HARVARD UNIVERSITY 1©© Ernst Mayr Library of the Museum of Comparative Zoology OCCASIONAL PAPERS MCZ LIBRARY MAY 1 8 1995 of the NATURAL HISTORY MUSEUM^ The University of KansaslRSlTY Lawrence, Kansas number 170, pages 1-42 7 december 1994 The Identities of the Colombian Frogs Confused with Eleutherodactylus latidiscus (Boulenger) (Amphibia: Anura: Leptodactylidae) John D. Lynch, Pedro M. Ruiz-Carranza, and Maria Cristina Ardila-Robayo School of Biological Sciences. The Uiiiversin' of Nebraska, Lincoln. Nebraska. 68588 USA. and Research Associate in Herpetology, Natural History Museum. The University of Kansas (JDL): Museo de Historia Natural. Instituto de Ciencias Naturales, Universidad Nacional de Colombia. Apartado Aereo 7495, Bogota, Colombia (MCA-R: PMR-C) ABSTRACT Eleutherodactylus latidiscus (Boulenger) is a lowland species found in western Ecuador and southwestern Colombia; the species is replaced geographi- cally (to the north) by E. cruentus (W. Peters). Eleutherodactylus latidiscus is assigned to the E. cruentus species group. The records attributed to E. latidiscus in Andean Colombia apply to other species (mostly E. supernatis Lynch and E. permixtus sp. nov.). Eleutherodactylus tamsitti Cochran and Coin is not a geo- graphic race of E. latidiscus and is redescribed. Eleutherodactylus permixtus. E. supernatis, and E. tamsitti are not each other's nearest relatives but are assigned to the E. devillei species group. At moderate and intermediate elevations on the western flanks of the Cordillera Occidental of Colombia and Ecuador, one finds a series of long-legged species (£. crenunguis Lynch. E. ocellatus Lynch and Burrowes, E. labiosus sp. nov., and E. orpacobates sp. nov.) allied to E. rubicundus (Jimenez de la Espada). a species found on the cis-Andean slopes of Ecuador. The Central American representative of this group is E. cerasinus (Cope). This series of six species is termed the E. cerasinus species group. Key words: Andes; Eleutherodactylus cerasinus group; E. cruentus group; E. devillei group; Eleutherodactylus latidiscus; Taxonomy. © Natural Histor>' Museum, The University of Kansas, Lawrence. ISSN:009 1-7958 2 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 RESUMEN Eleullierodactyhis latidiscus (Boulenger), una especie de tierras bajas, se encuentra en Ecuador occidental y Colombia suroccidental. Esta especie es remplazada geograficamente (al norte) por E. cruentits (W. Peters). Eleutherodactylus latidiscus es una especie del groupo criientus. Los registros de E. latidiscus de los Andes de Colombia pertenecen a otras especies (principalmente a E. pennixtus esp. nov. y E. supeniatis Lynch). Eleutherodactylus tamsitti Cochran y Coin no es una raza geogratica de E. latidiscus y aqui se redescribe. Eleutherodactylus permixtus, E. supernatis, y E. tamsitti pertenecen al grupo devillei. En los bosques nublados de los flancos occidentales de la Cordillera Occidental de Colombia y Ecuador, se encuentran una serie de especies con piemas largas {E. crenuuguis Lynch, E. occllatus Lynch y Burrowes, E. lahiosus esp. nov., y E. orpacobates esp. nov.) que tienen parentescos con E. ruhicundus (Jimenez de la Espada), una especie de las faldas amazonicas de los Andes del Ecuador. El representante centroamericano de este grupo es E. cerasinus (Cope). Esta serie de seis especies componen el grupo cerasinus. Palabras claves: Andes; Grupo de Eleutherodactylus cerasinus: Grupo de E. cruentus; Grupo de E. devillei: Eleutherodactylus latidiscus: Taxonomia. Aside from some debate conceiTiing the identification of certain frog species in Costa Rica and Panama (Dunn, 1933; Taylor, 1952), the first application of the name Eleutheroclactylus latidiscus (Boulenger) in any modern sense was by Cochran and Goin ( 1970), who reported specimens from Colombia, Ecuador (type locality only), and Panama (using Dunn's, 1933, early identifications). Within Colombia, Cochran and Goin reported material from the Cordilleras Central and Occidental (primarily high alti- tudes), the iniddle Magdalena valley, and the southern extreme of the Cordillera Oriental. The last record was described by them as a new subspecies, Eleutheroclactylus latidiscus tamsitti. Very large digital disks were noted in Eleutheroclactylus cruentus and E. latidiscus by Boulenger (1898), Dunn (1931; 1933), and Taylor (1952). Other large-disked forms from Guatemala and Mexico had been treated as the Eleutherodactylus alfredi Group earlier (Smith, 1939; Smith and Tay- lor, 1948). This condition represents one extreme in the near continuum of disk sizes found within the genus Eleutherodactylus (Lynch, 1976a; 1978). While most Eleutherodactylus species have relatively narrower digital disks (disk width about twice that of digit below disk), the large-disked species have disks approximately 3.0-3.5 times the width of the digit below the disk (Fig. 1 ). Species with relatively narrower digital disks have narrow disks or lack fleshy lateral fringes on the digits. Eleutherodactylus latidiscus was named by Boulenger (1898) on the basis of two large (49.7-53.4 mm snout-vent length [SVL]) adult females from the coastal lowlands of extreme northwestern Ecuador. Dunn (1933) used the name for E. cruentus in Panama. (Only 2 yr earlier, Dunn [1931] applied the name E. ventrimarmoratus to E. cruentus.) However, the MCZ LIBRARY COLOMBIAN ELEUTHERODACTYLUS '• ', •'. :JAr 1 8 1995 HARVARD NIVERSITY Fig. 1. Hands of Eleuthewdactyliis. A. E. latidiscus, ICNMHN 13340. B. E. tamsitti, ICNMHN 22949. C. E. pennixtiis, ICNMHN 22629. broadest misapplication of the name was made by Cochran and Coin ( 1970), who used the name for several species in Colombia (Lynch, 1980). Although not every specimen they reported has been located, they appar- ently examined only a syntype of E. latidiscus. Lynch (1976a) elevated their new subspecies, E. latidiscus tamsitti, to species status without pro- viding evidence in support of the taxonomic change. He wanted to empha- size that there was no evidence that E. tamsitti was the same species that had been named by Boulenger from the lowland Pacific forests. As mentioned by Lynch (1980), there are serious problems with Cochran and Coin's accounts for certain species. They reported the same individual (AMNH 38831) three times as three different species {E. cruentus, E. latidiscus. and E. vertehralis). Seventeen other specimens were reported as both E. latidiscus and E. vertebralis. Most of their material belongs to one of the subparamo species they confused with E. vertebralis; in turn, that species was confused with E. supernatis by Lynch (1980) and is described below. Beginning about 1979, Lynch consistently applied the name E. latidiscus to still a very different frog. As a result, the name E. latidiscus was used by Lynch and his coworkers (Lynch, 1979; Lynch and Duellman; 1980; Lynch and Miyata, 1980) for the species described below as E. lahiosus. Because Lynch associated Boulenger 's latidiscus with that undescribed species, he applied the name E. cruentus (W. Peters) to what we here call E. latidiscus. Savage ( 198 1 ) disputed some of Lynch's distri- butional claims for E. cruentus, but that species does range into northwest- ern Colombia (but not as far south as Ecuador). In describing E. loustes. Lynch (1979) suggested that it was allied to E. latidiscus as well. That suggestion derived from undue attention to the shape of the head as seen in large females. UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 Fig. 2. Dorsal views of heads of Eleutherodactylus laddiscits. A. ICNMHN 13336, male, 24.7 mm SVL. B. ICNMHN 13335, juvenile female, 26.9 mm SVL. C. ICNMHN 13340, adult female, 47.9 mm SVL. When one of us (JDL) first began to study the Ecuadorian frogs of the genus Eleutherodactylus, he was struck by the peculiar head shape of Boulenger's (1898) illustration of Hylocles laticliscus. Cochran and Coin (1970) emphasized the large head with flared lips in their touched-up illustration of one of the syntypes. Following study of the syntypes in 1972, Lynch searched for more recently collected specimens having the same peculiar morphology without realizing that the peculiar head shape was a sexually dimorphic trait (Fig. 2). The confusion persisted in part because he found relatively few sexually mature females and they were scattered through collections. It was not until 1984 that sufficient material was found to re-evaluate and reject the notions that Lynch had harbored for 20 yr. Among those now-rejected notions is the belated discovery that the frog he called E. cnientus from western Ecuador was E. latidiscus (Boulenger). Although several frog species have been associated with the name Eleutherodactylus latidiscus, apparently the only references that actually apply to the species are those dealing explicitly with the syntypes or the use of the name in faunal lists. (They were listed because the name was given for a frog in the Pacific lowlands; the reference implied is to Boulenger's original description and the type-specimens). The frogs that Lynch and Miyata (1980) called E. latidiscus from western Ecuador are not of that taxon. They emphasized two features evident in the syntypes (the fine granulation [shagreen] of the skin of the dorsum and the peculiar head shape [flared lips on proportionately large heads]). This combination of characteristics is most evident in females (whose lips are more flared than those of the males) and occurs in several species that resemble one another in being large, relatively slender frogs with relatively large digital pads (Fig. 1). COLOMBIAN ELEUTHERODACTYLUS MATERIALS AND METHODS Measurements follow Lynch and Duellman ( 1980). The following mea- surements are identified by abbreviations: SVL (snout-vent length), HW (head width), HL (head length, measured so as to estimate snout-occiput distance), lOD (interorbital distance), E-N (eye to nostril distance). Speci- mens were measured to the nearest 0.1 mm with dial calipers. Means are reported ±1 standard error of the mean. Crania were prepared either as double-stained preparations (following Dingerkus and Uhler, 1977) or by picking away musculature and fascia, followed by washing 0.5-3 min in warm dilute (25%) Clorox solution, and washing in cold water for 12 h. In the following accounts, the term tympanum refers to the structure evident on the side of the head of the frog. When the skin covering the tympanic annulus is not adpressed against the annulus. no tympanum is apparent externally (= tympanum concealed). In several Eleiitherodactylus, the annulus is only partially visible (Lynch and Duellman, 1980). In the following accounts, the terni odontophore is used to identify the projection bearing teeth that lies near (or at) the posterior extent of the dentigerous process of the vomer. In each case, these usages are consistent with the uses of such terms for more than a century in descriptions of frogs and to change terminology would lead to ambiguity. Specimens are identified by catalogue number and an abbreviation for the collection (following Leviton et al., 1985). Place names are arranged by country, department (or province), and municipio; in some cases, veredas (subdivisions of a municipio) are given. The specimens examined are listed in the appendix. SYSTEMATIC ACCOUNTS Between the confusion generated by Cochran and Coin (1970) and by Lynch and his coworkers (Lynch, 1979; Lynch and Duellman, 1980; Lynch and Miyata, 1980) over the name E. latidiscus (Boulenger), a variety of frog species has been associated in the literature. The sequence of system- atic accounts presented below is intended to order these taxa in such a way that some of the historical conflation can be sorted out. Three units are buried in this confusion and those units have geographic and phylogenetic components: (1) taxa from the Pacific lowlands, (2) Andean taxa (subparamo and high cloud forests), and (3) taxa from intermediate eleva- tions of the western slopes of the Andes of Ecuador and the Cordillera Occidental of Colombia. 6 univ. kansas nat. hist. mus. occ. pap. no. 170 The Identity of Eleutherodactylus latidiscus Boulenger (1898) named Eleutherodactylus latidiscus on the basis of two adult females from Cachabe, Provincia Esmeraldas, Ecuador. Dunn (1933) applied the name to what is now called E. cruentus (Savage, 1981) and Breder's (1946) use of the name is probably at Dunn's suggestion. Taylor (1952), reacting to Dunn's (1931; 1933) suggestions, pointed out that E. latidiscus and E. ventrimarmoratus were distinct from the Central American species that have large digital pads, but erred in reporting that these species (£. latidiscus and E. ventrimarmoratus) have smooth skin on the venter. Cochran and Coin's (1970) account off. latidiscus [latidiscus] is useful in that they provided a detailed description and illustration [albeit improved with pencil tracings] of one of the syntypes. However, because the species exhibits considerable sexual dimorphism (as does the closely allied E. cruentus), a redescription of the species is desirable. Eleutherodactylus latidiscus (Boulenger) Figure 3 Hylodes latidiscus Boulenger, 1898:121. Both syntypes of Hylodes latidiscus (BMNH 98.4.28.108-09. re-regis- tered as 1947.2.15.66-67) are adult females from "Cachabe ... Prov. Esmeraldas" (= Cachabi, 16 km SE Concepcion, 200 m; Peters, 1955:339). Cochran and Coin (1970:416-417, PI. 55) described BMNH 1947.2.15.67, whereas Boulenger (1898) used the other syntype as the basis for his description and illustration. Diagnosis. — ( 1 ) Skin of dorsum tuberculate in males, shagreen with scattered tubercles in females, that of venter areolate; dorsolateral folds absent; (2) tympanum distinct, round, its length 22-43% eye length; (3) snout subacuminate to rounded in dorsal view, angularly rounded in lateral view; lips flared in larger females; canthus rostralis concave in males, less distinct in females; (4) interorbital space narrower than upper eyelid; cranial crests absent; subconical tubercle on upper eyelid; (5) vomerine odontophores elevated, triangular in outline; (6) males lacking vocal slits; males with glandular nuptial pad on thumb; (7) first linger slightly shorter than second; Digits II-IV bearing broad disks; (8) fingers bearing lateral fringes; (9) ulnar tubercles absent; (10) small tubercle on heel, thick tuberclelike fold on distal tarsus; (11) two metatarsal tubercles; inner oval, four times size of rounded outer; supernumerary plantar tubercles indis- tinct; (12) toes bearing lateral fringes, no webbing; toe disks broad, slightly smaller than those of fingers; ( 1 3) dorsal pattern polymorphic, usually with pale lines along outer edge of W-shaped occipital mark; venter white with some brown stippling; groin bearing pale area (no spots); posterior surfaces of thighs brown with small cream spots; (14) adults moderate sized, males COLOMBIAN ELEUTHERODACTYLUS Fig. 3. A. Eleutherodactylus hybotragus, ICNMHN 13337, female. 36.5 mm SVL. B. E. latidiscus. ICNMHN 13340, female, 49.0 mm SVL. C. E. tamsitti, ICNMHN (JDL 17736). female, 50.5 mm SVL. D. E. supeniatis, ICNMHN 8069, female, 42.6 mm SVL. 21.9-25.9 (x = 23.8 ± 0.3. // = 16) mm SVL, females 35.2-53.4 (x = 42.3 ±1.3.//= 14) mm SVL. Eleutherodactylus latidiscus is most like E. cruenfus, but the two differ in that males of E. cruentus have pale spots edged with dark pigment in the groin and have concealed tympana. The tympanum is less prominent in female E. cruentus and is smaller than in E. latidiscus. The heel tubercle, tubercles along the outer edge of the tarsus, and ulnar tubercles are more prominent and larger in E. cruentus than in E. latidiscus. The dark pigment on the posterior surfaces of the thigh is darker with better definition of pale spots (if present) in E. cruentus. Description. — (See Table 1 for proportions.) Head wider than body (except in gravid females), longer than wide; snout shape variable sexually and by size (Fig. 2) — Males: snout subacuminate in lateral view, angularly rounded (= nearly truncate) in lateral view; nostrils protuberant, directed dorsolaterally; canthus rostralis prominent, concave; loreal region con- cave, sloping abruptly to lips; lips not flared. Females, snout subacuminate to rounded in dorsal view, angularly rounded in lateral view; nostrils weakly protuberant, directed dorsolaterally; canthus rostralis less distinct, straight to sinuous; loreal region concave, sloping gradually to flared lips. UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 00 +1 c a (U 1) o r- ^ ^ ri +1 ri 00 +1 00 f^ o ON '*_ >vd — ■ ^ +1 m 00 _• ;:r ^ — rj OS g, ^ t^ 00 00 00 "^ ri ON ri " +1 IJ 00 O O o r-i 00 00 ri +1 ON r- o oo ^ 00 'O „■ iri -^ O CI 11 -T +1 V On ri ON 00 o o ON — 1/3 > (U c T3 C o o [/5 > (U S Q O On -* r<^ 00 ITj ITi ri ON ri — ^ 00 d >n d VD d ^ ■ +1 1 +1 ri +1 1 +1 q — < r- — ; -^ -^ q o rj in vd r*-) 00 d r~^ d (N m r^ m r^i r-i 00 -* On +1 00 rj ri +1 ri 2; 00 ri +1 in r4 00 +1 3 in 7 o o^ On ON o 00 <^i T rj q ON ^^ 00 - — 1 00 1 — 1 00 d +1 in in rr] ri m T o +1 o in I r '5i) u n > CO r4 rr, r<", ro ON ri in ri d d n-] d — ' d ^ d f +1 7 +1 T +' 1" +1 ^_ NO ON ro — ; r-- — ; 00 inr-^NOdodoNdd o ri — ; d "^ "^ r-^ 00 m CO O ri — ; d 00 in r-^ ON r<-i r<-; OO rn r i d in 00 r^ ON oo ra NO rn d d -h' d "f +1 "f +1 00 ^ -^ ro t^ ON NO ON m r<~t m en Co > 1^ NO 00 r- ^ NO in NO 1 o +1 in o +1 ON 1 o +1 00 in 1 o +1 00 mj r~- ri ON m. ON (-<-l NO ri m, o ITj in in m, mi in NO r<-, ri in r^ m o m. ON NO in 1 O +1 ft in 1 o +1 mi 1 o +1 o m, O +1 ■xt- mj 1 o +1 o m On ■xt- m NO -* ^ NO ON o m, ^ ■* in o in 1^ rl -* ^ UJ NO r^ ri 00 O O Ol m^ rj o C/3 (U X oo B o o § o o B o o o o g o o c u o ;-i (L) CI, «1 C3 C o o O X) t2 C/5 c a UJ o H T3 .2 va "3 cS c3 t/3 a" 2 E OS o ■o 13 +-1 U W3 o u < o o o o -4—* C c n c (U ^ >- Z >H 3 r<-, IT) C I I a ^' ^' z z ^• GO W3 t« 1/3 t/3 CJ -J c oh I I rt ^ 00 J= (^ ^ CO - H Q C« C« Q Z^^Z o z ^;^ ^ OJ CJ ^ ^ I rt in ON j:: r<-i in 00 c I I -- in (U o T3 ^H CJ x> 3 CJ X3 iA> C3 3 X CJ N N 3 "flj QJ X H. 3 z V5 CJ c 4J C C/0 '55 _CJ _aj f— C CJ 28 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 Fig. mm. 11. Cranium of Eleiitherodactxhis ruhicundus, KU 177452. Scales = 3 Fig. 12. Elcuthciodactyhts lahiosiis (A) KU 131612, holotype and (B) ICNMHN 13243, male, 36.0 mm SVL. Eleutherodactylus orpacohates (C) ICNMHN 20249, female, 43.9 mm SVL and (D) amplectant pair, ICNMHN (JDL 18805-06), female 47.2 mm SVL. COLOMBIAN ELEUTHERODACTYLUS 29 Ecuador (redescribed by Lynch and Duellman, 1980). Eleuthewdactylus cerasinus, E. crenunguis, E. ocellatus, E. rubicundus, the undescribed frog Lynch confused with E. latidiscus. and another spe- cies from western Colombia form a small phenetic group — a group that we consider to include the nearest relatives of all the taxa. These animals superficially resemble the large-disked E. cruentus and E. latidiscus, but have longer legs and lack lateral fringes on the digits. Aside from some slight differences in color, these animals differ from one another in terms of expression of eyelid and heel tubercles, shagreen versus tuberculate skin of the dorsum, presence of nuptial pads and vocal slits in males, and the lengths of the inner digits of the hand (Table 3). Osteologically, the species (aside from E. ocellatus which has not been studied) are very similar as well. The shapes and sizes of skull bones agree among the five species for which skeletal material has been prepared and are distinguishable (as a group) from the species once confused with E. latidiscus (E. cruentus. E. permixtus. E. supernatis, E. tamsitti). Cranium. — (Based on Eleutherodactylus rubicundus) The nasals are long (Fig. 11) in comparison to those of E. cruentus. E. permixtus. E. supernatis, and E. tamsitti, and are closely juxtaposed along the anterior half to two thirds of their length. Posteriorly, the nasals are widely sepa- rated. The sphenethmoid is well ossified and extends well anterior beneath the nasals, reflecting the elongate snouts of species of this group. The frontoparietals are of equal width through the length of the orbit and nearly cover the frontoparietal fontanelle. No cranial crests are evident in large females. The crista paroticae are more slender than those seen in E. permixtus or E. supernatis (Fig. 10). The alary processes of the premaxillae are directed dorsally (with a slight posterior vector) and the palatal shelf is dissected moderately. The vomers are large and narrowly separated in their posterior half. The odontophores are well anterior of the neopalatines. The neopalatines are somewhat spatulate medially. The cultriform process is long, acuminate, and extends between the neopalatines. The alary pro- cesses are peipendicular to the cultriform process and are narrowly over- lapped by the median rami of the pterygoids. The pars facialis of the maxilla is only moderately high, but the posterior end of the maxilla (and quadratojugal) are low. The zygomatic ramus of the squamosal is nearly as long as the otic ramus and relatively high. The otic plate is narrow. Eleutherodactylus labiosus sp. nov. Figure 12 Eleutherodactylus latidiscus — Lynch, 1979:502-03; Lynch and Duellman, 1980:23-24, 64: Lynch and Miyata, 1980:6, 11. Holotype. — KU 131612, a juvenile (but see Remarks) female from La 30 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 Palma (junction of Highways 28 and 30), Provincia Pichincha, Ecuador, 920 m, obtained 8 August 1970 by John D. Lynch. Paratypes. — ICNMHN 13242-43, Colombia, Departamento de Valle del Cauca, Municipio Restrepo, Vereda Alegre, Campo Las Vegas, 200 m, ICNMHN 13241, Vereda Alegre, Rfo Calima; AMNH 102759, MCZ 90049-50, Hotel Tinalandia, 15 km SE Santo Domingo de los Colorados, Provincia Pichincha, Ecuador, 800 m. Diagnosis. — ( 1 ) Skin of dorsum shagreen, bearing low occipital folds, that of venter areolate; dorsolateral folds absent; (2) tympanum distinct, higher than long, its length 24-39% eye length; (3) snout subovoid in dorsal view, rounded in lateral profile; lips flared in adult females; canthus rostralis well defined, straight to weakly concave; (4) lOD narrower than upper eyelid; low cranial crests in adult females; conical tubercle on upper eyelid; (5) vomerine odontophores prominent, triangular in outline; (6) males with vocal slits, nuptial pads absent; (7) first finger slightly shorter than second; Fingers II-IV bearing broad discs; (8) fingers lacking lateral fringes; (9) ulnar tubercles absent; (10) small conical tubercle on heel, row of tubercles along outer edge of tarsus; (11) two metatarsal tubercles, inner oval, about four times size of rounded outer tubercle; supernumerary tubercles only at bases of toes; (12) lateral fringes and toe webbing absent; toes with broad discs, smaller than those of fingers; (13) brown above with darker markings; venter cream with brown reticulation; posterior surfaces of thighs dark brown with cream flecks; (14) adults moderate sized, males 35.4-50.8 (.f = 44.3 ± 1 .6, /; = 12) mm, females 48.5-52.3 {x = 50.4, // = 2) mm SVL. Eleutherodactylus labiosus is most similar to E. crenunguis, but differs in having prominent tubercles on the upper eyelid and a shorter thumb. Description. — Head as broad as body, slightly wider than long; HW 37.7^0.2% (J = 39.1 ± 0.2, // = 12) SVL in males, 36.7^1.4% (.r = 39.3 ± 0.4, // = 10) in females; snout suboval in dorsal view, rounded in lateral profile, long; nostrils weakly protuberant, directed dorsolaterally; canthus rostralis well defined, edge rounded, straight to weakly concave; loreal region concave, sloping to flared lips; E-N 85.4-1 16.4% (.v = 100.3 ± 2.4, n = 12) eye length in males, 87.9-110.2% {x = 98.1 ± 2.4, // = 10) in females; large, sometimes conical tubercle on posterolateral part of upper eyelid (anterior end of occipital W); edges of frontoparietals upturned, producing shallow furrow; interorbital space broad; upper eyelid 91.1- 137.9% {x = 1 19.7 ± 13.6, // = 12) lOD in males, 93.8-143.2% {x = 123.6 ± 4.9, n = 10) in females; supratympanic fold low; tympanum distinct, superficial, higher than long, separated from eye by 1.5 times its length; tympanum length 23.6-36.7% (x = 30.6 ± 1.3,//= 12) eye length in males, 25.0-38.8% (J = 31.1 ± 1.2, // = 10) in females; postrictal tubercles present, subconical, not prominent; choanae relatively small, longer than wide, not COLOMBIAN ELEUTHERODACTYLUS 31 concealed by palatal shelf of maxillary arch; vomerine odontophores me- dian and posterior to choanae, triangular in outline, each approximately twice size of a choana, separated medially by distance about one-sixth odontophore width, bearing a transverse row of 9-1 1 teeth; tongue longer than wide, its posterior border not notched, posterior one third to two fifths not adherent to floor of mouth; males with long vocal slits posterolateral to tongue. Skin of dorsum shagreen (closely packed flattened granules), least evi- dent on top of head and upper surfaces of limbs; low folds forming occipital W; apices of W extend anteriorly in form of X; dorsolateral folds absent, but low, indistinct folds beginning at posterior edge of head extend- ing short distances down flanks; flanks shagreen above, becoming areolate near venter; discoidal folds well anteriad to groin; venter finely areolate, throat more obviously areolate; skin below vent coarsely areolate; skin on undersides of limbs smooth; anal sheath absent; ulnar tubercles absent; palmar tubercle bifid, much larger than oval thenar tubercle; numerous low supernumerary palmar tubercles; subarticular tubercles elevated, round; no lateral keels on fingers; fingers long and slender bearing large expanded disks (disk of III-IV about 3 times widths of digits below disks, of l-II about twice); disks truncate with slight emargination; disk of third finger 1.75-2 times length of inner metatarsal tubercle; all disks bearing broad ventral pads; if Fingers I and II are adpressed equally, tip of I reaches two- thirds distance up the disk of II (inner two fingers approximately equal in length); nuptial pad absent on thumb of male. Small conical tubercle on heel; row of up to five smaller nonconical tubercles along outer edge of tarsus; folds and tubercles absent on inner edge of tarsus; inner metatarsal tubercle slightly more than twice as long as wide, outer metatarsal tubercle round, not elevated, about 25% size of inner; supernumerary plantar tubercles at bases of Toes II-IV; subarticular tubercles round, elevated, nonconical; toes with slight lateral keels, no webbing; disks and pads broad, smaller than those of fingers; fifth toe slightly longer than third toe when each is adpressed against fourth; tip of fifth toe reaches about halfway between distal and penultimate subarticular tubercles of Toe IV, whereas tip of third toe reaches to distal border of penultimate subarticular tubercle of Toe IV; heels overlapping when flexed hind limbs held at right angles to sagittal plane; shank 52.7-64.2% (r = 58.6 ± 1.0, /; = 12) SVL in males, 56.2-64.0% (j = 60.0 ± 0.9. /? = 10) in females. Color in preservative: Dorsum brown with pale occipital W, edged with dark brown, brown sacral chevron, interorbital bar; limb bars vague on thighs, bolder on other parts of limbs; bars narrower than interspaces, weakly oblique on shanks; posterior surfaces of thighs rich brown with few cream flecks to several small spots (KU 145000); faint canthal- 32 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 80 Fig. 1 3. Map of Colombian locality records of Eleuthewdactylus lahiosus (■) and E. orpacohates (•). supratympanic stripe; cream line along upper lip with dark brown patches above it; dorsal pigmentation set off from venter by heavy brown reticula- tion on nearly white background of flank; some individuals having brown flanks enclosing large white spots; this brown reticulum extending along anterior surfaces of thighs as an irregular line and along postaxial edge of underside of shank; ventral surfaces cream with few brown vermiculations on anterior belly, breast, throat. Color in life: Dorsal surfaces brown to dark brown with rust to orange highlights on warts and ridges; venter off-white to dirty cream with brown flecking to black marbling; throat usually darker than venter; labial bar cream to dull yellow; posterior surfaces of thighs fleshy-brown; iris pale green above, gray below, with a brown horizontal streak and black reticu- lation. (JDL field notes, May-June 1983). The iris of the holotype was COLOMBIAN ELEUTHERODACTYLUS 33 described as "bright reddish rust in upper quarter, gray over remainder, all reticulated with black" (JDL field notes, 8 August 1970). Etymology. — Latin, meaning large-lipped, in reference to the flaring of the lips in adult females. Distribution. — Found at low and moderate elevations from the Rio San Juan, Colombia, south to west-central Ecuador (Fig. 13). In general, Eleuthemdactylus lahiosiis and E. creminguis replace one another geo- graphically. The fonner occurs primarily at elevations below 1000 m, whereas the latter occurs at elevations of 800-1600 m. The two species are found sympatrically at Tinalandia and La Florida in Provincia Pichincha, Ecuador, but the lower cloud forests have been poorly collected in Ecuador and scarcely at all in southwestern Colombia. Remarks. — Only two adult females have been examined (AMNH 110871, ICNMHN 13241). The other females are anatomical juveniles (straight oviducts with only small ovarian eggs) despite the fact that some are larger than the "adults" (e.g., KU 145000 is 55.6 mm SVL). The holotype is a juvenile female 45.4 mm SVL and KU 145001 is approxi- mately the same size (45.5 mm SVL). The absence of females having moderate convolutions of the oviducts (what JDL terms "young females") suggests that this species may not exhibit a normal eleutherodactyline pattern of oviductal moiphology (thin, straight oviducts in juveniles, some bending of the oviducts in larger females, and extensive convolution and expansion in adult females, gravid or not). The absence of such a pattern is found in some other frog species (e.g., Rana blairi and Rana pipiens) in which convolutions occur during the reproductive season, but then regress. Eleuthewdactyhis orpacohates sp. nov. Figure 12 Holotype. — ICNMHN 20249, adult female from Quebrada Agudelo, Parque Nacional Natural Las Orquideas, Vereda Calles, Municipio Urrao, Departamento Antioquia, Colombia, 1410-1430 m, one of a series ob- tained 24 May 1988 by M. C. Ardila, J. D. Lynch, R M. Ruiz, and R. Sanchez. Paratypes.— ICNMHN 20250-63, collected with holotype: ICNMHN 20264-74, Quebrada de las Canoas, Vereda Calles, Municipio Urrao, 1770- 1870 m; ICNMHN 20244^8, Quebrada El Silencio, Vereda Calles, Municipio Urrao, 1480-1540 m; ICNMHN 20278-82, Quebrada Vironda, Parque Nacional Natural Las Orquideas, Vereda Calles, Municipio Urrao, 1400-1450 m; ICNMHN 20275-77, vicinity INDERENA cabaiia "Rio Calles," Parque Nacional Natural Las Orquideas, 1410-1525 m; CSJMHN 1922, 1925, 1929, 1933-34, 1940, Quebrada Sabaletas,Antad6, Municipio Ituango, Antioquia, Colombia, 1420 m. 34 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 Diagnosis. — (1) Skin of dorsum granular, with scattered larger tu- bercles, that of venter areolate; dorsolateral folds absent; (2) tympanum distinct, round, 22-37% eye length; (3) snout subacuminate in dorsal view, rounded in lateral profile; lips slightly flared in adult females; canthus rostralis straight or weakly concave; (4) interorbital space narrower than upper eyelid; low cranial crests in females; one to three subconical tu- bercles on upper eyelid; (5) vomerine odontophores elevated, triangular in outline; (6) males with white nuptial pad on thumb, lacking vocal slits; (7) first finger shorter than second; Fingers Il-IV bearing large discs; (8) fingers without lateral fringes; (9) ulnar tubercles small, nonconical; (10) nonconical tubercle on heel, row of tubercles along outer edge of tarsus; (11) two metatarsal tubercles, inner oval, three to four times size of outer; supernumerary tubercles only at bases of toes; (12) toes without lateral fringes or webbing, discs broad, smaller than those of fingers; (13) brown above with darker markings; venter off-white with gray spots or reticulum; posterior surfaces of thighs dark brown with minute white spots; (14) size geographically variable, in most populations, males 24.3-35.6 {x = 30. 1 ± 0.7. // = 18), females 43.6^8.4 (.v = 45.9 ± 0.4, n = 16) mm SVL; in southern Departamento Valle del Cauca, Colombia, males 25.7-29.7 (v = 27.7 ± 0.2, // = 21) mm, females 35.8-41.5 ( x = 39.6, n = 6) mm SVL. Eleutherodactylus orpacohates most closely related to E. cerasimts, E. crenunguis, E. labiosiis, E. ocellatus, and E. riihiciindus, but differs from E. cremmgids and E. lahiosiis in having nuptial pads in the male (vs. absent) and in lacking vocal slits (present in those species). Those species have shagreen skin of the dorsum, rather than the more granular skin with scattered tubercles of E. orpacobates. Eleutherodactylus orpacohates has conical eyelid tubercles as does E. labiosus (absent in E. crenunguis, E. ocellatus, and E. rubicundus), but is a smaller frog (comparable in size to E. ocellatus and E. rubicundus). Eleutherodactylus cerasinus is markedly smaller (Savage, 1981) and has obvious inner tarsal tubercles. Description. — (For proportions, /? = 1 8 for males, 16 for females.) Head as broad as body, longer than wide; HW 38.1-41.2% {x = 39.7 ± 0.2) SVL in males, 38.7^1.7% (.r = 40.1 ± 0.2) in females; snout subacuminate in dorsal view, rounded in lateral profile; E-N 77.8-100.0% {x = 90.8 ±1.5) eye length in males, 92.1-113.7% {x = 105.8 ± 1.6) in females; nostrils weakly protuberant, directed dorsolaterally; canthus rostralis straight, evi- dent but edge rounded; loreal region concave, sloping relatively gradually to lip; lips slightly flared in larger females; one to three conical tubercles on tuberculate upper eyelid; interorbital space broad, low cranial crests (up- turned edges of frontoparietals) in females; upper eyelid width 100.0- 139.4% (j= 118.3 + 2.4) lOD in males, 100.0-139.1% (z= 1 1 1.2 ± 2.5) in females; heavy supratympanic fold obscuring upper edge of tympanum; tympanum superficial, not prominent, separated from eye by distance equal COLOMBIAN ELEUTHERODACTYLUS 35 1.5-2 times tympanum length; tympanum length 22.4-31.1% {x - 27.6 ± 0.5) eye length in males, 24.5-37.2% (x = 31.3 ± 0.8) in females; one or two conical postrictal tubercles; choanae not concealed by palatal shelf of maxillary arch, moderately large; vomerine odontophores median and pos- terior to choanae, about size of a choana, separated on midline by distance nearly equal width of an odontophore; odontophores elevated, bearing transverse row of seven to nine teeth; tongue longer than wide, posterior third not adherent to floor of mouth, posterior edge notched; vocal slits absent in males. Skin of dorsum granular but beset with more pungent tubercles on lower back and flanks; dorsolateral folds absent; skin of venter areolate; discoidal folds well anteriad to groin; anal sheath absent; three to five nonconical ulnar tubercles; palmar tubercle bifid, much larger than oval thenar tu- bercle; supernumerary palmar tubercles numerous, low. diffuse; subarticular tubercles distinct, round, nonconical; indistinct lateral keels on III-IV; fingers bearing expanded disks, largest on III-IV, round apically; ventral pads broader than long, surrounded by grooves; first finger shorter than second; males bearing large, compact, white nuptial pad atop each thumb. Nonconical tubercle on heel; row of four or five outer tarsal tubercles; indefinite tubercle on distal portion of inner tarsal surface (otherwise, no inner tarsal folds or tubercles); inner metatarsal tubercle twice as long as wide, three to four times size of low outer metatarsal tubercle (longer than wide); low, indistinct supernumerary plantar tubercles at bases of Toes II- IV; subarticular tubercles round, nonconical; no lateral keels on digits; digits long, slender, bearing expanded disks; disks round apically, smaller than those of fingers; fifth toe longer than third when each is adpressed equally against fourth; tip of fifth toe not reaching base of distal subarticular tubercle of Toe IV, whereas tip of third toe reaches distal border of penultimate subarticular tubercle of Toe IV; heels overlapping when flexed hind legs held perpendicular to sagittal plane; shank 53.6-63.1% (x = 57.5 ± 0.6) SVL in males, 51.9-58.7% {x = 55.4 ± 0.6) in females. Color ill preservative: Dorsum brown with darker brown head and shoulders, some dark brown chevrons on lower back, continuing across thighs onto shanks; limb bars oblique on shanks, narrower than interspaces; indistinct pale postocular ridges; white flecks in oblique rows across upper flanks onto lower flanks; white flecks on upper surfaces of limbs; belly off- white with spots and short dashes of dark gray, forming a loose reticulum on anterior surfaces of thighs, lowest flanks; throat more heavily stippled with brown; dense reticulum (of dark gray or brown) on undersides of shank, anterior surfaces of thighs, flanks; posterior surfaces of thighs dark brown or black with small (much smaller than pad of thumb) white spots. Color in life: Dorsum pale to dark brown with brown markings and 36 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 17U cream (or yellow) flecks; some ridges and warts with rust highlights; throat washed with brown; crural stripe (when present) salmon to dull pale orange; venter dirty cream to off-white with gray to brown flecks, spots, or reticulum; posterior surfaces of thighs brown to black with cream to white flecks; underside of shank banded black and white; iris bright copper to rich brown with black flecks; concealed tunic of eye blue. Eleutherodactylus orpacobates exhibits considerable metachrosis. The color description given above is for animals observed during the day. At night, these animals are quite pale with the predominant ground color being cream (markings are brown). Measurements of holotype in mm. — SVL44.1; tibia 24.9; HW 17.7; HL 16.9; chord of HL 18.7; upper eyelid width 5.0; lOD 4.4; tympanum length 1.5; eye length 5.5; E-N 5.8. Etymology. — Greek (orpacos + hates), meaning one who frequents twigs. Most specimens have been found perched on twigs in cloud forests where they assume a Gastrotheca-like stance. Natural history. — Amplectant pairs are relatively common in this spe- cies. Each pair has been found perched on a liana or twig. No vocalization or other reproductive activity has been seen. Adults and juveniles are found at night perched on twigs (very occasionally on leaves) in wet cloud forest. Distribution. — Known from moderate elevations ( 1 140-2000 m) along the western flank of the Cordillera Occidental of Colombia from Paramillo (Antioquia) to northern Valle del Cauca (Fig. 13). Remarks. — Specimens from the most southern populations (Lago Calima and Yotoco, Valle del Cauca) are smaller than those over the remainder of the distribution. Even specimens from northern Valle del Cauca (Charco Azul, Mcpio. El Cairo) are as large as those from Antioquia. The small frogs from southern Valle del Cauca are larger than E. cerasinus, the smallest member of the group, and lack the inner tarsal tubercles of that species. Eleutherodactylus orpacohates has three pattern polymorphisms. The normal morph is described above (and see Fig. 12C, as well as male in Fig. 12D). One variant (female. Fig. 12D) has a crural stripe (the outer edge of the shank bears an orange stripe that obliterates bars on the shank). This variant is seen in 15 individuals (9 from the Lago Calima site, 1 from Charco Azul, 1 from the Orquideas samples, and 4 from Murri). The third variant is the middorsal stripe (a thin cream line from the tip of the snout to the vent) and is seen in 12 individuals (5 from Lago Calima, 2 from Pueblorico, 2 from Orquideas, 1 from Murri, and 2 from Ituango). One individual (KU 168087) has both the crural stripe and middorsal stripe. The Lago Calima site (1230 m) is near the Yotoco site (1590 m) and each sample consists of 47 specimens. Only the common variant is seen in the Yotoco sample. The difference in polymorph frequency between these COLOMBIAN ELEUTHERODACTYLUS 37 two sites suggests the absence of gene flow (an observation in agreement with Lynch's, 1992. conclusion for the better-collected E. erythropleura). Acknowledgments : Specimens were loaned, permission to prepare skel- etons was given, or working space extended by the following curators: L. Coloma. R. Crombie, W. E. Duellman, D. Frost, Hermano Daniel Gonzalez, A. Grandison, W. R. Heyer, R. F. Inger, A. G. Kluge, H. Marx, R. McDiarmid, K. Miyata, C. W. Myers, Hermano Niceforo Maria, R. Nussbaum, J. A. Peters, J. Restrepo, J. Rosado, V. Rueda, Hermano Marco A. Serna, H. Voris, C. Walker, E. Williams, G. Zug, and R. G. Zweifel. Our fieldwork during the past 13 years benefitted from the assistance and companionship of R Bernal, H. Carvajal, O. Pinto, J. Restrepo, J. Renjifo, V. Rueda. R. Sanchez, and M. Serna. Our fieldwork has been supported by the Universidad Nacional de Colombia, University of Nebraska, and a fellowship from the Fulbright Commission. Access to many areas was granted by officials of INDERENA; the help of F. Ramirez was especially important. LITERATURE CITED BouLENGER. G. A. 1898. An account of the reptiles and batrachians collected by Mr. W. F. H. Rosenberg in western Ecuador. Proc. Zool. Soc. London 1898:107- 126. Breder, C. M., Jr. 1946. Amphibians and reptiles of the Ri'o Chucunaque Drainage, Darien, Panama, with notes on their life histories and habits. Bull. Am. Mus. Nat. Hist. 86:375-436. Cochran, D. M., and C. J. Coin. 1970. Frogs of Colombia. Bull. U.S. Natl. Mus. (288): 1-655. DiNGERKUS, G., AND L. D. Uhler. 1977. Enzyme clearing of Alcian Blue stained whole small vertebrates for demonstration of cartilage. Stain Technology 52:229-232. Dunn, E. R. 1931. The amphibians of BaiTO Colorado Island. Occas. Pap. Boston Soc. Nat. Hist. 5:403^21. Dunn, E. R. 1933. Amphibians and reptiles from El Valle de Anton, Panama. Occas. Pap. Boston Soc. Nat. Hist. 8:65-79. Flores, G. 1988. Anew species oi Eleiitherodactylus (Amphibia: Leptodactylidae) from the Pacific slopes of the Ecuadorian Andes, with comments on the devillei assembly. Copeia 1988:110-116. Leviton, a. E., R. H. Gibbs, Jr., E. Heal, and C. E. Dawson. 1985. Standards in herpetology and ichthyology: Part I: Standard symbolic codes for institutional resource collections in herpetology and ichthyology. Copeia 1985:802-832. Lynch, J. D. 1976a. The species groups of the South American frogs of the genus Eleutherodactyhis (Leptodactylidae). Univ. Kansas Mus. Nat. Hist. Occas. Pap. (61): 1-24. Lynch, J. D. 1976b. New species of frogs (Leptodactylidae: Eleiitherodactylus) from the Pacific versant of Ecuador. Univ. Kansas Mus. Nat. Hist. Occas. Pap. (55): 1-33. 38 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 Lynch, J. D. 1978. A new eleutherodactyline frog tYom the Andes of northern Colombia (Leptodactylidae). Copeia 1978:17-21. Lynch, J. D. 1979. A new species of Eleutherodactylus from northern Ecuador (Amphibia: Leptodactylidae). Proc. Biol. Soc. Washington 92:498-504. Lynch, J. D. 1980 ["1979"]. The identity of Elciithemclacryliis vertchvalis (Boulenger) with the description of a new species from Colombia and Ecuador (Amphibia: Leptodactylidae). J. Herpetol. 13:411^18. Lynch, J. D. 1983. A new leptodactylid frog from the Cordillera Oriental of Colombia, pp. 52-57, in Anders, A. G. J, and K. Miyata (eds). Advances in Herpetology and Evolutionary Biology. Mus. Comp. Zool., Harvard Univ. Lynch, J. D. 1984. New frogs (Leptodactylidae: Eleutherodactylus) from cloud forest of the northern Cordillera Oriental, Colombia. Contrib. Biol. Geol. Mil- waukee Publ. Mus. (60): 1-19. Lynch, J. D. 1992. Distribution and variation in a Colombian frog, Eleutherodactylus erythroplewa (Amphibia: Leptodactylidae). Stud. Neotropi- cal Fauna Environ. 27:211-226. Lynch, J. D., and P. A. Burrowes. 1990. The frogs of the genus Eleutherodactylus at the La Planada Reserve in southwestern Colombia with descriptions of eight new species. Univ. Kansas Mus. Nat. Hist. Occas. Pap. (136): 1-31. Lynch, J. D., and W. E. Duellman. 1980. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Univ. Kansas Mus. Nat. Hist. Misc. Publ. (69): 1-86. Lynch, J. D., and K. Miyata. 1980. Two new species of Eleutherodactylus (Am- phibia: Leptodactylidae) from the lowlands and lower cloud forests of western Ecuador. Breviora (457): 1-12. Peters, J. A. 1955. Herpetological type localities in Ecuador. Rev. Ecuatoriana Entomol. Parasitol. 2:335-3^52. RivERO, J. A., and M. a. Serna. 1987. Tres nuevas especies de Eleutherodactylus (Amphibia, Leptodactylidae) de Antioquia, Colombia. Carribean J. Sci. 23:386- 399. Savage, J. M. 1981. The systematic status of Central American frogs confused with Eleutherodactylus cruentus. Proc. Biol. Soc. Washington 94:413—420. Smith, H. M. 1939. Mexican herpetological novelties. Proc. Biol. Soc. Washington 52:187-196. Smith, H. M., and E. H. Taylor. 1948. An annotated checklist and key to the Amphibia of Mexico. Bull. U.S. Natl. Mus. (194): 1-1 18. Taylor, E. H. 1952. A review of the frogs and toads of Costa Rica. Univ. Kansas Sci. Bull. 35:577-942. APPENDIX: SPECIMENS EXAMINED Eleutherodactylus cerasinus (80) COSTA RICA: Prov. Cartago: Turrialba, KU 28341-45. Prov. Heredia: Puerto Viejo, KU 36956. Prov. Limon: Estrella, KU 35373; La Lola, KU 37487; Suretka, KU 36945, 36957-64, 37036-38. PANAMA: Prov. Bocas delToro: 4.8-12.9 km WAlmirante, KU 1 14067-71; Cayo Nancy. 2-40 m, KU 1 14058; Isla de Colon, La Gruta, 20 m, KU 1 14059-61 ; S COLOMBIAN ELEUTHERODACTYLUS 39 end Isla Popa. 10 m. KU 114062; mainland, NW Isla Split Hill, KU 114063; mouth of Rio Cahuita. 1 m. KU 114056-57; Rio Changena, 650-830 m, KU 1 14013-55; Rio Claro at Rio Changena. 910 m. KU 1 1401 2; Peninsula Valiente, 10 m, KU 114064-66. Eleutherodactylus cruentus (36) COLOMBIA: Depto. Antioquia: Mcpio. Dabeiba, Campamento "Pantanos," Rio Amparrado, 805 m. ICNMHN 10565-72, 10580-83, 10612-17, 10619-20, 10634-39, 13869-74. Depto. Choco: Mcpio. Istmina, Quebrada Perado, ICNMHN 3218 (ID questionable, possibly = E. laticlisciis): Mcpio. San Jose del Palmar, Alto de Oso, 10 km W La Italia (Valencia), 1000 m, UVC 8257-58, 8260. Eleutherodactylus lahiosus (85) COLOMBIA: Depto. Cauca: Mcpio. El Tambo, La Costa, 1000 m, KU 145000-02. Depto. Choco: Quebrada Pangala, east bank lower Ri'o San Juan (ca. 17 km airline NE Palestina). AMNH 1 10871. Depto. Valle del Cauca: Mcpio. Dagua, Rio Anchicaya, 300 m, KU 168136; Mcpio. Restrepo, Vereda Alegre, Campo Las Vegas, 200 m, ICNMHN 13242^3, Rio Calima, ICNMHN 13241. ECUADOR: Prov. Carchi: cabeceras del Rio Baboso, near Lita, MECN LAC 138. Prov. Cotopaxi: on frontier of Bolivar province, ca. 7 km airline SSW El Corazon, 800 m, AMNH 104961. Prov. Pichincha : Centinela, 14.1 km SE Patricia Pilar. 550-600 m. MCZ 97575-91; Centro Cientitico Rfo Palenque, 47 km S Santo Domingo de los Colorados (road to Quevedo), KU 165895, MCZ 90053-54, 91954-56, 92113, 94461, 94866; along Highway 28 (old road to Quito), 16 km from junction with Highway 30, 1500 m, MCZ 90006; [Hotel] Tinalandia, 15-16 km SE Santo Domingo de los Colorados, 800 m, AMNH 102759, MCZ 88392-93, 88891-93, 8^9974-92, 89993-90004, 90049-52, 91953; La Florida, near Alluriquin, QCAZ 569-70, MECN LAC 210; Las Palmas, junction Highways 28 & 30, 920 m, KU 131612, 165894, 165896. Eleutherodactylus latidiscus (76) COLOMBIA: Depto. Cauca: Mcpio. El Tambo, Rio Munchique, 800 m, KU 145006, 1000 m. KU 145008; Mcpio. Timbiqui, Quebrada Guangui, AMNH 88947, 88950-53; Depto. Choco: Mcpio. Istmina, Quebrada Docordo, AMNH 87028-33; Mcpio. Tado, 2 km above Playa de Oro, AMNH 87034^2, upper Ri'o San Juan, AMNH 87044-50; Depto. Valle del Cauca: Mcpio. Dagua, Bajo Anchicaya, 300 m, UVC 561 1, 5757-61, 5763-65, 6612, 6827; Mcpio. Darien, 1.5 km W Lago Calima, 1230 m, KU 168010-11; Mcpio. Restrepo, Vereda Alegre, Campo Agua Bonita, 300 m, ICNMHN 13338, Campo Chanco, 460 m, ICNMHN 13335-36, 13343, Campo Las Vegas, 200 m, ICMNH 13339^2. ECUADOR: Prov. Carchi: cabeceras del Rio Baboso, near Lita, MECN LAC 33, 74-75; Prov. Esmeraldas: Cachabi, 16 km SE Concepcion, 200 m, BMNH 1947.2.15.66-67. Prov. Pichincha: Centinela, 14.1 km SE Patricia Pilar. 550- 600 m, MCZ 97555, 97557, 97574; Centro Cientifico Rio Palenque, 47 km S 40 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 Santo Domingo de Ids Colorados. MCZ 88391. 90014-16. 92111-12. 94455. 94457-58: [Hotel] Tinalandia, 15-16 km SE Santo Domingo de los Colorados. 800 m. MCZ 88422, 90017. 90330-33. 91221; La Florida, near Alluriquin. MECN LAC 206. 208; Santo Domingo de los Colorados. 500 m, KU 109059. Eleuthewdactylus orpacohates ( 1 86) COLOMBIA: Depto. Antioquia: Mcpio. Frontino. Corregimiento de Nutibara, Km 14 carr. Nutibara a La Blanquita. 2000 m. ICNMHN 16495-96, Km 16 Nutibara a La Blanquita. 1960 m, ICNMHN 16472, 16579, Km 18 Nutibara a La Blanquita, 1940 m, ICNMHN 16475-86, Km 16.5-17 Nutibata a La Blanquita, 1900 m. ICNMHN 16473-74. Km 23 Nutibara a La Blanquita, 1430 m, ICNMHN 16487-90. Km 27 Nutibara a La Blanquita. 1140 m, ICNMHN 16492-94; Mcpio. Ituango, Antado, Quebrada Sabaletas, 1420 m, MHNCSJ 1922, 1925, 1929, 1933-34, 1940, 2382, 2384-85, 2388, 2393. 2395. 2598, 2603, 2605; Mcpio. Urrao, Vereda Calles, Parque Nacional Natural Las Orquideas. vie. cabana Rfo Calles. 1410-1525 m, ICNMHN 20275-77. Quebrada Aqudelo. 1410-1430 m. ICNMHN 20249-63. Quebrada de las Canoas. 1770-1870 m. ICNMHN 20264-74. Quebrada El Silencio. 1480-1540 m, ICNMHN 20244-48. Quebrada Vironda. 1400-1450 m. ICNMHN 20278- 82. Depto. Risaralda: Mcpio. Pueblorico. 15 km NE Pueblorico. vie. La Trinidad. 1600 m, ICNMHN 27501 ; Km 1^ carr. Pueblorico-Villa Claret, Quebrada San Jose, 1520 m, ICNMHN (MC 3379. 3403); Vereda Tatama. Rio Tatama. 1800 m, ICNMHN (MC 3531-34); carr. Pueblorico a Villa Claret, Vereda La Trinidad, near La Trinidad, 1510 m, ICNMHN (MC 3748, 3751 ). Depto. Valle del Cauca: Mcpio. La Cumbre, Vereda Chicoral, corregimiento de Bitaco, Finca La Cataisa. 1 900 m. ICNMHN 2 1 580, 2 1 582, 2 1 585-86; Mcpio. Darien. 1 .5 km W Lago Calima. Rio Calima. 1230 m. KU 168083-99. 168101-31; Mcpio. Yotoco, Reserva forestal de Yotoco, Km 18, Buga-Loboguerrero Road, 1590 m, ICNMHN (JDL 11129-75). Eleuthewdactylus permixtus (526) COLOMBIA: Depto. Antioquia: Mcpio. Angelopolis, MLS 167; Mcpio. Belmira: El Verbal, 6 km N Belmira. 2720 m. ICNMHN 8932-35; Vereda Los Patos, Quebrada Los Patos. 3.7 km N Belmira. 2620 m. ICNMHN 8936-37; Mcpio. Jerico: MCZ 24896-97; Mcpio. Medellin: Medellin. AMNH 38831-35, ICNMHN 4113; Medellfn Valley, AMNH 38784. 38787; Serrania de las Valdias, Boqueron. 2800-3000 m, ICNMHN 4278-81. 6353-57; Corregimiento San Felix. 2400 m. ICNMHN 14263-73; 5.6-5.7 km WSW San Felix. 2860-2940 m, ICNMHN 23329-33; 6.6-8.1 km WSW San Felix, 2820-3100 m, ICNMHN 8929-31; Mcpio. San Pedro: AMNH 38757, 38759-60, 38762, 38763-64, MCZ 24903-06 (reported as "Sampedro," Cochran and Coin, 1970), MLS 138. 143. 145, 400(3). 425(3), 426(2) ; Vereda La Lana. Finca La Montanita. ca. 4 km E El Tambo, 2510 m. ICNMHN 23334-38; Mcpio. Santa Rosa de Osos: AMNH 14073. 38817 (Cochran & Coin. 1970, reported this specimen as being from Santa Rosa de Aso, Depto. Caldas), 39456-58; 7.3 km S Llanos de Cuiva, 2770 m, ICNMHN 8915; El Chaquiro, MLS 69; Mcpio. Remedies: Finca El Amparo, MLS 454; Mcpio. Sonson: MLS 186; 8 km E Sonson, 2780 m, ICNMHN 8938- COLOMBIAN ELEUTHERODACTYLUS 41 42; 12-12.5 km E Sonson, 2540-2560 m, ICNMHN 8943^4; Paramo de Sonson, 17 km E Sonson, 2800 m. ICNMHN 18685-716, 18813; Paramo de Sonson. 22 km E Sonson. 2540 m, ICNMHN 18717; Mcpio. Yarumal: 3.5 km N Llanos de Cuiva, 2700 m. ICNMHN 8907-14; 5 km N Llanos de Cuiva. 2650 m, ICNMHN 8916-28; Ventanas. MLS 347, 445. Depto. Caldas: Mcpio. Manizales: 17 km E Manizales, 2520 m, ICNMHN 22491-97; between crest of Cordillera Central and 5 km toward Manizales. 3700 m, ICNMHN 2024. 2026. 4734; MLS 155 ('"Paramo Las Letras. Caldas." is probably from this locality); Mcpio. Marulanda; Alto de las Cruces. MLS 189a, 322-323; Mcpio. Pensilvania; MLS 424; Mcpio. Salamina; MLS 325-26. 328-29. Montanita. MLS 365. 372. Depto. Quindio; Mcpio. Calarca: Cerro Campanario, 1 km S Cajamarca-Calarca Road, 3240 m, ICNMHN 8896-8906; Mcpio. Salento; Cairetera Salento-Cocora. Hda. Alaska. Finca Rincon Santo. 6 km N road. UVC 6993-7007. 7322. Depto. Tolima; "Quindio Mountains," MCZ 8220-21. 8223; "Carretera Fresno- Manizales" (probably = Paramo Letras. Mcpio. Herveo), MLS 318-20; Mcpio. Cajamarca: Corregimiento Anaime, 3190 m, ICNMHN 22752; Corregimiento Anaime. 3480 m." ICNMHN 22601-17; "La Linea," Km 111-112 Carretera Cajamarca a Calarca. 3040-3080 m. ICNMHN 8861-95; "La Linea," Km 1 13- 109 Carretera Cajamarca a Calarca, 3070-3340 m, ICNMHN 22784, 22591- 600; Mcpio. Herveo; Paramo de Letras, 3050 m, ICNMHN 945, 948^9, 952, UVC 6759; Paramo de Letras, 0.5 km above "El Paisa," 3100 m, ICNMHN 635^0, 642^9, 651-53. 2581-83. 2585-92, 2594-95; Paramo de Letras, Vereda Albania. 3200 m. ICNMHN 760-65. 771. 774, 3187-96, 3491-93, 6226-28. 6230-35. 6242-48, 6250-52; Paramo de Letras, Km 77-79 Carretra Mariquita a Manizales. 3310-3340 m. ICNMHN 18826-37; 15 km W Padua, 2420 m, ICNMHN 22498; Mcpio. Ibague: Nevado del Tolima, Inspeccion de Policia Las Juntas a El Rancho. 2400-2750 m. ICNMHN 4782-93; 1 1 km N Juntas. El Silencio. 2600 m. ICNMHN 8849-60; along Rio Combeima. between El Silencio and El Rancho. 2590-2750 m. ICNMHN 8798-8828. 22618-87, 22813-28; El Rancho. 2750 m. ICNMHN 3256-60, 3262-65, 3267-69, 3272- 73; Nevado del Tolima, by road, above El Silencio, 2830-2980 m. ICNMHN 22829-32; 1.5 hr by foot above El Rancho. 2800-2820 m. ICNMHN 8829- 8848. Eleiithewdactylus supernatis (63) COLOMBIA; Depto. Cauca; Mcpio. Coconuco. Paletara, 3030 m. ICNMHN 8036- 52. 8064-67. between Paletara and Quebrada Bujios. 2960 m, ICNMHN 8073- 75. Quebrada Ullucos. 27 km SSE Coconuco. 2930 m. ICNMHN 8053-63, Rio Negro, 23 km SSE Coconuco, 2960 m, ICNMHN 8068-71. Vereda Bellavista, ca. 2000 m. UVC 5589; Mcpio. Piiez. Km 34-35, carretera Belalcazar-Tacueyo, 2265-2400 m. ICNMHN 6485. 8381, Km 40^1, carretera Belalcazar-Tacueyo, 2780-2820 m, ICNMHN 6427-28, 6451, 6480, 6510, 6512-13, 6607, 6984, 7063-64, Km 46-49.5, carretera Belalcazar-Tacueyo, 3200-3300 m. ICNMHN 6839^0, 6842^4. Depto. Huila, Mcpio. San Jose de Isnos, Chupallal de Perico, 29.4 km by road NW San Jose de Isnos. 2600 m. ICNMHN 803 1-32. 39 km by road NW San Jose de Isnos, 2880 m, ICNMHN 8033-35. For localities in Ecuador, see Lynch (1980). 42 UNIV. KANSAS NAT. HIST. MUS. OCC. PAP. No. 170 Eleuthewdactylus tamsitti (53) COLOMBIA: Depto. Caqueta: Mcpio. Florencia, Vereda Gabinete, 8.6 km E Alto de Gabinete, 2040 m, ICNMHN 22948, Vereda La Portada, 35.2 km NW Florencia, 1 230 m, ICNMHN 238 1 8, 37.4 km N W Florencia, 1 350 m, ICNMHN 23850-58, Vereda Tarqui, 12.5 km E Alto de Gabinete, 1750 m, ICNMHN 22949-51. Km 48-*9 carretera Altamira-Florencia, 1655 m, ICNMHN 23632, Km 53-54 carretera Altamira-Florencia, 1350 m, ICNMHN 23633-41, 38.8 km NW Florencia, 1 370 m, ICNMHN 238 1 9-23, 23848, 39.3 km NW Florencia, 1410 m, ICNMHN 23824-28, 41.1 km NW Florencia, 1470 m, ICNMHN 23829^7, 41.8 km NW Florencia, 1530 m, ICNMHN 23849. Depto. Huila: Mcpio. Acevedo, near San Adolfo, on Ri'o Suaza, 1400 m, FMNH 69735, 69737. 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