with faint thin brown reticulation. The iris of living specimens appears as above or may be “pale blue green, heay- ily reticulated with black; with a dull reddish-brown horizontal streak” (J. D. Lynch field notes, 18 June 1968) or “gray with a faint green cast and a brown to reddish brown horizontal streak; some black flecking” (J. D. Lynch field notes, 26 July 1970). Measurements of holotype in mm.— SVL 32.5, shank 13.0, head width 12.5, head length 11.6, upper eyelid width 3.2, IOD 3.6, tympanum length 1.5, eye length 3.7. The holotype is a gravid fe- male (large yellow ovarian eggs and strongly convoluted oviducts). Natural history—At the four locali- ties where I collected E. riveti (Laguna de Zurucuchu, 8 km NW Biblian, 10 km S Cumbe, and Paramos de Raranga, SE of Cuchil) only two other species of Eleutherodactylus were found (E. cryo- 40 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY philius at Laguna de Zurucuchu and E. orestes south of Cuchil, Pdéramos de Raranga), although E. phoxocephalus was found at slightly lower elevations on the slopes of the Cuenca hoja. No direct evidence of reproductive activity has been found in spite of the great abun- dance of these frogs beneath rocks and in dirt banks. Gravid females have been collected whenever the species was found. E. riveti inhabits subpdramo and pd- ramo habitats between 2620 and 3420 m in the Andes of Ecuador south of Cuen- ca. The frogs were equally abundant in the predominantly grassy pdramos and in the shrubby subpdramos. These are the same type of ecologic situations in which to the north one encounters E. curtipes. Remarks.—I have twice misapplied names to this frog. In redescribing E. cajamarcensis (Lynch, 1969a), I indi- cated that E. riveti was the same as E. curtipes. This action was based on a translation of Despax’s (1911) descrip- tion, the illustrations he provided, and the discovery of some E. curtipes labeled as E. riveti in the collections amassed by the late James A. Peters. In 1968 I col- lected at Laguna de Zurucuchu in an attempt to collect topotypes of Hylodes w-nigrum. I assumed the frogs collected there to be w-nigrum and under that as- sumption applied that name to this spe- cies (Lynch, 1972a). The specimens from Zurucuchu did not fit the descrip- tion and following study of photographs of the holotype of H. w-nigrum gener- ously provided by Dr. Konrad Klemmer I realized that I had again misapplied a name to this population. Eleutherodac- tylus w-nigrum, as correctly pointed out by Cochran and Goin (1970) is a spe- cies of the fitzingeri group, previously called E. buergeri. Eleutherodactylus riveti lacks a pro- nounced frontoparietal fontanelle, has moderately large nasals in tenuous me- dian contact, and has a comparatively large part of the sphenethmoid exposed dorsally, characteristics also seen in E. curtipes and E. unistrigatus (Fig. 17). The frontoparietals appear to be fused to the prootics. The vomers are mod- erate-sized and are in contact with the elongate palatines. The anterior ramus of the parasphenoid does not reach the vomers or palatines and the parasphe- noid alae are slightly deflected posterior- ly but in contact with the median rami of the pterygoids. Distribution.—Both Andean cordil- leras and the connecting nudos at eleva- tions of 2620-3420 m surrounding the Cuenca hoja in southern Ecuador. Eleutherodactylus ruidus new species Fig. 18 Holotype—AMNH 17590, adult fe- male obtained at Molleturo, Provincia Azuay, Ecuador, 2317 m, on 5-19 June 1922 by G. H. Tate. Paratypes—AMNH 17588-89, 17591- 96, 17598-601, 17603, topoparatypes. Diagnosis—(1) skin of dorsum warty, that of venter coarsely areolate; no discoidal folds; (2) tympanum, ca- vum tympanicum, and plectrum absent; (3) snout round in dorsal view (tip feebly pointed), angularly round in lat- eral profile, short; (4) interorbital space as wide as upper eyelid; no cranial crests; (5) vomerine odontophores tri- angular in outline; (6) males lacking vocal sac and slits; (7) first finger short- er than second; digits bearing broad discs on dilated pads; (8) fingers bear- ing lateral fringes; (9) ulnar tubercles absent; (10) heel bearing non-conical tubercles, none on outer edge of tarsus; inner tarsal fold present; (11) two meta- tarsal tubercles, inner oval, twice size of round outer; supernumerary plantar tu- bercles only at bases of toes II-IV; (12) toes bearing lateral fringes; toe pads smaller than those of outer fingers; (13) brown above with darker scapular chev- ron, interorbital bar, postocular stripe, and labial bars; venter dusky cream with diffuse spotting on throat; groin, anterior and posterior thigh surfaces, concealed shank brown; (14) adults LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 4] moderate-sized, males 25.8-31.1 mm, fe- the surdus Artenkreis, E. baryecuus males 37.1-39.8 mm SVL. Lynch and E. surdus (Boulenger). Both In its most obvious characteristics, differ from E. ruidus in having cranial E. ruidus is similar to two members of crests, smooth skin on the dorsum, longer Fic. 17.—Skulls of Andean Eleutherodactylus. (A) E. unistrigatus, KU 111137; (B-C) E. curtipes, KU 109057; lines equal 5 mm. Fic. 18.—Eleutherodactylus ruidus, AMNH 17590; line equals 5 mm. 42 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY legs, and in color patterns. Too few data are available to assert that E. ruidus is a member of the surdus Artenkreis. In body proportions and other shape characteristics, E. ruidus resembles E. balionotus Lynch and E. riveti (De- spax); it differs from them in lacking the middle ear and vocal apparatus. The three are likewise similar in the features of the hands and feet, skin texture, pro- portions, and lack of cranial crests. Description—Head wider than body, wider than long; head width of males 30.2-33.8 percent SVL (x = 32.8, N = 7), of females 34.5-35.9 percent (x = 35.2, N = 7); snout round in dor- sal view (tip feebly pointed), angularly round in profile; E-N 64.7-88.9 percent eye length in males (x = 73.4, N = 7), 71.1-87.8 percent in females (x = 80.0, N = 7); nostrils not or weakly protu- berant, directed dorsolaterally; canthus rostralis rounded; loreal region concave, sloping to non-flared lips; upper eyelid 80.8-124.1 percent IOD (x = 94.9, N = 13); interorbital space flat (no cranial crests ); upper eyelid warty (not bearing enlarged tubercles); supratympanic fold prominent, extending from corner of eye to base of arm; no tympanum, cavum tympanicum, or plectrum; two _ large postrictal tubercles; choanae small, not concealed by palatal shelf of maxillary arch, round; vomerine odontophores ele- vated, triangular in outline, median and posterior to choanae, separated by a choanal width, each slightly larger than a choana, bearing 1-6 teeth in a trans- verse row; males have fewer vomerine teeth (1-3 per odontophore) than do females (2-6 per odontophore); number of vomerine teeth positively correlated (r = 0.778, N — 2 = 13, p < 0.01) with body size; tongue slightly longer than wide, rounded posteriorly, posterior one- fourth not adherent to floor of mouth; no vocal sac or slits in male. Skin of dorsal surfaces (including upper eyelid) warty, W-shaped ridge on occiput; some larger warts on flanks; no dorsolateral folds; venter coarsely areolate, throat smooth; discoidal folds relatively prominent; skin of thigh (ven- tral surface) coarsely areolate; pair of large warts ventrolateral to vent; anus opens at posterior level of thighs (in a short sheath); ulnar tubercles indistinct; palmar tubercle bifid, twice size of oval thenar tubercle; supernumerary palmar tubercles elongate; subarticular tuber- cles round, non-conical; digits bearing relatively prominent lateral fringes (most evident on inner margins of fin- gers); all digits bearing broad (wider than long) discs on pads; pad of thumb round, those of fingers II-IV broadly di- lated, all rounded apically, none bearing ungual notch; thumb shorter than sec- ond finger. One or two small round tubercles on heel, none on outer edge of tarsus; inner edge of tarsus bearing fold along distal one-half; inner metatarsal tubercle not compressed, its length twice its width, two times size of round, non-conical out- er metatarsal tubercle; plantar surface bearing small supernumerary tubercles at bases of toes II-IV; subarticular tuber- cles round except for most basal tuber- cle on toes IV and V (those smaller than others, longer than wide); toes bearing prominent lateral fringes (including outer edges of inner and outer toes); toes basally webbed only in that fringes coalesce basally; toes bearing discs (broader than long) on dilated pads (1.5 times width of digit); toe pads smaller than those of fingers; when legs are flexed at right angles to sagittal plane, heels touch; heel of adpressed hind limb reaches to posterior corner of eye; shank 42.1-46.5 percent SVL in males (x = 44.5, N = 7), 42.4-49.3 per- cent in females (x = 47.3, N = 7). In preservative brown with darker brown sacral chevron, interorbital bar, and dark stripes lateral to occipital W (latter is high-lighted with cream); prominent brown postocular (supratym- panic) stripe; 2-4 brown labial bars; can- thal stripe not evident; flanks suffused with brown, becoming more pale (dusky cream ) ventrally; limbs gray-brown with indistinct brown bars, if evident they LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 43 are relatively broad (about equal to in- terspaces) and transversely oriented; in some specimens (especially AMNH 17592) oblique barring of the flanks is evident. Venter dirty cream with some diffuse brown spotting on throat; edge of lip more densely brown giving im- pression of incomplete barring; under magnification all ventral surfaces are peppered with brown. Anterior and posterior thigh surfaces and ventral sur- face of shank rich brown; no anal tri- angle. Measurements of holotype in mm.— SVL 38.4; tibia 16.3; head width 13.4; head length 11.4; upper eyelid width 3.6; IOD 2.9; eye length 4.5; E-N 3.2. The holotype is a gravid female with large, yellow ovarian eggs and exten- sively convoluted oviducts. Etymology.—Latin, meaning rough, in reference to the skin texture. Natural history—G. H. Tate found these frogs beneath stones in pasture- land and scrubby growth; he noted the area had “very little forest.”. Two small females are not mature. AMNH 17602 is a juvenile female (21.1 mm SVL); AMNH 17597 (29.0 mm SVL) has mod- erate oviducal convolution and small eggs. Remarks.—The absence of auditory and vocal apparati in FE. ruidus suggest it is related to the E. surdus Artenkreis but I think the available data support an argument of relationship with E. balio- notus and E. riveti. The three are dicho- patrically distributed in the Andes of southern Ecuador. The three may repre- sent the southern vicariants of the more northern E. orcesi and E. thymelensis. Distribution Known only from the type-locality on the Pacific slopes of the Cordillera Occidental (2317 m). Eleutherodactylus spinosus new species Fig. 19 Holotype-—USNM 199891, an adult female obtained at Sapote, Provincia Morona-Santiago, Ecuador, 2470 m, by Peter Spoecker on 23 August 1962. Paratypes.—(all Provincia Morona- Santiago, Ecuador) USNM 199916, be- tween Cerro Negro and Pailas, 2652 m; USNM 199917, between Cerro Negro and Pailas, 2439-2561 m; USNM 199945- 66, El Cruzado, 2195 m; USNM 199967- 71, Loma de Puerco, 2226 m; USNM 199918-35, Pailas, 2195 m; USNM 199936-44, San Juan Bosco, 2195 m; USNM 199972, San Vicente, 2835 m; USNM 199892-915, Sapote, 2470 m. Fic. 19.—Eleutherodactylus spinosus, USNM 199950, line equals 10 mm. 44 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Diagnosis.—(1) skin of dorsum fine- ly tuberculate, that of venter coarsely areolate; no dorsolateral folds; (2) tym- panum prominent, its length 4-4 that of eye in males, %-% in females; (3) snout subacuminate in dorsal view, truncate in lateral profile, short; (4) interorbital space broader than upper eyelid; low cranial crests present; upper eyelid bear- ing 2-3 elongate tubercles; (5) vomerine odontophores oval to triangular in out- line; (6) males lacking vocal sac and slits; (7) first finger shorter than second; digits bearing broad discs on dilated pads; (8) fingers bearing narrow lateral fringes; (9) ulnar tubercles subconical; (10) heel and outer edge of tarsus bear- ing subconical tubercles; (11) two meta- tarsal tubercles, inner oval, 5-6 times size of round outer; many supernumer- ary plantar tubercles; (12) toes bearing lateral fringes; toe pads as large as those of fingers; (13) brown above with in- definite darker blotching; flanks paler with brown bars; groin and concealed surface of hind leg black enclosing white spots; venter dull cream with heavy brown reticulation; some specimens have pale cream lines (reticulation) on flanks; (14) adults moderate-sized, males 16.1-25.0 mm, females 28.3-34.5 mm SVL. Eleutherodactylus spinosus is prob- ably most closely allied to E. nigrogris- eus (Andersson). The two are readily distinguished in that E. nigrogriseus is smaller ( ¢ 2 19.3-26.0 mm, ? @ 28.5- 29.4 mm SVL), does not have prominent tubercles along the outer edge of the tarsus, has shagreened skin with scat- tered tubercles on the dorsum (not fine- ly tuberculate ), lacks enlarged tubercles on the eyelids, and has yellow spots (in life) on the posterior surface of the thighs. Description—Head narrower than body, broader than long; head width of males 37.2-41.5 percent SVL (x = 38.6, N = 19), of females 36.7-41.6 percent (x = 39.7, N = 21); snout subacuminate to subovoid in dorsal view, nearly trun- cate in lateral profile; snout short, E-N of males 66.7-86.2 percent eye length (x = 75.8, N = 19), of females 77.8- 97.1 percent (x = 86.4, N = 21); nos- trils weakly protuberant, directed dorso- laterally; canthus rostralis moderately sharp, concave; loreal region concave, sloping to lips; lips not flared in most specimens, weakly flared in large adult females; interorbital space nearly flat, frontoparietals weakly upturned, low cranial crests most evident in large fe- males; upper eyelid width 87.0-129.4 percent IOD in males (x = 104.6, N = 17), 81.2-103.1 percent in females (x = 94.8, N = 20); upper eyelid tuberculate, bearing 2-3 long, conical tubercles; temporal region sloping; supratympanic fold heavy, concealing upper edge of tympanum; tympanum prominent, round to higher than long, its length in males 17.2-27.6 percent eye length (x = 22.8, N = 19), in females 20.5-37.2 percent (xX = 28.3, N = 21); separated from eye by 1% times its length; choanae mod- erate-sized, round, not concealed by palatal shelf of maxillary arch; vomerine odontophores median and posterior to choanae, oval to triangular in outline, bearing 4-5 teeth in a transverse row on posterior edge of odontophore; odonto- phores 2-3 times size of a choana, nar- rowly separated (about one choanal width); tongue longer than wide, pos- terior % not adherent to floor of mouth, posterior edge not notched; males lack- ing vocal sac and slits. Skin of dorsum finely tuberculate, that of head and anterior back least so, heavy ridge from eye across scapular region; no dorsolateral folds; skin of venter coarsely areolate, discoidal folds prominent; no anal sheath; ulnar tuber- cles prominent, subconical; palmar tu- bercle bifid, larger than oval thenar tubercle; many supernumerary palmar tubercles, all smaller and less prominent than round, elevated, non-conical sub- articular tubercles; fingers bearing nar- row lateral fringes; fingers bearing discs (broader than long) on dilated, apically rounded pads; pad on thumb smallest, those on fingers II-IV as large as tym- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 45 panum; thumb shorter than second finger. Heel and outer edge of tarsus bear- ing large subconical tubercles; inner tarsal fold along distal 4-% of tarsus; in- ner metatarsal tubercle oval, twice as long as wide, non-compressed, 5-6 times size of round outer metatarsal tubercle; many supernumerary plantar tubercles, all smaller than non-conical subarticular tubercles; subarticular tubercles of toes smaller than those of fingers, slightly longer than wide; toes bearing lateral fringes; pads and discs of toes as large as those of outer fingers; heels of flexed hind legs touch; heel of adpressed hind limb reaches to eye; shank of males 50.2- 54.8 percent SVL (x = 52.4, N = 19), of females 47.8-55.4 percent (X = 51.3, N = 21). In preservative light to dark brown with black labial bars. In dark indi- viduals no dorsal pattern is evident; in more pale individuals, the dorsum is marbled (Fig. 19); the limb bars are narrow and oblique, and slanted bars are evident on the flanks. Innermost digits cream. Flanks and concealed sur- faces of limbs are black with cream spots (especially evident in axillae, groin, con- cealed thigh, and shank). Posterior thigh bears small cream flecks in males and small females, large cream spots in larger females. The venter is dull creamy-yel- low heavily spotted and reticulated with brown; the throat is generally darker than the venter. Twenty-six of the 98 specimens examined have pale cream reticulation on the flanks (Fig. 19). In life, E. spinosus is pale to dark brown with brown chevrons on_ back, scapular W, and limb bars. The venter is dull cream heavily reticulated with brown, to black with a gray reticulation. The posterior thigh, concealed shank, anterior thigh, groin, and axilla are black enclosing white spots. The flanks are dark brown reticulated with dull yellow. Measurements of holotype in mm.— SVL 34.3; tibia 17.2; head width 12.6; upper eyelid width 3.3; IOD 3.3; tym- panum length 1.2; eye length 4.2; E-N 3.6. The holotype is a gravid female with extensively convoluted oviducts and large, yellow ovarian eggs. This individual lacks the pale reticulation on the upper flanks. Etymology.—Latin, in reference to the tuberculate skin of the dorsum. Natural history.—Peters and his asso- ciates found specimens beneath rocks along trails and on elephant ear plants at night. The largest non-gravid females are 26.2-28.5 mm SVL; these females do not have extensively convoluted oviducts or large eggs. The largest juvenile fe- male (straight oviducts, small eggs) is USNM 199946 (25.7 mm SVL). Remarks.—Eleutherodactylus — spino- sus is currently known only from the eastern slopes of the Cordillera de Ma- tanga (where it is evidently most abun- dant between 2200 and 2500 m) and on the adjacent Cordillera del Condor. Lynch (1969b) recorded E. nigrogriseus from the eastern slopes of the Andes in Napo, Pastaza, and Tungurahua_proy- inces at elevations of 1150-1800 m. Pe- ters and his associates collected 16 speci- mens of E. nigrogriseus sympatrically with E. spinosus. Their specimens were taken at elevations between 2195 and 2835 m; most were found at night on vegetation in the montane forests. Eleutherodactylus cryptomelas ap- pears related to E. nigrogriseus and E. spinosus. Although fewer data are avail- able for E. cryptomelas, this species appears to occur at slightly higher ele- vations than does either of the other species and is more southern in its dis- tribution. Distribution —Known from the Ama- zonian slopes of the Cordillera de Ma- tanga at elevations of 1707-2835 m. Eleutherodactylus versicolor new species Fig. 13C Holotype-—KU_ 119858, an adult male obtained “15 km E Loja, 2800 m” (= 13.5 km E. Loja, 2800 m), a locality just east of the crest on the mountain range dividing Loja and Zamora-Chin- 46 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY chipe provinces, Ecuador, on 10 June 1968 by Robert W. Henderson and J. D. Lynch. Paratypes—KU 119859-71 and 119911-44, topotypes collected on 10 June and 13 June 1968 by R. W. Hen- derson and J. D. Lynch. Diagnosis.—(1) skin of dorsum sha- greened and finely tuberculate, that of venter areolate; no dorsolateral folds; (2) tympanum visible, round, its length two-fifths to one-half that of eye; (3) snout subacuminate in dorsal view, rounded in lateral profile; E-N greater than or equal to eye length; (4) inter- orbital space flat, no cranial crests; up- per eyelid width less than IOD; (5) vo- merine teeth and odontophores present, odontophores teardrop-shaped and ob- lique; (6) males lacking vocal slits and vocal sac; (7) first finger shorter than second; all fingers bearing discs on dilated pads; dilation ratios I: 1.5, II: 1.9, TH: 2.1, IV: 2.0: (8) fingers. with very poorly-developed lateral fringes or none; (9) ulnar tubercles present, not prominent; (10) inner edge of tarsus bearing a short ridge-like tubercle, outer edge bearing a row of small conical tu- bercles; no heel tubercles; (11) two metatarsal tubercles, outer conical, round, one-fourth to one-third size of oval inner; supernumerary plantar tu- bercles numerous, conical; (12) toes not fringed, no basal webbing; all digits bearing discs on dilated pads, pads not as large as those of fingers; (13) ground color usually brown on dorsal surfaces blotched with brown (chevrons, inter- orbital bar, canthal and supratympanic stripes); flanks barred or spotted; limbs barred; venter cream reticulated with brown; posterior surface of thighs brown with cream flecks; (14) adults small, males 19.3-25.2 mm, females 22.7-29.8 mm SVL. Aside from coloration, E. versicolor is similar to E. cajamarcensis and E. uni- strigatus. E. versicolor differs from both of these species in lacking a vocal sac and slits. Both E. cajamarcensis and E. versicolor differ from E. unistrigatus in having flattened warts on the upper eye- lids and distinct supernumerary plantar tubercles. In life, the three are readily distinguished—E. versicolor has a green ground color and lacks red areas in the groin and on the concealed thighs, E. cajamarcensis is usually brown and has red areas on the concealed thigh and in the groin, and E. wnistrigatus is tan, cream, or rarely brown above and cream below and lacks bright colors on the concealed thigh surfaces and groin. Description—Head as wide as or slightly wider than body, as wide as or slightly wider than long; head width 35.0-40.6 percent SVL (x = 37.9, N = 33); snout subacuminate in dorsal view, rounded in lateral profile; canthus ros- tralis sharp, straight or slightly concave; loreal region concave, sloping to lip; lips not flared; nostrils weakly protuberant, near tip of snout, directed dorsolaterally; snout of moderate length, eye length equal to or slightly greater than E-N; upper eyelid width 75.8-113.7 percent IOD (x = 90.3, N = 33), eyelid slightly wider in males than in females; inter- orbital space flat, no cranial crests; no frontoparietal fontanelle; tympanum vis- ible, prominent, round, its upper and posterior edges partially concealed by thick, glandular supratympanic fold; tympanum 38.9-50.2 percent eye length in males (x = 43.8, N = 16), 40.4-55.0 percent in females (x = 46.2, N = 17); tongue thick, longer than wide, posterior one-third not adherent to floor of mouth, weakly notched posteriorly; choanae small, round, well within border of jaws when palate is viewed from directly above; vomerine odontophores slanted, median and posterior to choanae, sep- arated medially by distance about equal to twice width of an odontophore; each odontophore bearing 2-4 teeth in a clump at posterior end of process; males lacking vocal sac and slits. Skin or dorsum shagreened and finely tuberculate; no dorsolateral folds; skin on upper eyelids more coarsely tubercu- late than that on rest of dorsum; skin on flanks and upper surfaces of limbs sha- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 47 greened and finely tuberculate, that on venter coarsely areolate; skin below vent and on underside of thighs areolate, that on undersides of forelimbs and shank smooth; discoidal folds prominent. Forearm bearing small ulnar tuber- cles; two palmar tubercles, partially fused, median largest and equal in size to oval thenar tubercle; few supernu- merary thenar tubercles; subarticular tubercles prominent, round, not conical, simple; fingers lacking lateral fringes or having only keel-like lateral fringes; digits bearing discs on dilated pads, pads wider than long, not emarginate; dilation ratios I: 1.2-1.8 (x = 1.5), II: 1.7-2.3 (x = 1.9), III: 1.8-2.4 (x = 2.1), IV: 1.7-2.4 (x = 2.0); first finger shorter than second (Fig. 20). Hind limbs moderately long, shank 48.4-58.4 percent SVL (x = 55.0, N = 33); heel weakly tuberculate, none en- larged; outer edge of tarsus bearing series of small tubercles which continue along outer edge of foot to base of toe V; inner edge of tarsus bearing a short ridge-like tubercle; inner metatarsal tu- bercle oval, about one and_ one-half times as long as wide, not compressed, three to four times as large as rounded, conical, outer metatarsal tubercle; plan- tar surface bearing numerous conical supernumerary tubercles; subarticular tubercles of toes like those of fingers but slightly smaller; toes Jacking lateral fringes or having only indistinct ridges along the edge of toes; toes not webbed except between toes IV and V; toes bear- ing discs on dilated pads, pads broader than long, slightly smaller than those of fingers. In preservative, E. versicolor has a ground color of tan, gray, or, more often, shades of brown. The ground color of the body lightens on the flanks and thighs. The dorsum is blotched with brown to nearly black and the blotches are usually outlined with cream. The dark markings on the face consist of a distinct canthal stripe, which continues behind eye as a supratympanic stripe, and labial bars. The tympanum is paler than the rest of the head and is cream with a bronze wash. The dorsal blotches continue onto the flanks as slanted bars. The flank bars are broken up into rows of spots in some individuals. A dark interorbital bar is present in all but the striped individuals (see below) and is bordered anteriorly by a pale interorbi- tal bar or patch of cream to brown. The forelimbs are strongly barred with dark brown usually edged with cream or tan. The hind limbs are barred like the fore- limbs except that the ground color on the thigh is more pale than that on the shank or tarsus. Most individuals have two bars on the thigh and two on the shank as well as one on the knee. The dark bars are about as wide as or slight- ly wider than the pale interspaces. An anal triangle is present but is not as dark as the limb bars. The anterior and pos- terior thigh surfaces and to a lesser ex- tent, the top of the thighs, are mottled brown with cream flecking. Many in- dividuals (24.4%) have one to 55 distinct enamel-like white spots on the flanks (the spots are outlined in black). The venter is cream to pale brown with a dense brown or black reticulation. The dark flecking on the venter is sometimes accentuated on the throat—these indi- viduals have dark bars on the margin of the lower jaws. The variegated pattern (Fig. 13C) is found in 93% of the specimens. Seven percent have a striped pattern. The striped individuals differ in pattern from the variegated individuals only in the pattern on the back and flanks. Most striped individuals have dark flanks de- limited dorsally by a sharp edge to the almost black flank. Between the dark dorsolateral areas are dark stripes or rows of spots. These individuals lack interorbital bars. One-half of the striped individuals have white spots on the flanks (2-6 spots per individual). In life, E. versicolor is green (or rarely tan or brown) with brown, red- dish brown, or black markings bordered with cream. A few individuals have a reddish-orange patch on the center of 48 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY the back and a reddish-orange inter- orbital bar. The concealed surfaces of the limbs and flanks are brown to black with reddish-pink flecks. The venter is cream to pale reddish-pink with black or brown reticulations. The iris is bronze-white with a reddish-brown hori- zontal streak and faint black reticula- tions. The enamel-like white spots of preserved individuals were dull white in life. Measurements of holotype in mm.— SVL 22.2, shank 12.4, head width 8.7, head length 8.5, upper eyelid 2.3, IOD 2.7, tympanum length 1.4, eye length 3.4. Etymology.—Greek, meaning varie- gated, in reference to the dominant color pattern. Natural history—No egg _ clutches were found but females with large eggs were collected at the same time equally large females with only small, white, Ovarian eggs were found. The speci- mens I collected between 10 and 16 June 1968 were found beneath stones and logs in sparsely wooded pasture or in cloud forest habitats at elevations be- tween 2750 and 3100 m. Based on lim- ited observations, it appears that E. ver- sicolor occurs at higher elevations than does E. cajamarcensis. Remarks.—Eleutherodactylus versi- color is considered a close relative of E. cajamarcensis and E. unistrigatus. The similarities between the three species are extensive, and they differ in color pat- tern, skin texture, size of digital pads (Fig. 20), and proportions. The skulls of the three species are not appreciably different—all have moderate-sized nasal bones in median contact but separated from the frontoparietals, no frontopari- etal fontanelle, no cranial crests, the frontoparietals fused to the prootics, moderate-sized vomers that are not in median contact, long anterior rami of the parasphenoid (reaching palatines and vomers ), and median rami of the ptery- goids in contact with the parasphenoid alae. B (an Fic. 20.—Hands of (A) Eleutherodactylus unistrigatus, KU 120019, and (B) E. versicolor, KU 119919; line equals 5 mm. LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 49 Distribution. —Intermediate eleva- tions (2750-3100 m) in the Andes of southern Ecuador. Eleutherodactylus vidua new species Fig. 13D Holotype—KU 120082, adult female obtained 15 km E Cuidad Loja, Provin- cia Zamora-Chinchipe, Ecuador, 2800 m, on 10 June 1968 by John D. Lynch. Paratypes.—KU 120083-88, 120090- 91, topotypes; KU 141994, 15 km E Loja, Provincia Zamora-Chinchipe, Ecuador, 2710 m; KU 120089, 13-14 km E Loja, Provincia Loja, Ecuador, 2850 m; KU 120092, 8-9 km N San Lucas, Provincia Loja, Ecuador, 3000-3100 m. Diagnosis.—(1) skin of dorsum fine- ly shagreened with ill-defined dorsolat- eral folds, that of venter areolate; (2) tympanum concealed beneath skin; (3) snout broadly rounded (obtuse) in dor- sal view, rounded in lateral profile, short; (4) interorbital space flat (no cranial ridges), wider than upper eyelid width; (5) vomerine teeth and dentigerous processes present; (6) males unknown; (7) first finger shorter than second, digits bearing narrow pads, dilation ratios 1.26-1.31 for outer fingers; (8) fingers lacking lateral fringes; (9) no ulnar tubercles or fold; (10) tarsus bearing ill-defined tubercle along inner edge, series of obscure tubercles along outer edge; no heel tubercles; (11) two metatarsal tubercles, inner slightly larger than outer; plantar surface bearing nu- merous supernumerary tubercles; (12) toes lacking lateral fringes and basal webbing; (13) no bright spots in axilla, groin, concealed limb surfaces, or on venter; venter cream with brown fleck- ing; dorsum tan to brown, mottled with darker brown or bearing cream dorso- lateral stripes; (14) females 18.0-23.1 mm SVL. Eleutherodactylus vidua is most sim- ilar to four other narrow-disced Andean Eleutherodactylus—E. ginesi, E. myersi, E. nicefori, and E. trepidotus—E. vidua differs from all of these in having the tympanum concealed and differs from E. myersi and E. trepidotus in lacking spots in the groin and on the venter. Vomerine teeth are not visible in E. my- ersi or E. trepidotus, but are visible in the other three species. E. ginesi lacks dorsolateral folds and is uniform black. Eleutherodactylus nicefori is a somewhat larger frog but like E. vidua is darker above and cream, flecked or reticulated with brown, below. Description—Head narrower than body, slightly wider than long; head width 36.2-39.9 percent SVL (x = 38.7, N = 8); snout obtuse in dorsal view, rounded and slightly sloping in lateral profile; snout short, eye length greater than E-N; canthus rostralis sharp, weak- ly concave; loreal region concave, slop- ing abruptly to non-flared lips; nostrils weakly protuberant, directed laterally, near tip of snout; upper eyelid width 65.6-81.4 percent IOD (x = 73.5, N = 8); interorbital space flat, no cranial crests present; frontoparietal bones in median contact for their entire lengths, no fontanelle; tympanum present, con- cealed beneath skin; tongue large, round, fleshy, not notched posteriorly, posterior one-third free; choanae small, round, completely visible when roof of mouth is viewed from directly above; dentigerous processes of vomerine bones present, round in outline, as large as or slightly larger than a choana, situated between and posterior to choanae, each bearing 2-4 teeth in a transverse row. Skin of dorsum finely shagreened with poorly defined dorsolateral folds and flattened warts on flanks; small scat- tered tubercles are present on the dor- sum and eyelids; skin of throat and ven- ter areolate, discoidal folds prominent; ventral surfaces of limbs smooth except on ventral face of thighs; forearm lack- ing ulnar tubercles; two palmar tuber- cles, inner largest, larger than oval thenar tubercle; palm bearing indefinite supernumerary tubercles; subarticular tubercles of fingers large, round, flat- tened, simple; fingers not bearing lateral fringes; digit tips bearing discs, weakly dilated, pads on outer two fingers slight- 50 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ly larger than those of inner fingers (Fig. 7C) and broader than long; dilation ra- tios I: 10 (x = 1.0), II: 10-12 (x = 1) Au 2-6 a4) PV 2-14 (x = 1.3); first finger shorter than sec- ond. Hind limbs short, shank 37.8-46.4 per- cent SVL (x = 42.3, N = 8); inner edge of tarsus bearing short, ill-defined tu- bercle separated from inner metatarsal tubercle; outer edge of tarsus bearing a series of weakly defined tubercles; no tubercles on heel; two metatarsal tuber- cles, inner about twice as long as wide, not compressed, one and one-half to two times size of rounded outer metatarsal tubercle; plantar surfaces bearing nu- merous poorly defined supernumerary tubercles; subarticular tubercles of toes smaller than those of fingers, round, flat- tened, simple; toes lacking lateral fringes and basal webbing; basal one-third of fifth toe fused to fourth toe; tips of digits weakly expanded, pads slightly broader than long, bearing discs. In preservative, the dorsum and limbs are gray to dark brown with a red- dish cast. Seven of the twelve individ- uals have prominent dorsolateral stripes. Of the five non-striped individuals, two are reddish-brown with very faint darker brown markings except on the limbs where brown bars are distinct. The other three non-striped individuals have extensive dark brown or blackish mar- bling on the dorsum and flanks; the ground color in these specimens is brown or gray-brown. In the seven striped individuals, the cream or white stripe begins at the tip of the snout and runs posteriorly along the canthus, on the outer edge of the upper eyelid, and down the body terminating above the thighs. The light stripe is edged by wider brown to black stripes (Fig. 13D). The center of the back is not spotted with dark brown or black. Two of the striped individuals are gray—the other five are brown to rust-brown. The limbs are barred with brown. All individuals have a dark brown to black anal patch edged laterally with cream or white stripes. The posterior surface of the thigh is dull brown with loosely-defined cream spots. The venter is cream with minute brown flecking. The throat differs from the rest of the venter in having more dense flecking (especially anteriorly). The lips are barred and a prominent dark brown or black canthal and supratympanic stripe is present; this stripe is usually outlined with cream. In life, E. vidua is tan to brown with brown markings. If dorsolateral stripes are present, they are cream. The tan individuals have a faint green wash. The venter is cream, dusky gray, or light brown. The posterior surfaces of the thighs are flesh colored to pale rose. The iris is gray with a horizontal brown streak and fine black reticulations. Wil- liam E. Duellman described the iris as “pale green above, reddish bronze _ be- low” (KU 141994, notes 21 July 1971). Measurements of the holotype in mm.—SVL 21.8, shank 9.0, head width 8.7, head length 7.9, upper eyelid 2.1, IOD 2.6, eye length 2.4. The holotype is an adult female with heavily convo- luted oviducts and large (1.8-2.4 mm in diameter), yellow, ovarian eggs. Its color pattern is of the striped morph. Etymology.—Latin, a widow, used as a noun in apposition. The choice of this name reflects my belief that this species is all-female, a notion requiring careful checking. Natural history—All but one indi- vidual (KU 120092) were collected be- neath rocks or vegetation on the Cor- dillera de Zamora, east of Loja, at eleva- tions between 2710 and 2800 m. When uncovered these small frogs would walk towards the nearest cover or crouch— none was seen to hop. No direct evi- dence of reproduction was discovered but one adult female (KU 120092) con- tained only small ovarian eggs whereas the other nine adult females contained large ovarian eggs and were presumably ready to deposit the eggs. A total of fifteen specimens has been obtained and each is a female. This LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 51 small sample is not adequate to assert that the species is all-female in view of two other cases of frogs of this genus in which males are rare. E. orestes exhibits a similar preponderance of females (one male: sixteen females) and E. trepidotus presents an even more pronounced bias (2 males: 81 females). In each of these cases the bias is clearly towards females and coupled with another case of ap- parent all-female species (Phrynopus peraccai), also from Andean Ecuador, stand in marked contrast to the other twenty to thirty species of high Andean Ecuadorian eleutherodactylines exhibit- ing nearly one to one sex ratios. Remarks.—Eleutherodactylus vidua has the least digital dilation of any An- dean Eleutherodactylus. The width of the digital pads of this species are com- parable with the bulbous digital pads seen in several species of the disc-less Andean genus Phrynopus. The skull of E. vidua (Fig. 11F-G) closely resembles that of E. orestes. The nasals are mod- erate-sized and separated, lying on a large sphenethmoid that extends ante- riorly nearly to the tip of the nasal bones. The frontoparietals are complete (no fontanelle) and are fused to the prootics. The zygomatic ramus of the squamosal is short (Fig. 12C). The vomers are moderate-sized and separated. The pal- atines are more normal in extent than those of E. orestes and E. colodactylus. The anterior ramus of the parasphenoid is comparatively elongate, approaching the condition seen in E. colodactylus (Fig. 8) and that seen in several Phry- nopus species (Lynch, 1975). The para- sphenoid alae are oriented at right an- gles to the anterior ramus and are nar- rowly separated from the median rami of the pterygoids. Distribution.—Eleutherodactylus vi- dua is known from the mountain crests east and north of Loja, Ecuador, at ele- vations of 2710 to 3100 m. Eleutherodactylus w-nigrum (Boettger) The southern extent of the distribu- tion of E. w-nigrum is in southern Ecua- dor. The species is widely distributed at moderate and intermediate elevations on both Amazonian and Pacific versants in Ecuador and Colombia and also in- vades the valleys of the Rio Cauca and Rio Magdalena in Colombia. The collections by Peters and his as- sociates on the Amazonian versant of the Cordillera Oriental east of Cuenca establish an altitudinal range of 1890 to 2604 m for E. w-nigrum, but in northern Ecuador it occurs at elevations between 1200 and 3000 m. This species will be treated in detail in a review of the spe- cies of the genus found on the Amazo- nian slopes of the Andes in Ecuador (in prep., Lynch and Duellman). DISCUSSION Relationships of the southern fauna. —The genus Eleutherodactylus ceases to be a component of the high Andean frog fauna south of the area here termed “southern Andean Ecuador.” The twenty species living within this area belong to two of the species groups recognized by Lynch (1976a). Two are members of the fitzingeri group (E. lymani and E. w-nigrum) and are closely related with- in that group. The other eighteen are referable to the wnistrigatus group. The intragroup relationships of the group are poorly known, in part because the group is so large (more than 100 species now recognized). As now understood, the relationships of the 18 species in the unistrigatus group found in southern Andean Ecuador are as follows: 1. E. cajamarcensis and E. versicolor are southern vicariants of northern spe- cies (E. coeruleus, E. lehmanni, and E. unistrigatus) found in southern Colom- bia and northern Ecuador. 2. E. balionotus, E. riveti, and E. ruidus form a vicariant series and are possibly the southern elements of a northern series (E. orcesi and E. thy- 52 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY melensis, northern Ecuador) which may be upland replacements of the wnistri- gatus series. E. percultus and E. pycno- dermis are very tentatively associated with the balionotus-riveti-ruidus series. 3. E. baryecuus is the southern ele- ment of an Artenkreis also containing E. devillei (Amazonian slope, northern Ecuador) and E. surdus (Pacific slope, northern Ecuador). 4. E. cryophilius is the southern ele- ment of the curtipes Artenkreis (buck- leyi and curtipes) distributed on the Central Cordillera of Colombia south through northern Ecuador. 5. E. cryptomelas and E. spinosus appear related to one another and to an Amazonian versant species, E. nigrogri- seus, which ranges northward through Ecuador; E. atratus may be allied to this assemblage. 6. E. phoxocephalus is apparently the southern member of a series of poorly known Pacific versant frogs in Colombia and Ecuador. 7. E. bromeliaceus is an upland rela- tive of E. lacrimosus which is found in Amazonian Brasil, Colombia, Ecuador, and Peru. 8. E. colodactylus may be related to E. proserpens and E. celator; the latter occurs on the Pacific versant in northern Ecuador. 9. E. orestes and E. vidua seem close- ly related and moderately well-separated from other species of the group. Their resemblance to E. ginesi (Venezuela), E. myersi and E. nicefori (Colombia), and E. trepidotus (northern Ecuador ) may be a reflection of relationship. Ecological segregation.—As a whole, the southern Andean fauna has affinities with northern Andean species although a northern connection is tenuous for the balionotus-riveti series and the orestes- vidua series. Only one species of the unistrigatus group, E. bromeliaceus, has obvious Amazonian affinities. Eleu- therodactylus lymani is related to a wide-spread Andean slopes species (E. w-nigrum) ranging through Colombia and Ecuador; as a whole, the fitzingeri group is a lowland (< 1000 m) group, but endemic, moderate elevation species are also present on the Cordillera Occi- dental and Sierra Nevada de Santa Mar- ta in Colombia, the Talamanca Range in lower Central America, and the Sierra Madre del Sur in nuclear Central Amer- ica. The twenty species may be assigned to ecogeographic units as follows: Pacific slope (cloud forest, 2000-3000 m): E. phoxocephalus and E. ruidus. Intercordilleran belt (relatively xe- ric, < 2600 m): E. cajamarcensis and E. lymani. Pdramo (> 3000 m, cool, moist): E. cryophilius and E. riveti. Subparamo (Amazonian _ slopes, 2800-3000 m): E. balionotus, E. orestes, E. percultus, E. pycnodermis, E. versi- color, and E. vidua. Amazonian slope (cloud forest, 1800- 2800 m): E. atratus, E. baryecuus, E. bromeliaceus, E. colodactylus, E. crypt- omelas, E. proserpens, E. spinosus, and E. w-nigrum. The ecogeographic contingencies readi- ly support the conclusion of a more di- verse Amazonian slope fauna. The rela- tive depauperity of the Pacific slopes is probably a reflection of the aridity im- posed by the proximity of the cold Humboldt current and partly a function of the pronounced absence of high alti- tude habitat on the western face of the Andes in southern Ecuador. With the exception of E. bromelia- ceus cited above, all of the southern Andean species appear related to north- ern species found on the higher slopes and in the intercordilleran belt (cloud forests, subparamos or pajonales, and paramos). The distributional and phy- logenetic data are suggestive of disper- sal or vicariance within altitudinal zones rather than between them and are there- fore consistent with Janzen’s (1967) general hypothesis, and concordant with the conclusions of Peters (1973) for Atelopus, that the most pronounced bar- riers are those of altitude (and the asso- ciated climates). LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 53 Correlates with species densities and diversities—Some of the greater rich- ness of species among eleutherodacty- lines of southern Andean Ecuador com- pared to that of northern Andean Ecua- dor is real. If an equal-sized area of pajonales and pdramos in northern An- dean Ecuador is compared to one in southern Andean Ecuador, the latter clearly has more species (6-8 species compared to 2-3); however, the pajon- ales and pdramos of northern Ecuador are relatively contiguous whereas, in southern Ecuador, these habitats are patchy. Thus, part of the greater rich- ness is a function of a continental islands or archepelagar paradigm (topographic relief of Simpson, 1964). Additionally, the Andes in southern Ecuador are less high than are those in northern Ecuador allowing slope species to occur on the subpdramo patches on the relatively low (ca. 2800 m) crests and peaks in south- ern Ecuador (viz., Abra de Zamora in Provincia Loja). Lastly, the pajonales and paramos of northern Andean Ecua- dor were subjected to extensive Pleisto- cene vulcanism (Sauer, 1957); such per- turbation would tend to reduce species diversity by rendering the habitat more uniform. North of the “faunal break” (between Guamote and Cafiar) noted by Lynch (1971, 1972a, b), one finds two wide-spread species, E. curtipes and E. unistrigatus, and ten other species of restricted distributions over an area some 8-10 times the size of southern Andean Ecuador. Species diversity computations re- quire abundance/species data. I have used data extracted from my own field- work in southern Ecuador to approxi- mate relative abundances even though collecting was seldom random. Field- work in 1968 and 1970 was designed to reveal what species were found in southern Ecuador and because most spe- cies were undescribed, selectivity in col- lection was minimized. Biases of pre- ferred microhabitat (of the animals and of the collectors) and of collecting schedules vs activity cycles of the or- ganisms are inescapable. However, to await an ecologist venturing into this area before anyone discusses relative abundance or species diversities is un- realistic; anecdotal data have some ap- plicability. In computing species den- sity values, I employed MacArthur's (1972) formula D, = 1 / & p?. The sim- plicity of computations and MacArthur’s assertion that the formula is more ap- propriate in community studies were the only reasons this formula was used in- stead of the more frequently used Shan- non-Weaver index. Species densities for 50 Andean lo- calities in Ecuador show that northern, and thus more often, higher elevation, sites exhibit lower densities than do the southern sites (Table 2). One could argue that there is a latitudinal (1°N to 3°S) effect but the altitudinal effect seems to explain the variation because lower elevation sites in northern Ecua- dor (on the Andean versants) have high species densities (e.g., Tandapi, Provin- cia Pichincha, 1450 m, 8 species; Pilald, Provincia Cotopaxi, 2500 m, 7 species). Species density is negatively correlated with altitude (r = —0.560, N — 2 = 48, Pe OOM): Species diversity is also negatively correlated with altitude (r = —0.524, N — 2 = 48, p < 0.01) as one might expect in light of the highly significant correlation between species density and altitude. I do not have D, values for any lowland sites but two Pacific slope localities have D, values comparable to those for the subparamo and cloud for- est localities in southern Andean Ecua- dor (Tandapi, D, = 2.6; Pilal6, D, = 4.6). The low species densities and di- versities at high Andean sites are not consistent with the area-diversity curves advanced by Simpson (1974) for An- dean floras or by Vuilleumeir (1970) for Andean birds. Lynch (1976b) suggested that re- source partitioning by the eight species of Eleutherodactylus found at a Pacific slope cloud forest locality (Tandapi) in central Ecuador might be mediated by 54 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY TABLE 2. Eleutherodactyline frog faunules (Eleutherodactylus and Phrynopus). Locality Altitude (m) St N? ID PE Carchi: Tulcan 3000 1 25 1.00 Volcan Chiles 3500 3 24 PRON / Paramo del Angél 3700 2 61 1.34 10-13 km SSE Tulcan 3020 3 37 2.59 Atal 2700 il 89 1.00 Imbabura: 1 km N Otavalo 2560 2} 13 1.16 Urcusiqui 3300 2 25 1.68 8 km NW Otavalo 3100 1 8 1.00 4 km W San Pablo 3050 1 44 1.00 Nudo de Mojanda 3400 3 9 2.79 Nudo de Mojanda 3440 ik 25 1.00 Nudo de Mojanda 3680 i to 1.00 Pichincha: % km N Cayambe 2820 il 47 1.00 Quito 2860 1 103 1.00 Volcan Pichincha 3700 1 37 1.00 6-7 km W Chillogallo 3000 3 29 1.78 Napo: E slope, Paso de Guamani 3650 2 94 1.07 Laguna de Papallacta 3350 4 163 1.39 Cotopaxi: Nudo de Tiopullo 3490 2 80 1.02 Mulalé 3000 1 25 1.00 Paramo de Milin 3900 1 34 1.00 Paramo del Apagua 3800 1 25 1.00 Tungurahua: 10 km SW Mocha 3700 2 29 Til 3 km SSW San Miguelito 2620 1 95 1.00 10 km W Cotaldé 3300 2 41 1.40 Chimborazo: 12 km SW Sta. Rosa 3400 1 25 1.00 18-20 km SW Sta. Rosa 3650 1 22 1.00 Urbina 3610 2 46 1.14 10 km W San Juan 3400 1 60 1.00 12 km SW Cajabamba 3800 1 24 1.00 Bolivar: crest E of Guaranda 3700 2 Be 1.04 SW slope Nevado Chimborazo 3800 9) 2 1232 [Faunal break noted by Lynch (1971, 1972a, b)] Canar: 8 km NW Biblian 3420 I 64 1.00 Azuay: Laguna de Zurucuchu 3200 2 47 1.04 Paramo de Matanga 3400 i Bill 1.00 8 km SSE Sevilla de Oro 3350 4 23 2.08 Cerro Negro 2930 3 ui 1.81 6 km W San Vicente 3110 3 31 2.28 3 km W San Vicente 2990 3 23 1.19 San Vicente 2835 8 78 2.44 3 km E Sevilla de Oro 2700 3 16 1.91 Suro Rancho 2700 4 59 2.32 bet. Sapote and Suro Rancho 2615 6 20 4.00 Sapote 2470 8 121 5.73 El Cruzado 2220 9 52 3.91 Pailas 2200 6 73 3.02 San Juan Bosco 2200 5 Al 2.66 Plan de Milagro 1700 3 8 2.46 Loja: 10 km S Saraguro 3120 5 26 2.41 Abra de Zamora 2800 10 296 2.27 1S = number of species 2N = number of individuals 3D, = 1/2p*, MacArthur’s (1972) diversity formula en LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 55 sexual dimorphism and narrower niches because interspecies g values were low (x = 1.14) compared to Hutchinson and MacArthur’s (1959) value of g = 1.28. Phi is a measure of the size difference necessary to avoid competition (size of larger/size of smaller) (Hutchinson and MacArthur, 1959). Crump (1974) noted marked sexual dimorphism and a “regu- lar” increase in sizes among the Eleu- therodactylus species found at an upper Amazonian rainforest site (Santa Ceci- lia, Provincia Napo, Ecuador). Using her data sets and augmenting them with my own, I find an even lower range of ¢ values for the Amazonian lowland fauna than that at Tandapi (Table 3). When the southern Andean fauna is broken into the four most obvious eco- geographic units, the more species-rich units on the Amazonian slope have ¢ values equally as low as that for the Amazon lowland fauna whereas those for the pdramo and the interandean-west slope faunules are more comparable to those of the Pacific lowlands and Tan- dapi. In all seven faunules, the degree of sexual dimorphism is identical (Table 3; sizes for the species found in southern Ecuador are provided in Table 4). In part, these low ¢g values are likely a function of pooling ecologic contingen- cies within localities (Lynch, 1976b, noted higher x ¢ for contingencies than for the entire faunule). However, the propriety of comparing x g values among TaBLe 3. Sexual dimorphism and ¢g values for seven Ecuadorian Eleutherodactylus faunules. Faunule S Dimorphism g Tandapi 8 Loja-W slope® 4 1.29-1.86(4); 1.45 Paramo” 3 1.25-1.51(3); 1.38 Ceja andina® 14 Bosque humédo* 10 Santa Cecilia i7/ Pacific lowlands 12 lrange (N); x + 2SE 120-1 °78(7 ): 1-38 == O15: 1.12-1:72(12); 1.39 += 0:09 1.18-1.72(8)); 1.39 = 0.14 11921-6515): 1239) == 0:07 1.21-1.81(10); 1.49 + 0.12 1.01-1.36(13); 1.14 + 0.06 1.06-1.55(6); 1.20 1.04-1.33(4); 1.21 1.01-1.29(22); 1.08 + 0.03 1.00-1.45(14); 1.09 + 0.04 1.00-1.17(27); 1.06 + 0.02 1.00-1.66(20); 1.14 + 0.07 * size data for species of these faunules are given in Table 4 TABLE 4. SVL of Eleutherodactylus in southern Andean Ecuador. Species rn) Le E. atratus 21.7 + 1.7(19): 27.4 + 1.3(10) E. balionotus 22.0(2) 28.1 + 0.6(7) E. baryecuus 29.0(5) 40.8 + 1.3(8) E. bromeliaceus 20.8 + 0.9(14) 26.5(5) E. cajamarcensis 22 Ae Orla) 23:8 == 10a) E. colodactylus Lik == 0!5(A6) 20.8 + 0.8(33) E. cryophilius 33.3(5) 45.1(5) E. cryptomelas 29:2 (A) Ape ui lih eg igidiy( Ved), sea be E. galdi 21.0 + 1.7(8) 3ir7ee a(S) E. lymani e249) 58.9 + 4.7(6) E. nigrogriseus 23.2 + 0.9(15) eT to E. orestes 19.8(1) 22.1 + 1.4(13) E. percultus 29.8(1) 38.2(1) E. phoxocephalus 27.2 = 0:7(29) Soo eeylel (29) E. proserpens 1867 == 10a) 22.0(4) E. pycnodermis 25.1 = 1.4(30) 38.0 + 1.1(40) E. riveti 24.1 + 0.6(35) 30.1 + 0.8(30) E. ruidus XAG) Se MAM 7/) 37.9 + 0.7(7) E. spinosus 20:1 == '0)6(34) 31.8 + 0.6(29) E. versicolor 23.0 + 0.8(16) 28.4 + 0.7(15) EUG eee SP leh) a 21.5 + 0.8(8) E. w-nigrum? 34.5 + 1.0(43) 59.2 + 2.6(9) ig + 2 SE(N) ? based on samples from the Amazonian slope in Provincia Morona-Santiago, Ecuador 56 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY several faunules is questionable in that the several faunules differ in the relative proportions of species of the fitzingeri and the unistrigatus groups. If g and the size of the larger of the two species are compared (Fig. 21), the two are significantly correlated (r = 0.70, N — 2 = 49, p < 0.01). This sample consists of 51 pair of species (using only fe- males). The correlation of ¢ values and size where only members of the fitzingeri group are involved is also highly signifi- cant (r = 0.83, N — 2 = 10, p < 0.01) whereas that where only members of the unistrigatus group are involved is not significant (r = 0.40, N — 2 = 37,p > 0.01, < 0.05). The correlation for uni- strigatus group species is significant at p = 0.05. Thus one might expect higher x g values in faunules having greater proportions of fitzingeri group species. However, using only seven faunules that is not true (Fig. 22); the correlation be- tween x g and the percent fitzingeri group species is non-significant (r = OAT IN = 2) 5.9 0/05). There isa highly significant negative correlation between x g and species density (Fig. 23) r= —=0:928. No) 2e-—" 5, pir 0.01). This is not a function of altitude (the correlation of x ¢ with altitude is not significant, r = 0.415, p > 0.05). Our knowledge of Andean eleutherodac- tylines is too fragmentary to decide if the approach to unity of ¢ values means smaller niches or increased niche overlap (the alternatives mentioned by Pianka, 1966) but the habitat segregation by the species in a Pacific slope cloud forest (Lynch, 1976b) suggests that the frogs have smaller niches. Comparisons with distributions of other Andean genera—The high An- dean frog fauna in northern Ecuador consists of Atelopus ignescens, Eleu- therodactylus curtipes and E. unistriga- tus, Gastrotheca riobambae, and Telma- tobius niger. Atelopus ignescens ranges south to at least the Abra de Zamora without a geographic replacement. Gas- trotheca riobambae and T. niger range south into the Cuenca hoya (both to the we? a 1 ° ee o— a A\ 4 20 30 40 50 85 SVL of Larger ? Fic. 21.—Magnitude of size difference (¢) as a function of body size in Eleutherodactylus. (4) ¢@ values where one (or both) species is a member of the fitzingeri group; (@) ¢ values for pairs of the unistrigatus group. e 1.2 e ————eeE — === —— 10 30 50 Per cent fitzingeri Gr Fic. 22—Magnitude of average ¢ in seven Eleutherodactylus faunules as a function of proportion of the faunule made up of members of fitzingeri group. IX 1 Fic. 23.—Magnitude of average ¢ in seven Eleutherodactylus faunules as a function of species density (S). LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 57 Nudo de Portete in Azuay Prov.). South of the Nudo de Portete, G. riobambae is replaced by three allopatric species, G. loyana, G. monticola, and G. psychro- philia (Duellman, 1974), and T. niger by two allopatric species, T. vellardi and an undescribed species (Trueb, pers. comm.). Eleutherodactylus curtipes ranges south to the Desierto de Palmira and is replaced on the pdramos ringing the Cuenca hoya by E. cryophilius. Eleutherodactylus wunistrigatus ranges south to the headwaters of the Rio Chambo; south of the Nudo de Azuay it is replaced by E. riveti (on the ranges surrounding the Cuenca hoya), E. caja- marcensis and E. versicolor (on the Cor- dilleras Chilla, Sabanilla, and Zamora), E. ruidus (on the Cordillera de Molle- turo), E. pycnodermis (on the Cordil- lera de Condorcillo), and E. balionotus and E. percultus (on the Cordillera de Zamora). In southern Andean Ecuador, frogs of the genus Colostethus are prom- inent and in the Loja hoya one encoun- ters Bufo spinulosus. High altitude iguanids of the genus Stenocercus exhibit a distributional pat- tern congruent with that of Eleuthero- dactylus (Fritts, 1974). Stenocercus guentheri ranges over northern Andean Ecuador, S. festae and S. simonsi occur in the Cuenca and Saraguro hoya, S. humeralis and S. ornatus occur on the north and west edges of the Loja hoya, and S. carrioni and S. rhodomelas range over the xeric Pacific slopes (Fritts, 1974). A less congruent pattern is seen in the teiid genus Pholidobolus where species’ range limits correspond with the Cordillera de Igualata and the Pastaza Valley and, to a lesser extent, the Nudo de Azuay, where P. prefrontalis leaves interandean habitats and inhabits the Pacific slopes (Montanucci, 1973). In some respects, the species of Ate- lopus, Eleutherodactylus, Gastrotheca, and Telmatobius are exposed to the same general selective regimes and eco- geographic barriers, and one might an- ticipate concordant variation among the four genera. On the other hand, Eleu- therodactylus differs from the others in exhibiting direct development; intuitive- ly, they seem more sensitive to environ- mental variation. However, Scott (1976) implied that Eleutherodactylus, freed as they are of the uncertainty of water, might be more uniformly populated (than frogs having a tadpole stage) en- couraging freer gene flow (and _ less fragmentation of populations). The parapatric (rather than dichopatric) distributions of most Artenkreisen of Eleutherodactylus suggests that gene flow is inhibited by ecogeographic bar- riers. The high densities of Eleutherodac- tylus and the putative non-patchiness of their microhabitat (at least in the low- lands, Heyer and Breven, 1973) suggest that a parapatric speciation model func- tions because these frogs exhibit a very low vagility. The lack of parapatric spe- ciation among the frogs now grouped as A. ignescens may be a result of search- ing patterns by the female of an amplec- tant pair for a suitable ovoposition site (Peters, 1973). Similarly, the apparent lack of speciation by G. riobambae may result from the admixture of local popu- lations coming from vocal attraction by calling males and/or dumping of larvae in suitable Andean ponds by the female. The restrictions on reproductive sites for Atelopus and Gastrotheca presumably override the impact of slight ecogeo- graphic barriers whereas Eleutherodac- tylus are more sensitive to those barriers because they may deposit their eggs where they are living rather than need- ing to move to ovoposition sites. SUMMARY The frog fauna of the Andes of southern Ecuador consists of no fewer than 21 species of eleutherodactylines belonging to the genera Eleutherodac- tylus and Phrynopus. A single species of Phrynopus and twenty species of Eleu- therodactylus occur in the Andes south of the Nudo de Azuay and north of the 58 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Huancabamba depression in northern Pert. Of the eighteen representatives of the unistrigatus group of Eleutherodac- tylus found in the region, sixteen are named as new species. With a single exception, the affinities of the 20 species of Eleutherodactylus are with high altitude and moderate alti- tude species found in northern Ecuador and southern Colombia. Eleutherodac- tylus bromeliaceus is most similar to a species of the Amazon basin, E. lacri- mosus. The greater diversity of eleuthero- dactylines in the Andes of southern Ecuador as compared to northern Ecua- dor may be a function of 1) the Pleis- tocene vulcanism of northern Andean Ecuador, 2) habitat contiguity in north- ern Andean Ecuador, or 3) the invasion of swbparamo habitats in southern Ecua- dor by species found on the Andean slopes at more northern latitudes. In general, the Andean species of Eleutherodactylus are parapatrically dis- tributed, and the distribution areas of the southern species are smaller than those of the largely sympatric represen- tatives of the genera Atelopus, Gastro- theca, and Telmatobius. These observa- tions suggest that Eleutherodactylus are less vagile and more sensitive to eco- geographic barriers than are the frogs of the genera Atelopus, Gastrotheca, and Telmatobius, all of which are Jarger than most Eleutherodactylus and have re- strictions on oviposition sites (all have aquatic tadpoles). RESUMEN La fauna anfibia de los Andes del sur del Ecuador esta representada al menos por 21 especies de eleutherodactilinos pertenecientes a los géneros Eleuthero- dactylus y Phrynopus. Entre el Nudo de Azuay, por el norte, y la Depresion de Huancabamba (norte del Pert), por el sur, se encuentra una sola especie de Phrynopus y 20 de Eleutherodactylus. Dieciseis especies del grupo wnistrigatus de Eleutherodactylus que se encuentran en esta regidn son descritas como nuevas. Con una excepcidén, todas éstas espe- cies de Eleutherodactylus muestran afin- idad con especies de moderada y de pronunciada elevacién del norte del Ecuador y del sur de Colombia. Solo hay una excepcién, Eleutherodactylus bromeliaceus, la cual es mas similar a una especie de la cuenca amazonica, E. lacrimosus. La mayor diversidad de eleuthero- dactilinos en los Andes Sud-ecuatori- anos, comparada con aquella del norte ecuatoriano, puede ser funcién de: 1) Vulcanismo pleistocénico de los Andes del norte del Ecuador, 2) continuidad ecolégico-ambiental en la regién norte de los Andes ecuatorianos, 6 3) la inva- sidn de los ambientes subparamicos en el sur del Ecuador por especies que se encuentran en las laderas andinas en latitudes mas nortinas. En general, las especies andinas de Eleutherodactylus se encuentran distri- duidas parapatricamente; las areas de distribucién de las especies surefas son mas pequefias que aquellas de los ge- neros Atelopus, Gastrotheca, y Telmato- bius que son ampliamente simpatricos. Estas observaciones sugieren que _ los Eleutherodactylus son menos vagiles y mas sensitivos a las barreras ecogeo- graficas que los sapos de los géneros Atelopus, Gastrotheca, y Telmatobius, los cuales en general son de mayor ta- mano y que ademas tienen restricciones en cuanto a los sitios de oviposicion por tener renacuajos acuaticos. LITERATURE CITED Acosta-SouLis, M. 1968. Divisiones fitogeo- graficas y formaciones geobotanicas del Ecuador. Imp. Casa Cultura Ecuat., Quito. 309 pp. Barpour, T. 1921. New Central American frogs. Proc. New England Zool. Club 10:25-31. Barsour, T., and Nosie, G. K. 1920. Some LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 59 amphibians from northwestern Peru, with a revision of the genera Phyllo- bates and Telmatobius. Bull. Mus. Comp. Zool. 63:395-427. BouLeNcER, G. A. 1882. Catalogue of the Batrachia Salientia s. Ecuadata in the collections of the British Museum. 2nd Ed. British Museum (Natural History), 503 pp. BouLencer, G. A. 1900. Descriptions of new batrachians and reptiles collected by Mr. P. O. Simons in Peru. Ann. Mag. Nat. Hist. (7)6:181-186. Cocuran, D. M., and Gorn, C. J. 1970. Frogs of Colombia. Bull. U.S. Nat. Mus. (288 ):1-655. Crump, M. L. 1974. Reproductive strategies in a tropical anuran community. Univ. Kansas Mus. Nat. Hist. Misc. Publs. (61) :1-68. Despax, M. 1911. Reptiles et batraciens de L’Equateur recueillis par M. le Dr. Rivet. Mission de ]Equateur (Arc de méridien équatorial) 9(2):18-43. DuUELLMAN, W. E. 1974. A systematic review of the marsupial frogs (Hylidae: Gas- trotheca) of the Andes of Ecuador. Occas. Pap. Mus. Nat. Hist. Univ. Kan- sas (22,):1-27. Fritts, T. H. 1974. A multivariate evolu- tionary analysis of the andean iguanid lizards of the genus Stenocercus. San Diego Soc. Nat. Hist. Memoir (7):1-89. Heyer, W. R., and Berven, K. A. 1973. Spe- cies diversities of herpetofaunal samples from similar microhabitats at two tropi- cal sites. Ecology 54:642-645. HutcuHinson, G. E., and MacArtuur, R. H. 1959. A theoretical ecological model of size distributions among species of ani- mals. Amer. Nat. 93:117-125. JANzEN, D. H. 1967. Why mountain passes are higher in the tropics. Amer. Nat. 101:233-249. Lyncu, J. D. 1964. A small collection of anu- ran amphibians from Panama, with the description of two new species of Eleu- therodactylus (Leptodactylidae). Journ. Ohio Herp. Soc. 4:65-68. Lyncu, J. D. 1968a. Two new frogs of the genus Eleutherodactylus from eastern Ecuador (Amphibia: Leptodactylidae). Journ. Herpetol. 2:129-135. Lyncu, J. D. 1968b. Systematic status of some andean leptodactylid frogs with a description of a new species of Eleu- therodactylus. Herpetologica 24:289- 300. Lyncu, J. D. 1969a. Taxonomic notes on Ec- uadorian frogs (Leptodactylidae: Eleu- therodactylus). Herpetologica 25:262- 274. Lyncu, J. D. 1969b. The identity of the frog, Pseudohyla_ nigrogrisea of Ecuador. Bull. So. California Acad. Sci. 68:219- 224, Lyncu, J. D. 1970a. Identity of two andean Eleutherodactylus with the description of a new species (Amphibia: Lepto- dactylidae). Journ. Herpetol. 3:135- 143. Lyncu, J. D. 1970b. A new eleutherodacty- line frog from Amazonian Ecuador. Proc. Biol. Soc. Washington 83:221-225. Lyncu, J. D. 1971. Redescriptions of three little-known Eleutherodactylus from northwestern Ecuador (Amphibia: Lep- todactylidae). Trans. Kansas Acad. Sci. 73:169-180. Lyncu, J. D. 1972a. Two new species of frogs (Eleutherodactylus: Leptodactylidae) from the paramos of northern Ecuador. Herpetologica 28:141-147. Lyncu, J. D. 1972b. Systematics and ecology of robber frogs in western Ecuador. Yearbook Amer. Philos. Soc. 1971:332- 333. Lyncu, J. D. 1974a. New species of frogs (Leptodactylidae: | Eleutherodactylus) from the Amazonian lowlands of Ecua- dor. Occas. Pap. Mus. Nat. Hist. Univ. Kansas (31) :1-22. Lyncu, J. D. 1974b. A new species of Eleu- therodactylus (Amphibia: Leptodacty- lidae) from the Pacific lowlands of Ecuador. Proc. Biol. Soc. Washington 87:381-387. Lyncn, J. D. 1975. A review of the andean leptodactylid frog genus Phrynopus. Occas. Pap. Mus. Nat. Hist. Univ. Kan- sas (35) :1-51. Lyncu, J. D. 1976a. The species groups of the South American frogs of the genus Eleutherodactylus _ (Leptodactylidae). Occas. Pap. Mus. Nat. Hist. Univ. Kan- sas (61):1-24. Lyncn, J. D. 1976b. New species of frogs (Leptodactylidae: | Eleutherodactylus) from the Pacific versant of Ecuador. Occas. Pap. Mus. Nat. Hist. Univ. Kan- Sasw (oe nd-ooe Lyncu, J. D., and ScHwarrz, A. 1972. Taxo- nomic disposition of some 19th century leptodactylid frog names. Journ. Herpe- tol. 5:103-114. MacArtuour, R. H. 1972. Geographical ecol- ogy. Patterns in the distribution of spe- cies. Harper and Row, New York. 269 Ppp. Monranuccr, R. R. 1973. Systematics and evolution of the andean lizard genus Pholidobolus (Sauria: Teiidae). Univ. Kansas Mus. Nat. Hist. Misc. Publs. (59): 1-52. ParKER, H. W. 1932. The systematic status of some frogs in the Vienna Museum. Ann. 60 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Mag. Nat. Hist. (10)10:341-344. Peters, J. A. 1973. The frog genus Atelopus in Ecuador (Anura: Bufonidae). Smith- sonian Contrib. Zool. (145) :1-49. Pranka, E. 1966. Latitudinal gradients in spe- cies diversity: a review of concepts. Amer. Nat. 100:33-46. Rutuven, A. G. 1922. The amphibians and reptiles of the Sierra Nevada de Santa Marta, Colombia. Univ. Michigan Mus. Zool. Misc. Publ. (8):1-69. Sauer, W. 1965. Geologia del Ecuador. Min- isterio de Educacién, Quito. 383 pp. SavaGE, J. M. 1965. A new bromeliad frog of the genus Eleutherodactylus from Costa Rica. Bull. So. California Acad. Sci. 64:106-110. Scorr, N. J. 1976. The abundance and diver- sity of the herpetofaunas of tropical for- est litter. Biotropica 8:41-58. Simpson, B. B. 1974. Glacial migrations of plants: island biogeographical — evi- dence. Science 185:698-700. Smpson, G. G. 1964. Species density of North American recent mammals. Syst. Zool. 13s 5- ta: Taytor, E. H. 1952. The frogs and toads of Costa Rica. Univ. Kansas Sci. Bull. 35: 577-942. VuILLEUMEIR, F. 1970. Insular biogeography in continental regions. In the northern Andes of South America. Amer. Nat. 104:373-378. APPENDIX: SPECIMENS EXAMINED Eleutherodactylus atratus (43 spec.) ECUADOR: Morona-Santiago: El] Cruza- do, 2195 m, USNM 199690-99; San Vicente, 2835 m, USNM_ 199712; Sapote, 2470 m, USNM_ 199701-10, 199713 (cleared and stained skeleton); 2 km W Sapote, 2561 m, USNM_ 199711; between Sapote and Suro Rancho, 2604-2622 m, USNM 199683-89; Suro Rancho, 2683 m, USNM 199675-82; % km W Suro Rancho, 2744 m, USNM 199700. Zamora- Chinchipe: Abra de Zamora, 2850 m, KU 165236-38. Eleutherodactylus balionotus (10 spec.) ECUADOR: Loja: 13.5 km E Loja, Abra de Zamora, 2800 m, KU 142135-44. Eleutherodactylus baryecuus (17 spec.) ECUADOR: Morona-Santiago: Cerro Ne- gro, 2927 m, USNM 199715; El Cruzado, 2195 m, USNM 199723-24; San Juan Bosco, 2195 m, USNM 199725; between San Juan Bosco and El Cruzado, 2226 m, USNM _ 199717-22; San Vicente, 2805-2835 m, USNM _ 199729; 3 km W San Vicente, 2988 m, USNM 199730; Sapote, 2470 m, USNM_ 199726-28; between Sevilla de Oro and Méndez (probably between the crest and Pailas), USNM 199716; Suro Rancho, 2683 m, USNM 199714. Eleutherodactylus bromeliaceus (20 spec.) ECUADOR: Morona-Santiago: El Cruza- do, 2195 m, USNM 199738-39; Mirador, 1982 m, USNM_ 199735; Pailas, 2195 m, USNM 199732-34, 199743-45; Plan de Milagro, 1707 m, USNM 199736-37, 199746-47; % km E Sa- pote, 2393 m, USNM 199740, 199748-49; 1 km E Sapote, 2332 m, USNM 199741; mountain above Sapote (to the south), 2500 m, USNM 199742, 199750; between Sapote and Suro Rancho, 2622 m, USNM 199731. Eleutherodactylus cajamarcensis (137 spec.) ECUADOR: Loja: 5 km NE Cariamanga, 1870 m, KU 141896-909; 12 km NE Catacocha, 2060 m, KU 141890-95; 9 km E Loja, 2660 m, KU 119949-50; 12% km S Loja, 2250 m, KU 165190-92: 6 km N San Lucas, 2760-2900 m, KU 120007-15, 165193-99; 7-8 km N San Lucas, 2940-3000 m, KU 119951-120005, 120006(5), 120022-24 (cleared and_ stained skeletons); 13 km E Veracruz, 2250 m, KU 141860-89. PERU: Cajamarca: pre-Inca ruins near Huambos, MCZ 5407. Piurd: summit of Cor- dillera between Chanchaque and Huancabam- ba, 3100 m, KU 135495, 135502. Eleutherodactylus colodactylus (113 spec. ) ECUADOR: Azuay: 8 km ESE Sevilla de Oro (on Azuay-Morona-Santiago frontier), 3140 m, USNM 198458. Morona-Santiago: E] Cruzado, 2195 m, USNM 198436; Pailas, 2195 m, USNM 198431-33, 198459; San Juan Bosco, 9195 m, USNM 198434-35, 198460-61; imme- diate environments of San Vicente, 2790-2820 m, USNM 198465-79; 1 km W San Vicente, 2851 m, USNM 198462-64; 1 km E Sapote, 2332 m, USNM 198457; mountain above Sa- pote to the south, 2500 m, USNM 198456; % km W Sapote, 2546 m, USNM_ 198437-55. Zamora-Chinchipe: Abra de Zamora, 2800 m, KU 142151-59 (142155 cleared and_ stained skeleton), 165219-21; 14 km E Loja, 2770 m, KU 142160-61; 15 km E Loja, 2710 m, KU 142162-64. PERU: Piurd: 33 km SW Huancabamba, 2745-3050 m, LSUMZ 32368-414. Eleutherodactylus cryophilius (10 spec.) ECUADOR: Azuay: Bestidn (southwest- LEPTODACTYLID FROGS OF THE GENUS ELEUTHERODACTYLUS 61 ern slopes Cerro Bestién), 3080 m, AMNH 13970; Laguna de Zurucuchu, 3200 m, KU 120091; between Sevilla de Oro and Azuay- Morona-Santiago frontier, 3354 m, USNM 199994. Morona-Santiago: on crest between Azuay and Morona-Santiago provinces (ap- proximately 8 km ESE Sevilla de Oro), 3384 m, USNM 200390; San Vicente, 2835 m, USNM 199995-96; 6 km W San Vicente, 3110 m, USNM 199993, 200391-93. Eleutherodactylus cryptomelas (8 spec.) ECUADOR: Loja: 8-9 km N San Lucas, 3000-3100 m, KU 120095-96. Morona-Santi- ago: Sapote, 2470 m, USNM 198480-82; 2 km W Sapote, 2560 m, USNM 198483. Zamora- Chinchipe: 15 km E Loja, 2710 m, KU 141992-93. Eleutherodactylus lymani (77 spec. ) ECUADOR: Azuay: 4 km SW Catavina, 1600 m, USNM JAP 3541; Rio Minas, 20 km W Santa Isabel, 1250 m, USNM JAP 3625; 55.4 km E Pasdaje, ca. 1000 m, KU 152009. El Oro: E) Chiral, 1630 m, AMNH 13961, 13964; Cordillera de Chilla, Llanos de Guavos, AMNH 13738; Guainche, AMNH 16256; Pinas, AMNH_ 16257; Portovelo, 610 m, AMNH 16334, 16339, 16341-44. Loja: Loja, 2150 m, BM(NH) 1931.2.12.1-3/RR 1947.2.15.99-101, KU 119502; 8 km NE Loja, 2300-2500 m, USNM GOV 8734-35, 8737, 9533-39, 9541, WCAB 39912-13; 2 km E Loja, 2210 m, KU 119504-12; 3 km E Loja, 2100 m, USNM JAP 2455; 7 km E Loja, 2500 m, KU 119503; 10 km W Loja City, 3000 m, USNM 98931; 7.6 km S Loja, 2210 m, KU 141962-64; 9 km S Loja, 2230 m, KU 165539-40, 165231 (lot of eggs); 12.2 km S Loja, Rio Malacatos valley, road to Vileabamba, 2275 m, KU 141292; 17 km NE Macara, 1240 m, KU 141965; Pta. Santa Ana, 1100 m, AMNH 13734, 13966, 13973; Rio Puyango, AMNH 16211-15, 16224. PERU: Cajamarca: Bellavista, MCZ 5423, 5426-27, 5429, 5431, UMMZ 55774(3); Pa- lamba, MCZ 5436; Perico, MCZ 5408, 5410, 5412, 5415, 5418-21, 5422. Eleutherodactylus orestes (20 spec. ) ECUADOR: Azuay: 2.1 km S Cutchil, 2720 m, KU 141468; 3.1 km S Cutchil, 2730 m, KU 141467; 5 km ESE Sevilla de Oro, 2957 m, USNM JAP 6580-81. Loja: Saraguro, 2570 m, KU 141995; 9.5 km S Saraguro, 3120 m, KU 141469-72, 151052 (cleared and stained skeleton); 10 km S Saraguro, 3100 m, KU 141996-97; 11 km NE Urdaneta, 2970 m, KU 141998-2003. Zamora-Chinchipe: Abra de Za- mora, 2850 m, KU 120094, 165550. Eleutherodactylus percultus (2 spec.) ECUADOR: Zamora-Chinchipe: Abra de Zamora, 2850 m, KU 166057-58. Eleutherodactylus phoxocephalus (193 spec. ) ECUADOR: Azuay: 10 km SW Victoria del Portete, Parque Nacional Portete de Tar- qui, 2700 m, KU 131281-82. Canar: 18.4 km NW El Tambo, 2960 m, KU 142118-30. Coto- paxi: Pilalé, 2320 m, KU 142075-103, USNM JAP 3208-11, 3213-14, 3329-36; Pilal6, 2460 m, KU 131404-78, 131485-88, 131698; Pilald, 2580 m, KU 131480-84, 131699-716. Loja: Saraguro, 2500 m, KU 135460-62; 3.3 km NNE Saraguro, 2400 m, KU 142117; 2 km S Sara- guro, 2680 m, KU 142114-16; Rio Zamora, 6.5 km N Loja, 2060 m, KU 142113. Pichincha: 4 km W Chiriboga, 2120 m, KU 142072-74; Finca Santa Lucia, 7.7 km E Chiriboga, 2120 m, KU 142063-71; Los Alpes, 2500 m, KU 140876-77; San Ignacio, 10 km E Tandapi, 2030 m, KU 109137. Zamora-Chinchipe: 15 km E Loja, 2710 m, KU 142104-12. Eleutherodactylus proserpens (27 spec.) ECUADOR: Morona-Santiago: FE] Cruza- do, 2195 m, USNM 199751; Plan de Milagro, 1707 m, USNM 198485, 198504; 1 km S Plan de Milagro, 1707 m, USNM 198503; Rio Piun- tza, Cordillera del Condor, 1830 m, KU 147044-46; % km E Sapote, 2393 m, USNM 199752; 1 km E Sapote, 2332 m, USNM 198499-501, 199753; mountain above Sapote (to the south), 2500 m, USNM 198496-98; % km W Sapote, 2546 m, USNM 198486-91; between Sapote and Suro Rancho, 2622 m, USNM 198484, 198492-95, 198502. Eleutherodactylus pycnodermis (137 spec. ) ECUADOR: Azuay: 3 km E Sevilla de Oro, 2713 m, USNM 199851-53; 5 km ESE Sevilla de Oro, 2957 m, USNM_ 199854, 199867; between Sevilla de Oro and Cerro Negro, 3110 m, USNM 199868; crest at Azuay- Morona-Santiago frontier, ca. 8 km SE Sevilla de Oro, 3354 m, USNM_ 199855-59, 199866; crest at Azuay-Morona-Santiago frontier, ca. 8 km SE Sevilla de Oro, 3384 m, USNM 199758- 59, 199860-65. Morona-Santiago: between Cerro Negro and Pailas, 2652 m, USNM 199755-57, 199869-71; San Vicente, 2805-35 m, USNM 199754, 199815-50, 199884, 199888- 90; San Vicente, 2851 m, USNM 199813-14, 199885-87; 3 km W San Vicente, 2988 m, USNM 199794-812, 199882-83; 6 km W San Vicente, 3100 m, USNM 199787-93, 199880- 81; Suro Rancho, 2683 m, USNM 199760-66; 62 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY % km W Suro Rancho, 2744 m, USNM 199767- 86, 199872-79. Eleutherodactylus riveti (318 spec. ) ECUADOR: “Equateur’ MHNP 1902-357. Azuay: Cerro Negro (E of Sevilla de Oro), 2927 m, USNM JAP 6545, 6695-98; Cuenca, USNM 146277; 9 km S Cumbe, 3300 m, KU 131077-79; 10 km S Cumbe, 3350 m, KU 131080-145; 21.6 km S Cumbe, 3310 m, KU 141988-91; 28.6 km S Cumbe, 3190 m, KU 141986-87; 2.1 km S Cutchil, 2720 m, 141438-42; 3.1 km S Cutchil, 2730 m, 141433-34; 3.5 km S Cutchil, 2785 m, KU 141424-29: 9.6 km S Cutchil, 2935 m, 141430-32; 11.5 km S Cutchil, 2820 m, KU 141435-37; Laguna de Zurucuchu, 3200 m, KU 119812-56, 119857 (cleared and stained skeleton); 3 km E Sevilla de Oro, 2713 m, USNM JAP 6445-49; 5 km ESE Sevilla de Oro, 2957 m, USNM JAP 6582-87; 8 km ESE Sevilla de Oro, 3140 m, USNM JAP 6704; crest of Azuay-Morona-Santiago frontier, ca. 8 km SE Sevilla de Oro, 3354 m, USNM JAP 6482, 6484; crest at Azuay-Morona-Santiago frontier, ca. 8 km SE Sevilla de Oro, 3384 m, USNM JAP 6566-67. Canar: % km N Biblian, 2620 m, KU 141418; 8 km (by road) NW Biblian, 3100 m, KU 141419-23; 8 km NW (airline) Biblian, 3420 m, KU 131146-209; 15 km SSE Cafar, road to Azoques, UMMZ 132917; 20 km NE Gun, 2927 m, USNM JAP 6395, 6686, 6407, 6410, 6700. Morona-Santi- ago: Paramo de Raranga, 12 km S (airline) Cutchil, 3400 m, KU 120025-70; 8 km S (air- line) Cutchil, 3040 m, KU 120071-79, 120080- 81 (cleared and stained skeletons), 120303; 3 km W San Vicente, 2988 m, USNM JAP 7732; 6 km W San Vicente, 3110 m, USNM JAP 7878-79, 7881-96. Eleutherodactylus ruidus (16 spec. ) ECUADOR: Azuay: Molleturo, 2317 m, AMNH 17588-6083. Eleutherodactylus spinosus (100 spec.) ECUADOR: Morona-Santiago: between Cerro Negro and Pailas, 2652 m, USNM 199916; between Cerro Negro and _ Pailas, 2439-2561 m, USNM 199917; El Cruzado, 2195 m, USNM 199945-66; Loma de Puerco, 2226 m, USNM 199967-71; Pailas, 2195 m, USNM_ 199918-35, 199973-77; 1 km S Plan de Milagro, 1707 m, USNM_ 199978; Rio Pitntza, Cordillera del Condor, 1830 m, KU 147039; San Juan Bosco, 2195 m, USNM 199936-44, 199979-82; San Vicente, 2835 m, USNM_ 199972, 199988-89; Sapote, 2470 m, USNM_ 199891-915, 199986-87; between Sa- pote and Suro Rancho, 2604-2622 m, USNM 199983-95. Eleutherodactylus versicolor (177 spec.) ECUADOR: Loja: 13.2 km E Loja, 2770 m, KU 141443-45; 8-9 km N San Lucas, 3000- 3100 m, KU 119945. Zamora-Chinchipe: Abra de Zamora, 2800 m, KU 119948 (cleared and stained skeleton), 119872-910, 141446-48, 142011-12, 165593-645; 14 km E Loja, 2775 m, KU 119946-47 (cleared and stained skele- tons), 119858-71, 119911-44, 141449-60, 141465-66, 142004; 15 km E Loja, 2685-2710 m, KU 141461, 142005-09; 15.4 km E Loja, 2645 m, KU 141462-63; 16.6 km E Loja, 2560 m, KU 141464; 18 km E Loja, 2510 m, KU 142010. Eleutherodactylus vidua (18 spec.) ECUADOR: Azuay: 32 km S Cumbe, 3180 m, KU (WED 47603). Loja: 8-9 km N San Lucas, 3000-3100 m, KU 120092. Za- mora-Chinchipe: Abra de Zamora, 2800 m, KU _ 120082-91, 120093 (cleared and stained skeleton), 141994, 165648-51. Herp QL668.E257 L92 Leptodactylid frogs of the genus El Ha iii iaenani 62 3 RECENT MISCELLANEOUS PUBLICATIONS UNIVERSITY OF KANSAS MUSEUM OF NATURAL HISTORY* 52. 53. 50. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. Reproductive cycles in lizards and snakes. By Henry S. Fitch. Pp. 1-247, 16 fig- ures in text. June 19, 1970. 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