Frogs of the Eleutherodactylus Biporcatus Group (Leptodactylidae) of Central America and Northern South America, Including Rediscovered, Resurrected, and New Taxa

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3357, 48 pp., 21 ®gures, 2 tables March 26, 2002

Frogs of the Eleutherodactylus biporcatus Group (Leptodactylidae) of Central America and Northern South America, Including Rediscovered, Resurrected, and New Taxa

JAY M. SAVAGE1 AND CHARLES W. MYERS2

CONTENTS Abstract ...... 2 Resumen ...... 2 Introduction ...... 2 Methods ...... 3 Taxonomic History ...... 4 Rediscovery of Lithodytes gulosus Cope, 1875 ...... 6 Identity of Strabomantis biporcatus W. Peters, 1863 ...... 10 Application of the Name Lithodytes megacephalus Cope, 1875 ...... 14 Resurrection of Hylodes rugosus W. Peters, 1873 ...... 15 Systematic Synopsis: the biporcatus Species Group ...... 17 Key to Species ...... 17 Eleutherodactylus aphanus Campbell ...... 17 Eleutherodactylus biporcatus (W. Peters) ...... 21 Eleutherodactylus gulosus (Cope) ...... 24 Eleutherodactylus megacephalus (Cope) ...... 29 Eleutherodactylus opimus, new species ...... 34 Eleutherodactylus rugosus (W. Peters) ...... 38 Is the Eleutherodactylus biporcatus Group Monophyletic? ...... 40 Acknowledgments ...... 42 Appendix: Specimens Examined ...... 42 References ...... 45

1 Research Associate, Division of Vertebrate Zoology (Herpetology), American Museum of Natural History; Pro- fessor Emeritus, Biology, University of Miami. 2 Curator Emeritus, Division of Vertebrate Zoology (Herpetology), American Museum of Natural History.

ABSTRACT A revision of the broad-headed frogs of the biporcatus species group of Eleutherodactylus s.l. has a wholly unexpected nomenclatural consequence. Eleutherodactylus biporcatus (W. Peters, 1863) is not from ``Veragua'' (western Panama) as originally thought, but is the proper name for the Venezuelan frog heretofore known as E. maussi (Boettger, 1893). Three names are resurrected from synonymy for Central American species currently mas- querading under the misapplied name biporcatus, and a fourth species is described as new: (1) The rediscovery of Eleutherodactylus gulosus (Cope, 1875) shows it to be a large montane frog occupying an apparently small range in the borderland of Costa Rica and Panama. (2) Eleuth- erodactylus rugosus (W. Peters, 1863) is a smaller species occurring on the Paci®c versant of southwestern Costa Rica and western Panama; Lithodytes pelviculus Cope and L. ¯orulentus Cope are synonyms of E. rugosus. (3) Eleutherodactylus megacephalus (Cope, 1875), an intermediate-sized frog ranging from Honduras to central Panama, is the more common species to which the name biporcatus has usually been applied. Available material from the western half of the Isthmus of Panama was too sparse to decide if another (unnamed) species is being included under the name megacephalus. (4) The name biporcatus also has been used for Eleutherodac- tylus opimus, new species, which occurs from central Panama to western Colombia. Based on the condition of the m. adductor mandibulae, the Venezuelan Eleutherodactylus biporcatus s.s. (E. maussi, auctorum) belongs to the Middle American clade of Eleutherodac- tylus (subgenus Craugaster). However, preliminary data on karyotypes, as well as morpho- logical differences, cast doubt on the closeness of E. biporcatus to the other species studied. The monophyly of the ``biporcatus group'' therefore remains to be tested.

RESUMEN La revisioÂn de las ranas de cabeza ancha del grupo biporcatus de Eleutherodactylus tiene una consecuencia nomenclatural completamente inesperada. Eleutherodactylus biporcatus (W. Peters, 1863) no es de ``Veragua'' (PanamaÂ occidental) como se creõÂa previamente, sino que es el nombre propio de la rana venezolana conocida hasta ahora como E. maussi (Boettger, 1893). Se resucitan tres nombres de la sinonimia para especies centroamericanas previamente ocul- tos bajo el nombre incorrecto de biporcatus, y se describe una cuarta especie nueva: (1) El redescubrimiento de Eleutherodactylus gulosus (Cope, 1875) indica que es una rana grande, de montanÄa, que tiene una distribucioÂn geograÂ®ca pequenÄa en la frontera entre Costa Rica y PanamaÂ. (2) Eleutherodactylus rugosus (W. Peters, 1863) es una especie maÂs pequenÄa que ocurre en las tierras maÂs bajas del PacõÂ®co del suroccidente de Costa Rica y del occidente de PanamaÂ; Lithodytes pelviculus Cope y L. ¯orulentus Cope son sinoÂnimos de E. rugosus. (3) Eleutherodactylus megacephalus (Cope, 1875), una rana de tamanÄo intermedio y distribuida desde Honduras hasta PanamaÂ central, es la especie que maÂs comunmente se ha llamado biporcatus. El material disponible de la mitad occidental del Istmo de PanamaÂ no es adecuado para decidir si hay alguna otra especie (sin nombre) incluida bajo el nombre megacephalus. (4) El nombre biporcatus tambieÂn se ha usado para Eleutherodactylus opimus, especie nueva, que ocurre desde PanamaÂ central hasta Colombia occidental. BasaÂndose en la condicioÂn del m. adductor mandibulae, Eleutherodactylus biporcatus s.s. (E. maussi auctorum) de Venezuela pertenece al clado mesoamericano de Eleutherodactylus (subgenero Craugaster). Sin embargo, informacioÂn preliminar sobre cariotipos, asõÂ como di- ferencias morfoloÂgicas, hacer dudar del parentesco de E. biporcatus con las otras especies estudiadas. La mono®lia del ``grupo biporcatus'' debe ser corroborada.

INTRODUCTION spots in a dark reticulum. On the Atlantic side of Costa Rica, the little frogs are further In the lowlands and on the adjacent slopes characterized by a coloring of red or orange of lower Central America, some of the most on the posterior venter and undersides of the commonly seen leaf-litter anurans are small, hands and feet. These frogs have been re- broad-headed, diurnal, toadlike Eleuthero- ferred to as Eleutherodactylus biporcatus by dactylus that usually have large white ventral most recent authors following Dunn (1931). 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 3

However, the impressions gained of these an- imens from the Berlin Museum, through the imals based on diurnal observations of the courtesy of Dr. Rainer GuÈnther. small juveniles are illusionary, for the adults Serendipitously, the two lines of research are large (to 70 mm SVL), seldom-seen noc- converged on the occasion of the 62nd an- turnal predators. They may typically hide in nual meeting of the American Society of Ich- burrows during the day, and come to the sur- thyologists and Herpetologists at the Ameri- face at night to ambush passing prey, includ- can Museum of Natural History, in June ing large arthropods and even small frogs 1991, when conjoint examination of the Ber- and lizards. lin type material convinced us of the need Unlike most species in this huge genus of for this collaborative analysis. In its course, over 500 species, males seemingly do not vo- we examined extensive series of biporcatus- calize.3 As a result, adult males, which are like frogs, the type specimens of all nominal usually located by their sounds in other spe- Central American taxa thought to be syno- cies, are underrepresented in collections; at nyms of E. biporcatus, and the types of valid well-sampled sites, females are obviously related species. Our primary goal in the study disproportionately represented among adults. is to determine the systematic status, distri- Reproductive behavior is unknown, but eggs bution, and relationships of E. biporcatus are probably laid in the leaf litter where the and its allies and to establish a proper no- juveniles abound. menclature for included valid taxa. The present study has developed from two Our manner of presentation of the results different lines of research, each questioning of study departs somewhat from traditional the allocation of all lower Central American systematic papers where an a posteriori log- specimens of these broad-headed frogs to a ical framework is imposed on the not nec- single species. Savage has had a longtime essarily logical sequence of investigation. commitment to elucidating the composition Rather, as in the genre of detective ®ction, of, and relationships within the genus Eleuth- we have tried for the most part to present each clue leading to our conclusions in the erodactylus, especially in Central America. order of discovery. As in such tales, there For some time he had thought that more than were several real surprises and a few false one species was subsumed under the name leads as well. Eleutherodactylus biporcatus, but believed that a review of broad-headed frogs nomi- nally of that taxon from throughout its range METHODS was necessary to con®rm or reject this hy- Description of the external morphology of pothesis. Myers was particularly interested in the included taxa follows, where appropriate, the genus in the context of its species and the characters de®ned in Lynch and Duell- their distribution on the Panamanian Isthmus. man (1980, 1997) and Savage (1975, 1987). His own ®eldwork in western Panama had The presence or absence of paired dorsal led him also to conclude that more than one ridges (plicae) is important in the taxonomy species probably was represented by the of this group. For convenience, we use the monolithic E. biporcatus. In the process of term suprascapular plicae for the anterior beginning to resolve this problem, Myers ridges, even though they may extend poste- was able to borrow several critical type spec- riorly from the eyes to midbody in some species. One species has paravertebral plicae 3 This is an inference based on never having seen one positioned at midbody, separate from the su- calling. The males of all but one species in the group prascapular plicae. The posterior or latero- do lack vocal slits (hence vocal sacs), but absence of these structures is not a reliable predictor of voiceless- sacral plicae, when present, are often less ness. Vocal sacs have been lost in diverse species of well de®ned, sometimes comprising only frogs that, however, retain well-developed advertisement slightly raised, linear rows of small tubercles. calls (e.g., Myers, 1982: 4±5 [Dendrobates reticulatus]; Measurements were taken with a dial cal- Zweifel, 2000: 72±73 [Liophryne similis]). And voice in female frogs does not depend on such structures (see iper to 0.01 mm and rounded to the nearest Natural History in systematic section, under Eleuthero- 0.1 mm. Head-body length is indicated by dactylus biporcatus). the abbreviation SVL, which may be read ei- 4 AMERICAN MUSEUM NOVITATES NO. 3357 ther as snout-vent length or amphibian stan- time for much of what is now western Pan- dard length (distance from the tip of the ama, including today's provinces of Bocas snout to posterior margin of vent); for these del Toro, ChiriquõÂ, and Veraguas. Interest- frogs there is no practical difference between ingly, in the same paper he described an un- the de®nitions, inasmuch as the vent opens related Eleutherodactylus as Hylodes goll- in a vertical plane. Measurements, taken meri, based on two frogs purportedly from mostly by the ®rst author over a period of Caracas, Venezuela (lectotype ZMB 3168, time, involve specimens both poorly and well paralectotype ZMB 36301; originally iden- preserved. For this reason and, especially, ti®ed by Lichtenstein and von Martens because signi®cant population samples of [1856] as Hylodes martinicensis from ``Ca- adults are nearly lacking, we provide only vacas''). simple quantitative values without statistical Subsequent study (Rivero, 1961; Savage, analyses. 1987) suggested that an inadvertent exchange The color descriptions emphasize colors in life, which are based on our ®eld notes and of data may have occurred between the color slides. In the Systematic Synopsis these aforesaid type specimens of Hylodes goll- summarize variation in the most obvious meri and some types of Phrynidium (ϭ Ate- and/or distinguishing features. Most bright lopus) crucigerum Lichtenstein and von Mar- colors, especially reds, yellows, and greens tens (1856). Although a few extant ``types'' are lost more or less rapidly on preservation, of Phrynidium crucigerum are from the pub- while darker ones tend to be intensi®ed. Con- lished type locality ``Veragoa'', others are sequently the general pattern, uniform, spot- conspeci®c with a species (usually called ted blotched, striped, etc., is similar in life Atelopus cruciger) inhabiting the Cordillera and preservation. Other aspects of coloration de la Costa of Venezuela.4 On the same hand, and pattern markedly affected by preserva- the types of Hylodes gollmeri are represen- tion beside those just mentioned are indicat- tatives of a rather common but distinct Pan- ed where appropriate. amanian species of Eleutherodactylus.A

TAXONOMIC HISTORY 4 In a recent paper, LoÈtters et al. (1998) con®rmed that The frogs of the genus Eleutherodactylus specimens labeled (in ZMB) as syntypes of the name DumeÂril and Bibron, 1841 were referred to crucigerum are from both Panama and Venezuela. They various genera during the 19th and early 20th however selected a nonlabeled specimen (ZMB 3380) from ``Veragoa'' (western Panama) as the lectotype and century (G. Myers, 1962). Original species synonymized Phrynidium crucigerum with the Central descriptions and subsequent references to the American Atelopus varius.LoÈtters et al. proposed a sce- taxa discussed below used the names Hylo- nario whereby ZMB 3380±-a single specimen which has des Fitzinger, 1843, Lithodytes Fitzinger, ``no indication of type status'' collected by J. Warszew- icz±-must have been cataloged with at least four other 1843, and Strabomantis W. Peters, 1863, un- specimens that may subsequently have been lost or ex- til (and even after) Stejneger (1904) conclud- changed (crucigerum supposedly was based on ®ve ed that most species in the nominal Hylodes specimens from Veragoa). and Lithodytes should be placed in Eleuth- They asserted that Venezuelan specimens could not erodactylus. Many workers of this period ap- have been used for the original description because of a difference in skin texture. However, as is well known, plied the name Hylodes to all these frogs, but early authors commonly based descriptions on single Cope (1862: 153±154) divided that genus specimens drawn from whatever sample of ``cotypes'' into several groups, including Hylodes, with (syntypes) was at hand. typically granular venters, and Lithodytes, Considering the poor original description and all the assumptions made by LoÈtters et al. as to what is or might with smooth venters, a distinction no longer have been one of the original types, they might have recognized. better served stability by designating a syntype that was In 1863, Wilhelm Peters erected a new cataloged as such and that would have preserved the genus and species for a broad-headed frog well-known name Atelopus cruciger for the Venezuelan frog. The matter would now seem nomenclaturally dis- from ``Veragua'' that he called Strabomantis posed of, except for the arguable question as to whether biporcatus. Veragua was the name of a the ``lectotype'' was indeed part of the syntype series as Province of New Granada and used at that postulated by LoÈtters et al. 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 5 similar error in locality data associated with Cabello, on the Caribbean coast of Venezue- old Berlin Museum type specimens will be la.5 shown later in this paper to have serious im- Noble (1918: 329±331) used the name plications for the provenance and identity of Eleutherodactylus rugosus for examples of Strabomantis biporcatus. this group from Nicaragua and placed Lith- Peters (1873) later described a second odytes megacephalus and L. pelviculus in its broad-headed form as Hylodes rugosus from synonymy. He concluded that his ``large se- ``Chiriqui'' without reference to his earlier- ries of specimens shows conclusively that E. named Strabomantis biporcatus from Vera- megacephalus is but the adult of E. rugo- gua. The name ChiriquõÂ was used at that sus''. Noble retained L. gulosus as a valid time for western Veragua Province and is species of Eleutherodactylus similar to but essentially equivalent to today's Panamani- larger than megacephalus/rugosus. an provinces of Bocas del Toro and Chiri- Stejneger (1904) had revived usage of the name Eleutherodactylus for frogs placed in quõÂ. the Hylodes and Lithodytes of previous au- Shortly thereafter, Cope (1875) recognized thors, a course followed by most subsequent two additional broad-headed taxa, Lithodytes workers. Dunn (1931) used the name Eleuth- megacephalus and Lithodytes gulosus, col- erodactylus biporcatus for the Panamanian lected on W. M. Gabb's expedition to the species and, according to his unpublished southern Cordillera de Talamanca of Costa notes, regarded all names based on Central Rica. A few years later Cope (1877: 89) de- American material of the group available at scribed Lithodytes pelviculus, ``said to have that time to be synonyms, but he reluctantly been found on the west coast of Central continued to recognize E. gulosus as valid. America''. Boulenger (1882: 198±220) and Taylor (1952), in his monograph of Costa Brocchi (1882: 49±59) simultaneously Rican anurans, recognized Eleutherodactylus placed Strabomantis in the synonymy of Hy- rugosus (pp. 730±734) and E. ¯orulentus lodes, in which they recognized all ®ve of (pp. 765±766) as valid species. He accepted the broad-headed species mentioned above. Noble's (1918) synonymy of Lithodytes me- Cope later (1893: 336) maintained Lithodytes gacephalus with E. rugosus, but questioned for the three species that he had named (gu- whether L. gulosus and L. pelviculus should losus, pelviculus, megacephalus) and debe included there. scribed another (Lithodytes ¯orulentus) from Savage (1973, 1976, 1980) and Savage Boruca in the Paci®c drainage of southwest- and Villa (1986), in¯uenced by Dunn's un- ern Costa Rica. published synonymy, regarded all Central GuÈnther (1900±1901 [1885±1902]) treated American names for this group to be based biporcatus (p. 231), rugosus (p. 233), ¯oru- on specimens of E. biporcatus. Myers and lentus (p. 234), megacephalus (p. 239), and Rand (1969) and Rand and Myers (1990) fol- gulosus (p. 240) as distinct taxa of Hylodes. lowed Dunn (1931) in using E. biporcatus GuÈnther (op. cit.) also had intended to treat for Panamanian members of this group. Lithodytes pelviculus as a valid species of Lynch (1975), in the most recent review of the taxonomy of these frogs, likewise con- Hylodes, as shown by its inclusion as species cluded that these various names represented no. 23 in his key on page 227 (where it is only one species in Central America, namely misspelled ''petriculus'' [®de erratum on p. Eleutherodactylus biporcatus. He included xx]), but subsequently changed his mind and that species in his Eleutherodactylus bipor- allocated the name to the synonymy of H. megacephalus (p. 239). 5 A South American broad-headed frog unrelated to Nieden (1923: 432±439, 466) followed the biporcatus group but confused with it is the nominal GuÈnther's (1901) assignments for the Central Limnophys napaeus JimeÂnez de la Espada, 1870, which American species, and also picked up the authors have placed in the synonymy of Eleutherodac- name Hylodes maussi Boettger (1893) for a tylus biporcatus. Lynch (1975: 30±31) reviewed the evidence and allocated napaeus to the synonymy of South American broad-headed species appar- Eleutherodactylus cornutus in the sulcatus species ently related to this complex, from Puerto group. 6 AMERICAN MUSEUM NOVITATES NO. 3357 catus group along with E. cerastes, E. cor- within the genus. The South American clade nutus, and E. necerus of northwestern and is characterized by having a m. adductor upper Amazonian South America, and E. bu- mandibularis posterior subexternus present, foniformis of lower Central America and and the mandibular ramus of the trigeminal northwestern South America, but he placed nerve (V3) passing external (lateral) to the the Venezuelan E. maussi in the separate sul- muscle (``S'' condition). The Central Amer- catus group. This grouping was reiterated in ican clade (subgenus Craugaster Cope, Lynch's (1976) discussion of the South 1862: 153) has a m. adductor mandibularis American species groups of Eleutherodacty- externus super®cialis, and the trigeminal lus. nerve passing mediad (internal) to the mus- Lynch (1975, 1976) regarded the forms in cle (``E'' condition). This conclusion re- his E. biporcatus group as closely related quired a complete renovation of the two since they shared the features of very broad broad-headed species groups, because only heads (head width Ͼ 42% SVL), ®nger I Ͼ Eleutherodactylus biporcatus (sensu lato), E. ®nger II, and smooth venters. Lynch (1975) bufoniformis, E. ¯orulentus, E. necerus, and indicated that another cluster of broad-head- E. maussi have the E condition; these were ed South American Eleutherodactylus, the assigned to the biporcatus group. Except for sulcatus group, was somehow related to the E. maussi, the remaining South American biporcatus group, but in the 1976 paper he broad-headed forms all have the S condition placed the two groups in different infrage- and currently are placed in the sulcatus neric units within the genus. group.6 Subsequently, Lynch (1981) expanded the In 1980, Jonathan A. Campbell collected sulcatus group to include the nominal mono- several biporcatus-like frogs from the moun- typic genus Amblyphrynus (Cochran and tains of eastern Guatemala. Savage examined Goin, 1961) and a new form to bring its com- this material in 1980 and expressed his opin- position to ®ve species: Eleutherodactylus ion that it was doubtless a diminutive new helonotus (Ecuador), E. ingeri (Colombia), member of the biporcatus group. Campbell E. maussi (Venezuela), E. ruizi (Colombia), (1994), after obtaining additional material, and E. sulcatus (Amazonian Ecuador and described the species as Eleutherodactylus Peru). aphanus. Savage and DeWeese (1979, 1981) and Lynch and Duellman (1997), in a major Savage (1984, 1987), while acknowledging treatment of Eleutherodactylus relationships, the utility of Lynch's (1976) attempt to bring placed Eleutherodactylus bufoniformis and some kind of order to the classi®cation of E. necerus in a separate bufoniformis species Eleutherodactylus, criticized Lynch's system group, leaving the biporcatus group with as essentially phenetic and based on trivial three recognized species (aphanus, biporca- external features much given to homoplasy. tus, maussi). We accept the biporcatus and They preferred to emphasize the condition of bufoniformis species groups for purposes of jaw muscles and karyology over ®nger the present study, which focusses on the ®rst length and ventral texture. group. Lynch (1986) responded to this challenge with a thorough examination of the jaw mus- REDISCOVERY OF LITHODYTES cle characters of the m. adductor mandibulae GULOSUS, COPE, 1875 of Eleutherodactylus, ®rst discovered by Starrett (1968) and touted by Savage and In February of 1976, Myers collected a se- DeWeese (1979, 1981) as systematically in- ries of unusual juvenile Eleutherodactylus formative. Lynch's work has broad signi®- (®g. 1) in leaf litter near the site of the pre- cance to the systematics of all Eleutherodac- tylus and their allies, but only those aspects 6 The sulcatus species group now includes nine spe- relating to the present paper are summarized cies according to the latest revision (Lynch, 1997). It is a South American assemblage. Only one species occurs here. (barely) in political Central America, namely E. laticor- The principal result of Lynch's 1986 study pus Myers and Lynch (1997) from the Panamanian-Col- was the recognition of two major clades ombian frontier. 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 7

this enigmatic frog. Found at elevations between 1000 and 1220 m, on and near the continental divide, the adults were distinct from lower-elevation E. biporcatus in lacking dorsal plicae and in details of coloration. The question then naturally arose as to whether these frogs represented an undescri- bed species or were conspeci®c with one of the several taxa previously named from western Panama and southern Costa Rica, but then regarded as junior synonyms of E. biporcatus. Crucial to resolving this question was examination of the type specimens of each available name. Consequently, Myers borrowed the type series of Strabomantis biporcatus and the holotype of Hylodes rugosus from the Berlin Museum. A preliminary comparison of these animals with the newly collected material left him with little doubt Fig. 1. The ®rst specimen collected during the that the Fortuna frogs could not be associated rediscovery of Lithodytes [Eleutherodactylus] gu- with either of the species represented by the losus in 1976Ða century after its original descrip- Berlin types. tion (Cope, 1875)Ðat the proposed Fortuna Dam But Myers was astonished to ®nd that the site in western Panama. A juvenile female type series of S. biporcatus represented a (AMNH 95035) of 32 mm SVL, less than half species different from the frogs usually grown; note the pustulate dorsum, a juvenile con- called E. biporcatus for more than half a cen- dition (compare adult in ®g. 12). [From a trans- tury, and that they were also unlike any other parency by C. W. Myers.] known species from Central America. This puzzlement led to consultation with Savage on the occasion mentioned in the introduc- sent-day Fortuna Dam (then being surveyed tion to this paper, with the latter fully agree- for construction), at 1120 m, on the Paci®c ing with Myers' conclusions that the Fortuna versant of western Panama (®g. 2). Although frogs: (1) were not conspeci®c with the spe- reminiscent of the common broad-headed ju- cies usually called E. biporcatus in Central veniles identi®ed with Eleutherodactylus bi- America; (2) were not conspeci®c with the porcatus, they differed from them most ob- types of S. biporcatus or H. rugosus; (3) and viously in lacking well-de®ned dorsal plicae that the types of S. biporcatus were not rep- and in having a different ventral coloration. resentatives of the species usually referred to In addition, the dorsum was conspicuously as E. biporcatus in Central America! covered by numerous small pustules. Direct Further discussion led us to the hypothesis comparison with juvenile E. biporcatus of that the Fortuna frogs might well be exam- the same size strongly suggested that these ples of one of the other of Cope's nominal frogs were not conspeci®c with typical low- species, Lithodytes gulosus or L. megace- land E. biporcatus.7 phalus, also from an upland area in the same Later visits to nearby areas by Myers and mountain range as Fortuna but on the Atlan- John Daly in July of 1982 and January 1983 tic slope in southern Costa Rica. The possi- led to the capture of two large females, one bility also existed that both these names were subadult female, and additional juveniles of based on a single taxon conspeci®c with the Fortuna examples. 7 Jungfer (1988: 16) later used the name Eleuthero- Cope's (1875: 110±112) original descrip- dactylus biporcatus for a juvenile specimen from the Fortuna regional population, but his accompanying pho- tions clearly differentiated the holotypes of tograph represents a species (E. megacephalus) from an- Lithodytes gulosus and L. megacephalus as other locality. follows: 8 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 2. Views of the Fortuna Dam site in the upland valley of the RõÂo ChiriquõÂ, in 1976, prior to road and dam construction (access was by trail or helicopter). Upper: Survey and research camp of the Instituto de Recursos HidraÂulicos y Electri®cacioÂn (IHRE) above RõÂo ChiriquõÂ (at right), on northern base of Cerro Fortuna, about 1000 m above sea level. Lower: View looking westward across the subsequently ¯ooded RõÂo ChiriquõÂ to Cerro Pinola (center), from 1060 m on north slope of Cerro Fortuna. [Photographs by C. W. Myers, March 2±3, 1976.] 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 9

The types of both putative species are gulosus with the two adult females (AMNH housed at the United States National Muse- 124370, AMNH 124372), 84 and 72 mm um of Natural History and are part of the W. SVL, respectively, and a subadult female M. Gabb collection made in Costa Rica in (AMNH 124371, 46 mm SVL) from the For- 1873±1874. The holotype of both Lithodytes tuna Dam area of Panama leaves little doubt gulosus (USNM 32590) and of Lithodytes that they represent the same species. The megacephalus (USNM 32579) are from most striking similarity is the absence both ``Pico Blanco'', LimoÂn Province, Costa Rica, of the suprascapular plicae and the laterosa- 6000 ft. However, as pointed out by Savage cral tubercle and/or ridge system. The plicae (1970), Gabb's expedition did not ascend are characteristic of all other named forms of Pico Blanco (ϭ Cerro Kamuk), but actually biporcatus-like frogs, and laterosacral tuber- climbed to the summit of Cerro Utyum, the cles or ridges are found in some of the other next peak to the northwest. species of the group (®g. 3). The type specimen of Lithodytes gulosus In other features, the adult females from is in excellent condition and agrees well with Panama agree closely with the holotype and Cope's original description. Unfortunately Cope's description of Lithodytes gulosus in the type of Lithodytes megacephalus has suf- having a uniform dark dorsum and posterior fered serious decay and today consists only thigh surface. In addition they have the chin of skin fragments and disarticulated bones. and throat pale or gray-brown, pale venters The maceration of this specimen is relatively mottled with gray-brown or with dark brown recent, as Noble (1918) and Dunn (unpub- spotting, and the seat patch is inconspicuous. lished notes) examined it while it was still The subadult female differs from the oth- whole. Noble regarded the specimen as a ers in some features of coloration, including large adult conspeci®c with Eleutherodacty- distinct transverse dark bars on the limbs, lus rugosus and slightly different from the some pale tan spotting on the posterior thigh type of L. gulosus, which he retained as surface, the dark seat patch mark restricted Eleutherodactylus gulosus. Dunn (1931) ap- to the area around the vent, the chin and parently considered both names as synonyms throat gray with small white spots, the un- of Eleutherodactylus biporcatus, the types of derside of the leg suffused with yellow, and which he had examined in Berlin in 1928. the venter, groin and anterior thigh surface Although the condition of the holotype of marbled blackish gray on white (life colors). Lithodytes megacephalus makes direct com- This specimen is somewhat intermediate in parisons impossible, Cope's (1875) descrip- coloration and dorsal pustulation between the tion and illustration (reproduced herein as large adults and the series of juveniles. ®g. 6) clearly demonstrate that it is not con- Unlike adults, juveniles (AMNH 95035± speci®c with the adult Fortuna frogs. Unlike 95046, 124365±124369; 18±32 mm SVL) them it had well-developed suprascapular have dorsally pustulate skin, a lighter dorsum and laterosacral ridging, the posterior thigh with obscure darker mottling, and greenish surface was black, marbled distally with ash, ventral coloring; juveniles also differ notably and the dark seat patch was continuous onto from adults in that traces of suprascapular the lower posterior surface of the thigh. plicae can be discerned on close examination Comparison of the holotype of Lithodytes in most specimens (®g. 13). A juvenile male 10 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 3. Diagrammatic representation of dorsal-ridge patterns in the Eleutherodactylus biporcatus group. A: E. gulosus, no de®nite dorsal plicae in adults (although weak, short anterior suprascapular plicae are discernible in most juveniles; see ®g. 13). B: E. biporcatus s.s. (maussi auctorum), paired suprascapular and weak laterosacral plicae. C: E. rugosus, paired suprascapular plicae, no laterosacral plicae. D, E: E. megacephalus has chalice-shaped or hourglass-shaped suprascapular plicae, with geographic variation in the laterosacral plicae or linear series of closely set tubercles; the V-shaped laterosacral pattern (D) is common from Nicaragua through Costa Rica, tending to be replaced in western Panama by nonconverging (E) parallel ridges. Two additional species, Eleutherodactylus aphanus (®g. 8) and E. opimus (®gs. 17±19), resemble E. rugosus (C) in lacking paired laterosacral plicae or tuberculate ridges, although these species are overall much less tuberculate than rugosus (®g. 20).

(KU 115221, 28 mm SVL) from the Atlantic found in allied forms (see Systematic Syn- versant of Panama (juncture of RõÂo Claro and opsis) further supporting recognition of RõÂo Changena, Bocas del Toro, 910 m) also Eleutherodactylus gulosus (Cope, 1875) as a appears to be conspeci®c with the holotype valid species. and Fortuna Dam series; it has a pustulate dorsum and resembles the subadult female in IDENTITY OF STRABOMANTIS coloration. BIPORCATUS W. PETERS, 1863 We conclude that there can be little doubt that the Costa Rican holotype of Lithodytes The syntypic series of Strabomantis bi- gulosus and recent specimens from western porcatus consists of four specimens, ZMB Panama represent a species distinct from all 3330 (3 adults) and ZMB 3322 (a juvenile), related broad-headed Eleutherodactylus;itis all purportedly from ``Veragua''. The adults, the only one in which dorsal ridging is com- hereafter referred to as ZMB 3330A, B, and pletely lacking (adults) or very vague (ju- C, are about 67, 60, and 55 mm SVL and veniles). In addition, this species attains a the juvenile is 32 mm (®g. 4). much larger size than any other broad-head- These frogs resemble the several broad- ed form (table 2). Features of juvenile and headed Eleutherodactylus described from adult coloration are also different from those Central America in having cranial crests on 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 11

Fig. 4. The type series of Strabomantis biporcatus W. Peters, 1863, in dorsal and ventral view. Left to right: ZMB 3330[A]&, 3330[B]& (lectotype), 3330[C](sex? ), 3322 juv. The type locality was given as ``Veragua'' (western Panama), but based on current knowledge, these specimens must have been collected in Venezuela. 12 AMERICAN MUSEUM NOVITATES NO. 3357

TABLE 1 Measurements (in mm) Made a Century Apart on the Two Largest Syntypes of Strabomantis biporcatus Peters, 1863

the frontoparietal bones, and resemble all ex- volved two other names based on Berlin Mu- cept E. gulosus in the presence of well-de- seum specimens: Phrynidium crucigerum veloped suprascapular plicae. They differ Lichtenstein and von Martens 1856, and Hy- most obviously from any broad-headed lodes gollmeri W. Peters, 1863, originally de- Eleutherodactylus known from the region in scribed from Veragoa and Caracas, respec- having a tarsal fold, greatly enlarged, conical tively. Subsequent research (Rivero, 1961; subarticular tubercles under the ®ngers, and Savage, 1987) demonstrated that some of the de®nite plantar tubercles. syntypes of the former Atelopus cruciger and Because of these discrepancies, we consid- the type of Eleutherodactylus gollmeri were ered the possibility that some mix-up in data conspeci®c with a Venezuelan and a Central had occurred, given the former custom at the American species, respectively.8 Berlin Museum of having the identi®cation Aided by this clue, we compared the syn- and specimen number on a slip of paper types of Strabomantis biporcatus with placed in the bottle but unattached to the Eleutherodactylus maussi Boettger, 1893, the specimens. However, direct comparison of only known broad-headed species of the ge- the syntypes with Peters' (1863) description nus in Venezuela. And indeed, specimens revealed no essential difference between from a series (AMNH 70533±70552, 70554± them. Peters gave measurements for the larg- 70558) of E. maussi, along with the rede- est examples, which he listed as ``A'' and scription of that form by Lynch (1975), ``B'', and we remeasured the largest syntypes agreed in every signi®cant feature with the (ZMB 3330A, 3332B) as shown in table 1. syntypes of Strabomantis biporcatus (®g. 5). In view of the length of time in preser- Signi®cant characters included: large size vative and the present softened condition of (adult females about 49±74 mm SVL), the specimens, these measurements are in reasonable agreement. This, in combination 8 But as already discussed (footnote 4) LoÈtters et al. (1998) chose a presumed syntype of Phrynidium cruci- with the close concordance between other gerum from Panama as ``lectotype'' and placed the name characteristics of these specimens and the in the synonymy of Atelopus varius, unfortunately leav- original description, convinces us that they ing the name cruciger as unavailable for Venezuelan are the syntypes of Strabomantis biporcatus populations. (Added in press: LoÈtters and La Marca [Bull. Zool. Nomencl. 58: 119±121, June 2001] have as labeled. At this point, we recalled the sit- now petitioned to conserve the name Atelopus cruciger uation described above under Taxonomic through selection of a neotype, collected in 1930, that History, where erroneous locality data in- is not part of the original type series.) 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 13

Fig. 5. The South American frog heretofore known as Eleutherodactylus maussi (Boettger, 1893). Left to right: AMNH 70539, 70552, 70556, adult females from Rancho Grande in the Coastal Range of northern Venezuela. Compare with the syntypes of the earlier named Strabomantis biporcatus W. Peters, 1863, in ®gure 4. slightly arched vomerine odontophores, a su- gesting that he was not differentiating coarse prascapula and paravertebral ridge system, granules from other small knobs or tubercles laterosacral tubercle or ridge system promi- found on the body. nent to absent, conical subarticular tubercles Although the three adult syntypes super- under the ®ngers, an inner tarsal fold, weak ®cially appear to have smooth ventral sur- toe fringes, low inner and prominent outer faces, close examination reveals traces of metatarsal tubercles, and planter tubercles coarse granulation on the venters. The skin present. on the throat is smooth in the largest speci- At ®rst glance, the only apparent point of men, as stated by Peters, but is coarsely gran- disagreement between the syntypes of Stra- ular on the other two. The juvenile has an bomantis biporcatus and de®nitive Eleuth- obviously coarsely-granular venter but a erodactylus maussi involved the texture of smooth throat. The condition of ventral gran- the ventral integument. Eleutherodactylus ulation frequently is distorted by preserva- maussi is characterized by having the ab- tion, especially if the frog was originally dominal surface coarsely granular or areo- poorly ®xed in weak alcohol (modern for- late. In his original description of S. biporcatus, Peters (1863: 406) noted that ``der malin preservation also often tends to dimin- KoÈrper erscheint glatt, ist aber bei genauerer ish granulation). Our judgement is that all the Betrachtung allenthalben mit kleinen syntypes had the venters areolate (coarsely WaÈrzchen bedeckt . . . Eben so zeigt die granular) in life. Bauchseite kleine WaÈrzchen, waÈhrend die Although no collector is identi®ed by Pe- Kehle glat ist''. [The body seems smooth, al- ters (1863) in the original description, the though . . . very small knobs can be discov- Berlin Museum catalog lists the source as ered . . . The ventral side exhibits no small Warszewicz, who also was responsible for knobs, while the throat is smooth.] Peters Panamanian syntypes of Atelopus cruciger,a mentioned that the kleinen WaÈrzchen (small name also applied to a Venezuelan frog warts, tubercles, or knobs) were denser and (footnotes 4, 8, and associated text). Curator somewhat larger on the posterior thigh, sug- Rainer GuÈnther provided the following in- 14 AMERICAN MUSEUM NOVITATES NO. 3357 formation (in letter to Myers, September 27, odytes pelviculus Cope, 1878; and Lithodytes 1995): ¯orulentus Cope, 1893. As we will show in a later section, the [Some of] the types of Atelopus cruciger were indeed collected by the same collector named Warszewicz holotypes of the ®rst, third, and fourth names . . . as [were] the types of Strabomantis biporcatus are based on an essentially allopatric Paci®c . . . In our archives we found a report by Lichtenstein population con®ned to the humid forests of saying that he (obviously personally) purchased from southwestern Costa Rica and adjacent south- Warszewicz ``6 GlaÈser mit verschiedenen Tieren in western Panama. Our evidence establishes Weingeist, die er auf der Cordillera von Veragoa in bisher noch gar nicht besuchten ThaÈlern gesammelt this population as a valid species. This raises zu haben erklaÈart''. From the listed specimens (some the questions as to whether the name Lith- clearly with marine origin) however it becomes clear odytes megacephalus is an available name that Warszewicz obtained his material not only in based on a specimen from an Atlantic slope Veragoa (sometimes Veragua) but also from else- where. Some archivist noted on the deed of Warszew- population in Central America. icz already: Panama, Brasilien and Peru.... From As previously pointed out, the holotype of our documents it follows further that most (?) of the Lithodytes megacephalus (USNM 32579)9 Warszewicz collection was sold by auction in the from the Atlantic versant of Costa Rica is in British Museum . . . In our herpetological collections Catalogue all the specimens obtained from Warszew- such a deteriorated condition as to defy fur- icz have as collection site Veragoa or Veragua, which ther study. Nevertheless, the original descrip- after all is not always correct. tion and ®gure (Cope, 1875: 110±113, p1. 23, ®g. 11, 11a) leave little question that it The agreement of the syntypes of Stra- bomantis biporcatus and representatives of represents Atlantic versant frogs long re- the nominal taxon Eleutherodactylus maussi ferred to as Eleutherodactylus biporcatus. in all diagnostic features forces us to con- The principal features supporting this conclu- clude that biporcatus is not a Central Amer- sion are the presence of well-developed su- ican frog as long assumed. The proper name prascapular and laterosacral ridging (®g. 6), for the Venezuelan species must be Eleuth- the posterior thigh surface black, marbled erodactylus biporcatus (Peters, 1863) under with ash, with the black pigmentation essen- the principle of priority (International Com- tially continuous to the vent (®g. 6), the light mission on Zoological Nomenclature, 1999: dorsal color, and the size (70 mm SVL). All art. 23). The name maussi is little used, and other features described or ®gured by Cope its replacement will not lead to signi®cant are consistent with this conclusion. In addi- destabilization in nomenclature. tion, Noble (1918) and Dunn (1931) both examined the holotype while it was in good condition and concluded that it was conspe- APPLICATION OF THE NAME ci®c with frogs ®tting their concepts, respec- LITHODYTES MEGACEPHALUS COPE, tively, of Eleutherodactylus rugosus and E. 1875 ``biporcatus'' in Central America. The discovery that Strabomantis biporca- A problem with this interpretation centers tus is a senior subjective synonym of Hylo- on the elevational data associated with the des maussi not only requires that the Vene- holotypes of both Lithodytes megacephalus zuela species be known as Eleutherodactylus and Lithodytes gulosus. Cope stated that the biporcatus, but has another major taxonomic former was collected ``on a spur of Pico repercussion as well. For 70 years, following Blanco, at 6000 feet elevation'' and the latter Dunn (1931), the name Eleutherodactylus bi- ``from the same locality as the last species.'' porcatus has been applied almost universally As previously pointed out (Savage, 1970), to common lowland broad-headed frogs that the Gabb expedition that collected both spec- occur from Honduras to Colombia. With the imens actually climbed Cerro Utyam, not recognition that Eleutherodactylus gulosus is a valid upland species, four other names are 9 Savage (1970) either through a lapsus calami or ty- pographical error listed Cope's type of this species as potentially applicable to the lowland frogs. Hylodes megalocephalus. This stands as an incorrect These are: Hylodes rugosus Peters, 1873; subsequent spelling of the species name, and it has no Lithodytes megacephalus Cope, 1875; Lith- status in nomenclature. 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 15

Fig. 6. Original illustration of the holotype of Lithodytes megacephalus, reproduced same size from Cope's (1875) ®gures 11 and 11a on plate 23 (ϭ pl. ``1'' in ®g. legends on page facing pl. 23).

Pico Blanco (Cerro Kamuk), the next peak types are from the same locality (Cerro to the southeast. Utyam) in a loose sense, they probably were The elevation of 6000 ft (1829 m) is well not collected at the same place. above the known limit for nominal Eleuth- In spite of the perceived dif®culty associ- erodactylus ``biporcatus'' on the Atlantic ated with the elevational data, we are con- slope. On the other hand that elevation may vinced on the basis of available evidence that well be within the range for Eleutherodac- the name Eleutherodactylus megacephalus tylus gulosus, which we now know occurs at (Cope, 1875) is the valid name for many least between 1000 and 1220 m. We do have frogs referred previously to E. biporcatus some question as to whether they both are (sensu Dunn) in lower Central America. found as high as 1830 m but it is possible Even so, as demonstrated in this paper, the that they are marginally sympatric at a lower previous ``biporcatus'' auctorum was a com- elevation. posite of at least three species. Even as re- It is not altogether clear to us from the stricted herein, Eleutherodactylus megace- original descriptions that Cope's statement phalus may still be a composite (see species that the holotype of Lithodytes gulosus being account under Remarks). ``from the same locality. . . '' as the type of L. megacephalus meant anything more than RESURRECTION OF HYLODES that they were both collected on Cerro RUGOSUS W. PETERS, 1873 Utyam. However, the USNM catalog lists both as being from ``Pico Blanco, Costa The holotype of Hylodes rugosus (ZMB Rica, 6000 ft.'' Savage (1970) also listed 7812; ®g. 7) is a subadult, 21 mm SVL, hav- both as from ``spur of Cerro Utyam, 6000 ft. ing prominent suprascapular ridging, no la- (1830 m)''. We now suspect that Gabb's ®g- terosacral rows of tubercles, a pair of distinct ure of 6000 feet is an overestimate of the broad dorsal stripes, vivid black and light elevation where the specimen of L. megace- thigh bars, and a contrasting pattern of light phalus was taken, and that while both holo- and black groin markings. In all these fea- 16 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 7. Holotype of Hylodes rugosus W. Peters, 1873. ZMB 7812 in dorsal and ventral view, a juvenile from ``Chiriqui'', western Panama. Scale divisions ϭ mm. tures, which in combination differentiate it and groin markings are a particularly distinc- from Eleutherodactylus gulosus and E. me- tive feature. gacephalus, it closely resembles samples of The Golfo Dulce series and E. megace- broad-headed frogs of this complex from the phalus most closely resemble one another Golfo Dulce region of southwestern Costa within the biporcatus group in general hab- Rica and adjacent Panama. Although the type itus and in having well-developed suprascap- locality ``Chiriqui'' is ambiguous because it ular plicae. In the Golfo Dulce frogs, most was applied to both Atlantic and Paci®c ver- individuals have a well-developed medially sant Panama at the time of the species' de- directed branch from the anterior arm of each scription, the holotype is obviously from the suprascapular ridge, but this is never seen in Paci®c slope population. Examination and E. megacephalus. In addition, the upper sur- comparisons with material from the Golfo face of these frogs is covered by many rel- Dulce of the holotypes of Lithodytes pelvi- atively large wartlike pustules, while the pus- culus Cope, 1878 (26 mm SVL) and the de- tules present in juvenile E. megacephalus are scription of Lithodytes ¯orulentus Cope, small and tend to be pointed, with the dor- 1893 (type lost but 35 mm SVL) show that sum of adults being smooth to shagreened these names are synonyms of L. rugosus. All with only a few widely scattered pustules. examples from southwestern Costa Rica Eleutherodactylus megacephalus also has share the features mentioned above except paired laterosacral series of pustules that are that the light (reddish brown in life) dorsal usually fused into ridges, which are lacking stripes are occasionally absent. The vivid in Golfo Dulce material. Most frogs in the black and light (scarlet in life in adults) thigh last sample have distinctive paired broad dor- 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 17 sal stripes, which are never present in E. me- occasional males (18 pairs of chromosomes), gacephalus. and 2N ϭ 35 in most males (16 pairs and 3 Since the Golfo Dulce population is com- unpaired chromosomes); N.F. ϭ 42. See ®g- pletely allopatric to those here assigned to ure 21 and closing discussion. Eleutherodactylus megacephalus, and there is no evidence of intergradation in the char- KEY TO SPECIES acteristics distinguishing them, we conclude 1. Venter smooth; no tarsal fold; no vocal slits that two species are represented. Conse- in adult males; dorsal plicae present or ab- quently we recognize Eleutherodactylus ru- sent ...... 2 gosus (Cope, 1893) of the Paci®c slope of Venter coarsely areolate; an inner tarsal fold southwestern Panama and adjacent Costa present; vocal slits in adult males; paraver- Rica as a valid species within the biporcatus tebral plicae at midbody separate from su- group. prascapular plicae (®g. 3B)...... E. biporcatus 2. Dorsum of adults and juveniles with distinct SYSTEMATIC SYNOPSIS: THE paired suprascapular plicae (®gs. 3C±3E, BIPORCATUS SPECIES GROUP 18)...... 3 Dorsum of adults lacking suprascapular pli- INCLUDED SPECIES: Eleutherodactylus cae (®g. 3A), but characteristically con®g- aphanus Campbell, 1994; E. biporcatus (W. ured, weakly developed anterior plicae pre- Peters, 1863); E. gulosus (Cope, 1875); E. sent in most juveniles (®g. 13) ...... megacephalus (Cope, 1875); E. opimus, new ...... E. gulosus species; and E. rugosus (W. Peters, 1873). 3. Posterior dorsum lacking laterosacral system See table 2 for a tabular comparison of the of either V-shaped or parallel plicae or included species. closely set tubercles (e.g., ®gs. 3C, 18) . . DEFINITION: Robust, toadlike eleutherodac- ...... 4 tylines with broad heads (head width 39± Posterior dorsum with a de®nite laterosacral system of either V-shaped (®g. 3D) or non- 55% of SVL in adults, usually Ͼ 42%) hav- converging (®g. 3E) parallel rows of tuber- ing paired frontoparietal crests that are visi- cles or tuberculate ridges ...... ble externally in adults of most species; head ...... E. megacephalus outline from above semicircular or ovoid to 4. Posterior surface of thighs not contrastingly broadly rounded; upper eyelids tuberculate; marked with black and red (or white), al- tympanum external, distinct, annulus tym- though sometimes barred in black with tan panicus prominent, round in males, ovoid interspaces; dorsum smooth or with scat- and higher in females; supratympanic fold tered low pustules ...... 5 from eye towards arm; no inguinal glands; Posterior surface of thighs contrastingly no male nuptial pads; vomerine odonto- marked with black bars, separated by red interspaces (white in juveniles) in life; dor- phores triangular, or slightly arched lying sum strongly tuberculate with numerous posterior and medial to large choanae, the large warts and ridges...... patches narrowly separated medially; tibia ...... E. rugosus 46±60% of SVL; disks present on all digits, 5. Venter immaculate; size small, adult males not expanded on ®ngers, slightly expanded 18±22 mm SVL, adult females 35±44 mm on toes; ®ngers short, slender; relative length ...... E. aphanus of appressed ®ngers III Ͼ I Ն II ϳ IV; basal Venter variously marked with brown and/or toe webbing absent or very slight; relative large light spots; size moderate to large, length of appressed toes IV Ͼ III Ͼ V Ͼ II adult males probably Ͼ30 mm SVL, adult Ͼ I, the third much longer than ®fth; jaw females to at least 69 mm ...... E. opimus muscle formula DFSQAT ϩ e. KARYOLOGY:2Nϭ 20; N.F. ϭ 36 in two examined species (Eleutherodactylus mega- Eleutherodactylus aphanus Campbell cephalus and E. opimus), and 2N ϭ 20; N.F. Figures 8, 9 (map) ϭ 38 in two other species (E. gulosus and E. Eleutherodactylus aphanus Campbell, 1994: rugosus). In Eleutherodactylus biporcatus, 296±302, ®gs. 1±3. Holotype: UTA A-33405, the karyotype is 2N ϭ 36 in females and an adult male from 12.0 km WSW Puerto San- 18 AMERICAN MUSEUM NOVITATES NO. 3357

TABLE 2 Characteristics of the Eleutherodactylus biporcatus Groupa 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 19

TABLE 2 Continued.

to TomaÂs, Department of Izabal, Guatemala, no accessory palmar tubercles; 2 to several 786 m. ¯at ulnar tubercles; heel with 1±3 small tu- DIAGNOSIS: This, the smallest member of bercles; sole smooth; no de®nite lateral ridg- the biporcatus group (®g. 8), is distinguished es or fringe on toes; inner metatarsal tubercle from its allies by the combination of an im- oval, outer tubercle much smaller, round; no maculate venter, presence of suprascapular tarsal fold; prominent outer tarsal tubercles plicae, and lack of laterosacral ridges or tu- in juveniles, weak to well-developed in bercle series. Eleutherodactylus aphanus is adults. most similar to E. opimus, but differs in ven- COLORATION: Adult males olive-brown, tral coloration, and aphanus may also differ pale brown, orange or yellowish orange in having weakly keeled digits (Campbell, above; often with peach or orange markings 1994: 298). on side of head; posterior thigh surface gray MORPHOLOGY: Adult males 18±22 mm with white mottling; venter yellow to bright SVL, adult females 35±44 mm (mean SVL chartreuse; adult females brown to gray ϭ 19.8 mm, 4 (, 40.2 mm, 3 &); tibia 48± brown above; posterior thigh surface dark, 58% of SVL; head width 45±50% of SVL; almost black with pale lavender mottling; cranial crests present, visible externally in venter purplish, except immaculate cream in some adult females; dorsal surfaces basically juveniles. smooth; de®nite suprascapula plicae forming NATURAL HISTORY: The only information a chalice- or hourglass-shaped system; ¯anks is provided in Campbell's original descrip- smooth; no de®nite laterosacral system of tubercles or ridges; snout outline from above tion. Eleutherodactylus aphanus is an in- ovoid; nostrils not protuberant; upper eyelid habitant of upland wet forest. The three with one to several tubercles; width less than largest females (35±44 mm SVL) contained IOD; tympanum large, about 70±90% of eye yolked oviducal eggs 1.4±2.4 mm in diam- length in males, 50±70% in females; vomer- eter. ine odontophores triangular, lying posterior DISTRIBUTION: Eastern Guatemala (Depto. and mediad to large choanae; patches nar- Izabal) in the premontane zone of the Mon- rowly separated medially; subarticular tuber- tanÄas del Mico and adjacent part of the Sierra cles ovoid, obtuse; thenar tubercle low, elon- de las Minas, in a known elevational range gate; palmar tubercle somewhat larger, bi®d; of 591±786 m (®g. 9). 20 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 8. Adult Eleutherodactylus aphanus Campbell, a light colored male and a darker female from Depto. Izabal, Guatemala. Upper: Male paratype (UTA-A 24480, 22 mm SVL). Lower: Female paratype (UTA-A 33414, 44 mm SVL). [Photographs courtesy of Jonathan A. Campbell.] 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 21

Eleutherodactylus biporcatus (W. Peters) laterosacral ridge system of small tubercles Figures 3B, 4±5, 10, 11 (map), 21E, F or a few large, irregularly arranged black- Strabomantis biporcatus Peters, 1863: 405±407. tipped warts, absent in some individuals or Lectotype: Herewith designated ZMB 3330B, occasionally lengthened posteriorly to form an adult female from ``Veragua'', in error for a V-shaped ridge system (e.g., AMNH northern Venezuela. 70547); venter coarsely areolate; snout trun- Hylodes maussi Boettger, 1893: 39±40. Lecto- cate from above; nostrils not protuberant; type: SMF 3821 (not seen),10 from Puerto Ca- snout short, slightly obtuse in pro®le; eye bello, [Carabobo State], Venezuela, [near sea moderate, about equal to loreal distance (E- level; probably a general locality or shipping N); upper eyelid with many moderate-sized point as suggested by La Marca, 1992: 39]. NEW supraocular tubercles or only a few large SYNONYMY. Eleutherodactylus cornutus: Gines, 1959: 109 ones; superciliary tubercles moderate-sized (®g.), 112. and widely spaced; eyelid width almost equal Eleutherodactylus cornutus maussi: Rivero, 1961: to IOD; tympanum large, equal to or larger 54. than length of eye in males, three-fourths to Eleutherodactylus maussi: Lynch, 1975: 31. equal in females; vomerine odontophores slightly arched, lying posterior and mediad DIAGNOSIS: This is the most distinctive of to choanae; patches narrowly separated me- the species discussed here in having a coarse- dially; subarticular tubercles rounded and ly areolate venter, a distinct inner tarsal fold, projecting on all ®ngers; thenar tubercle and vocal slits in adult males. None of these raised, oblong; palmar tubercle larger, features is present in the other taxa covered strongly bi®d, outer segment oblong, much in this review. smaller than inner; de®nite large accessory MORPHOLOGY: Adult males about 30±38 palmar tubercles; ulnar tubercles evident in mm; adult females 49±74 mm (mean SVL ϭ juveniles, weak or absent in adults; heel with 36.1 mm, 6 (, 59.5 mm, 9 &); tibia 48±57% scattered small tubercles; sole of foot with of SVL; head width 39±54% of SVL; cranial large plantar tubercles; weak keels on toes; crests rather low, no raised bosses at poste- inner metatarsal tubercle prominent, an elon- rior end of crests and no distinct otic crests; gate ovoid; outer metatarsal tubercle round, dorsum smooth to tuberculate (®g. 10) below; a well-developed inner tarsal fold. tween major ridges and/or tubercle series; ju- COLORATION: Adult and juvenile Eleuth- veniles much more tuberculate than adults, erodactylus biporcatus are similar in dorsal often with short ridges of coalesced tubercles coloration except that juveniles have more especially laterally; ¯anks tuberculate to dark markings. Upper surface brown; plicae granular; paired suprascapular plicae usually outlined by black, or dark pigment reduced distinct, forming a chalice- or hourglass- to scattered spots; large tubercles in latero- shaped system, often connected or nearly sacral area black, outlined by cream; dark connected by a medially directed arm from bordered, light interorbital bar in some ju- each plica; a completely separate pair of usu- veniles but not in large adults; upper lip uni- ally distinct )(-shaped or posteriorly diverg- form or usually barred with darker pigment ing paravertebral plicae at midbody; a weak on the brown ground color; lower margin of 10 Mertens (1967: 39) listed this specimen as holotype, orbit usually marked by two or more black, which, in this case, probably quali®es as designation of light-edged spots; light supratympanic fold, lectotype ``by inference'' (Internatl. Commission., 1999: usually bordered by black below; no black art. 74.6). As assumed by Mertens, Boettger (1893: 39± spot on tympanum; no complex black and 40) probably based his description on the single speci- white marking on the groin, which is similar men presumed to be adult. But Boettger mentioned having two young specimens also. Such ambiguity as to in color to venter and ¯ank; upper limb sur- whether or not ``a nominal species-group taxon was es- faces barred with darker pigment on brown tablished on a single specimen'' is commonplace in the ground color; bars much more evident in ju- older literature, although the additional specimens now- veniles than in adults; posterior thigh surface adays are usually considered as syntypes (``cotypes'' formerly). One of Boettger's juvenile specimens may have dark brown with many tiny yellowish spots; been exchanged to the Museum of Comparative Zoology dark seat patch around vent; no contrasting as a ``cotype'' (specimen cited by Lynch, 1975: 42). series of dark and light bars on inner surface 22 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 9. Central America, showing distributions of Eleutherodactylus aphanus Campbell and Eleuth- erodactylus megacephalus (Cope). of shank; edge of throat brown with numer- the ®eld and in cages, remaining in the same ous tiny cream spots; a series of light spots spot for several days at a time and sometimes along edge of lower jaw in juveniles; venter returning to the same site after once leaving light tan to whitish with numerous small it. The hiding and resting place of one spec- whitish spots, one on each ventral areolation; imen (presumed & 50 mm SVL) ``was under undersides of limbs similar to throat in col- leaf litter and in a small cavity in the clay oration. just large enough to accommodate . . . bot- NATURAL HISTORY: Heatwole (1962) gath- tom of the cavity was covered with frag- ered data on Eleutherodactylus biporcatus mented leaves''. This frog was observed for (then known as E. cornutus maussi) over a nearly three weeks before it abandoned the period of several months in the cloud forest site. Heatwole (1962: 5) cautioned that his or montane rain forest of Rancho Grande. He observations might have re¯ected a tendency found that specimens were sedentary both in of the frogs ``to return to the same place after 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 23

Fig. 10. Eleutherodactylus biporcatus (W. Peters). An unsexed subadult (KU 166369, 37 mm SVL) from Rancho Grande, Venezuela. [From a transparency by W. E. Duellman.] activity periods rather than [strictly] seden- disturbances that are commonly encountered in other tary habits as frogs forced to move often re- anurans such as in¯ating with air or voiding urine when captured. An additional behavioral pattern . . . turned to their original location before the deserves comment. When the litter was removed from next observation period.'' above the female attending eggs, she in¯ated herself, Based on examination of ovaries, Heatwo- raised up on all four legs, lowered her head and le (1962: 2) thought that the species bred in arched her back in a manner reminiscent of the de- fense±®ght reaction of Bufo (Noble 1931). Whenever both rainy and nonrainy seasons. He found a she was touched she emitted the startling shriek female of 60 mm SVL attending a clutch of which had led to her discovery. Neither females not 45 eggs in a cavity under leaf litter: ``The attending eggs nor males displayed this striking be- cavity was just large enough to accommodate havior pattern. the eggs and female, and appeared to have Males are not known to vocalize, although been made by her . . . On top of her was a the presence of vocal slits is suggestive that layer of litter 4 cm thick.'' Two of the eggs, they may (see also footnote 3). Courtship or measuring 4.4 and 8.4 mm, were unpig- amplexus has not been reported for the spe- mented and contained advanced embryos cies. Heatwole (1962: 4) found specimens in with large yolk reserves; partial data were the stomachs of two colubrid snakes (Den- obtained on development and hatching. New- drophidion and Liophis). ly hatched froglets averaged 10.7 mm SVL DISTRIBUTION: Eleutherodactylus biporca- (10±11.4 mm, N ϭ 14). tus (s.s.) is endemic to northern Venezuela, Heatwole discovered the aforesaid nest af- where it seems to be known only from a few ter tapping a pile of litter and eliciting a premontane localities in the Cordillera de la ``screeching sound'' from beneath (1962: 2). Costa and in the SerranõÂa del Interior (®g. 11; He went on (1962: 4) with comments on de- earlier maps in Lynch [1975: 18] and La fense mechanisms and maternal care: Marca [1992: 112], as E. maussi). [Eleutherodactylus biporcatus] has the responses to REMARKS: Eleutherodactylus biporcatus 24 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 11. Locality records for Eleutherodactylus biporcatus (W. Peters) in the Cordillera de la Costa and SerranõÂa del Interior of northern Venezuela, and Eleutherodactylus opimus, new species, in eastern Panama and western Colombia.

(s.s.) is known mainly from the high wet for- employ of the U.S. Fish and Wildlife Ser- est at Rancho Grande (see Beebe and Crane, vice.) 1947) in the Cordillera de la Costa, but one Heatwole (1962: 1±2) reported a maxi- available specimen extends the range over mum size of 30.2 mm SVL for a male in a 100 km southeastward to the SerranõÂa del In- preserved sample of 20 frogs (presumably 10 terior. This frog (KU 185672), a well-pre- of each sex), and a maximum of 74 mm SVL served adult female 63 mm SVL, was col- in a ®eld-encountered female (presumably lected by Scott J. Maness on the night of released). He postulated that sexual maturity February 10, 1974 from ``rd → 1.5 K, Guz- in females was attained at about 60 mm SVL manera trial [trail?], P[arque] N[acional] ``as no individuals smaller than this con- Guatopo, Miranda'' (according to a list pre- tained mature ova''. Our small sample (table pared by Maness for specimens sent to KU, 2) indicates a larger size for males (to 38 courtesy of J. E. Simmons). It was collected mm) and a smaller size for sexually mature at the same time as a specimen of Bufo ``ty- females. phonius'' (KU 185715); the elevation was not recorded and no other data are available. Eleutherodactylus gulosus (Cope) (Maness at that time was a Peace Corp Vol- Figures 1, 3A, 12±13, 14 (map), 21A unteer stationed at Rancho Grande; later, he Lithodytes gulosus Cope, 1875: 112±113. Holo- and a companion were killed ®ghting a ®re type: USNM 32590, an adult female from in Florida scrubland in 1981, while in the ``Spur of the Pico Blanco'', Costa Rica, ``at 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 25

Fig. 12. Eleutherodactylus gulosus (Cope). An adult female (AMNH 124370, 84 mm SVL) from continental divide above upper Quebrada de Arena, 1160±1220 m, ChiriquõÂ, Panama. [From a transparency by C. W. Myers.]

6000 feet elevation'', error for Cerro Utyum, (®g. 13), which have pustulate skin (®gs. 1, CantoÂn de Talamanca, Provincia de LimoÂn, 13). Eleutherodactylus gulosus further differs Costa Rica, 1829 m (see Savage, 1970, 1974). from its geographically closest relatives (E. Hylodes gulosus: Boulenger, 1882: 211. megacephalus, E. rugosus) in having the Eleutherodactylus gulosus: Noble, 1918: 330. Eleutherodactylus rugosus (part): Taylor, 1952: black ``seat patch'' marking restricted to a 730. small area immediately surrounding the vent, Eleutherodactylus biporcatus (part): Lynch, 1975: and in lacking conspicuous pale areas in a 23. Jungfer. 1988: 15 (part [photograph ϭ E. dark reticulum across the chest. megacephalus]). MORPHOLOGY: Adult males unknown (per- DIAGNOSIS: The smooth- or nearly smooth- haps about 40±60 mm SVL?), adult females skinned adults of this large species (®g. 12) 72±103 mm; tibia 49±57% of SVL; head are immediately distinguished from all other width 46±52% of SVL; frontoparietal crests members of the group by the complete ab- well developed in larger specimens (but not sence of dorsal ridging or plicae on the body, posteriorly knobbed), and a de®nite palpable although distinctly shaped suprascapular pli- otic crest, terminating well above the supra- cae are faintly indicated in most juveniles tympanic fold as an externally expressed 26 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 13. Eleutherodactylus gulosus (Cope). Dorsal view of a juvenile (AMNH 124368, 19.7 mm SVL) showing characteristically con®gured anterior pair of short, weak suprascapular plicae (arrows), which disappear during ontogeny, ϫ4.8. The lateral edges of the plicae are emphasized by dark pigment. The transverse dark line between the posterior ends of the plicae conceals (in some specimens) a weak tubercular connecting ridge. The pustulate nature of the dorsal skin in juvenile E. gulosus is even more evident in life (see ®g. 1). boss in larger specimens; dorsal surfaces width less than IOD; tympanum relatively smooth or with scattered low tubercles in small, 35±52% of eye length in 10 juvenile adults, but covered with many small pustules males (xÅ ϭ 45.1%), 34±51% in 8 juvenile in juveniles; no suprascapular ridges in females (xÅ ϭ 39.8%), 42±54% in a subadult adults, but most juveniles in the range of 18± and two adult females (xÅ ϭ 49.7%); tympa- 32 mm SVL usually with distinctly shaped num an anteriorly tipped vertical oval (ap- but weakly developed anterior suprascapular pearing nearly round in one juvenile, AMNH plicae (®g. 13); no V-shaped or parallel ridg- 95042), tympanum length 71±94% of height es or rows of enlarged tubercles in laterosa- (xÅ ϭ 81.6%, N ϭ 21); vomerine odonto- cral region of either adults or juveniles; phores triangular, lying posterior and mediad ¯anks granular, with weak lateral ridges in a to choanae, patches narrowly separated me- few specimens (AMNH 124371±124372); dially; subarticular tubercles ovoid and ¯at- venter smooth; snout outline from above tened under ®ngers, ovoid and slightly pro- truncate; snout short, slightly obtuse in projecting under toes; thenar tubercle large, ®le; nostrils not protuberant; eyes moderate, elongate; palmar tubercle low, bi®d to tri®d; length about equal to loreal distance (E-N); no de®nite accessory palmar tubercles; no ul- upper eyelid with a few large tubercles nar tubercles; heel with 1 or 2 weakly en- mixed with many small pustules in juveniles, larged tubercles; sole smooth; no de®nite lat- 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 27 eral ridge or fringe on toes; a prominent, in life irregularly and inconspicuously mot- large, oval inner metatarsal tubercle, outer tu- tled pale tan and black (tan and brown in bercle low, much smaller; no tarsal fold; tar- preservative) below posterior extensions of sus smooth. the dark limb bands; ``seat patch'' small, COLORATION: Two adult females (72, 84 comprising a discrete blackish mark imme- mm SVL) from Panama uniform dark brown diately around vent or obscured by dorsal above in life and in preservative; subadult color in largest adults; tongue light yellow in female (46 mm SVL) orange-brown dark- an adult; inside lower lip light yellow and ened by vague black mottling in life, lighter base of tongue yellowish white in juveniles; brown in preservative with some obscure buccal cavity otherwise white; series of small black spots on posterior dorsum and dark light spots along lower lip margin except in bars on limbs; juveniles lighter brownÐin- large adult females; iris bronzy brown with dividually variable in life (blackish brown, dense ®ne black venation in adults, venation brown, light orange-brown, or yellowish less evident in juveniles. Examined in good brown)Ðwith dull darker brown spotting light, the iris for one series of juveniles and mottling on back and darker brown bars (AMNH 95035±95046) was described in on limbs; anterior dorsum of juveniles some- ®eldnotes as times relatively free of dark pigment, show- bronze on upper half, turning (without sharp demar- ing as a light area between suprascapular pli- cation) slightly duller or paler bronze on lower half, cae, which are edged in dark pigment, with and marked overall with dense, poorly de®ned black a transverse dark line connecting the poste- venation; a few individuals have the vaguest hint of rior ends of the plicae (®g. 13); usually a a dark horizontal streak through the pupil. light interorbital bar bordered by brown pos- NATURAL HISTORY: Except for descriptions teriorly in juveniles; upper lip uniform to of its habitat in Panama, very little is known weakly barred with brown and tan in juve- about Eleutherodactylus gulosus. Juveniles niles; uniform or with small light lip spots in were common at an elevation of 1000 m, on adults; lower border of orbit marked with al- the forest ¯oor by day at the Fortuna Dam ternating small black and lighter spots in site (in 1976 before construction of the dam adults, much duller or absent in juveniles; had been started), at the narrow downstream supratympanic fold black in adults, colored end of the valley of the upper RõÂo ChiriquõÂ. similar to rest of head in juveniles; tympa- Specimens later were found in the upper part num colored like rest of head in juveniles, of this humid highland valley at Quebrada de uniform dark brown or with a black spot in Arena and on the continental divide above adults. the upper Quebrada de Arena. As mentioned No complex contrasting mark in the groin, under Remarks below, Myers and Duellman which tends to be weakly and vaguely mot- (1982: 12±14) described the geography of tled brown on white (marbled blackish gray the highland valley of the upper RõÂo Chiri- on white in a subadult female); in life, un- quõÂ. Myers et al. (1984: 13) described the derside of head nearly uniform light brown habitat on the continental divide (Ͼ 1100 m) or gray-brown in adult females, gray with as a cool cloud forest that small white spots and an irregular midgular light stripe in a subadult female; underside receives almost daily fog that is frequently accom- of upper arm, hind limbs, and venter mottled panied by rain and mist. This cloud forest is a facies of high-canopied lower montane rain forest. Tree or spotted with brown on white in adults; growth is very dense, and there is a frequently dense throat and chest of juveniles greenish gray herbaceous and fern layer at ground level. There is a with white speckling on throat and white modest ``moss layer'' on the trunks of trees, which spots on chest; undersurfaces of limbs and support a profusion of epiphytic growth, including posterior venter bright greenish yellow with- bromeliads. out markings in juveniles; posterior surface Except for one individual picked up at of thighs of adults nearly uniform silvery night, all specimens were on the surface of gray in life below obscure dorsal dark bars the leaf litter by day. The largest Panamanian (dull light spots or short vertical dashes noted female (®g. 12) was sitting placidly on the in preservative); posterior thigh of juveniles ground on a narrow ridge, next to a log 28 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 14. Costa Rica and western Panama, showing localities for Eleutherodactylus gulosus (Cope) and Eleutherodactylus rugosus (W. Peters). where the collector was having lunch. A few of their moisture orographically; this moist individuals were kept for a while in captivity air funnels through the narrowing valley, at the American Museum, where they fed profoundly affecting the climate and biota on readily on crickets and appeared to be sit- this part of the Paci®c versant. Eleuthero- and-wait predators. One adult female (72 mm dactylus gulosus is probably a primarily At- SVL) also ate baby mice. lantic-versant species as it is not known from DISTRIBUTION: Humid premontane slopes the relatively well-collected Paci®c slopes of of the Cordillera de Talamanca in southern Panama west of the Fortuna region. Costa Rica and extreme western Panama, The ontogenetic change in the dorsal in- with an elevational range of 1000±1220 m, tegument is striking in this species. Juveniles perhaps to 1830 m (®g. 14). are strongly pustulate (®g. 1) and have faint REMARKS: As pointed out by Myers and but usually discernible short, anteriorly situ- Duellman (1982), the area of the upper val- ated suprascapular plicae (®g. 13), whereas ley of the RõÂo ChiriquõÂ in Panama (where this the few adults available have essentially species was rediscovered), although on the smooth dorsa (®g. 12). The subadult female Paci®c slope, has a strongly Atlantic-versant (AMNH 124371, 46 mm SVL) retains fewer, herpetofauna. This results from its location less conspicuous pustules, and the suprascap- to the south of a low section of the conti- ular ridging has virtually disappeared. Adults nental divide, which dips below 1200 m have on each side a conspicuous otic knob above the upper RõÂo ChiriquõÂ Valley. As- or boss showing prominently behind the eye cending air currents from the Atlantic versant and above the tympanum (®g. 12). This spill into the valley before having lost most knobÐthe external expression of the raised 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 29 terminus of the otic crestÐis discernible in large (females) frogs (®g. 15) having well- the larger juveniles, being positioned imme- developed suprascapular plicae and a de®nite diately under the suprascapular plica. laterosacral system of linearly arranged tu- There also is ontogenetic change in dorsal bercles or tuberculate ridges, and with the coloration. Adults (®g. 12) become an over- dark seat patch mark continuing onto the all darker color, and the rear of the thigh, thigh. Unlike the more rugose Eleutherodac- which is inde®nitely mottled or spotted with tylus rugosus, the posterior thigh surface in pale tan in juveniles, turns silvery gray in E. megacephalus is not contrastingly barred adults, the largest of which had also acquired with black and red (white in small juveniles) silvery gray coloring along the lower sides but may be uniform or dark with some small and anterior face of the thigh. Dark barring yellow spots or mottling. on the hind limbs is evident in juveniles (®g. MORPHOLOGY: Adult males 30±43 mm 13), obscure or obsolete in large specimens. SVL, adult females 50±70 mm (mean SVL Ontogenetic change in ventral coloring is ϭ 33.6 mm, 10 (, 61.6 mm, 10 &); tibia striking. Juveniles have the underside of the 46±60% of SVL; head width 44±55% of head and chest greenish gray, with white SVL; large knobs or bosses at posterior end speckling on throat and with larger white of cranial crests; distinct bony otic crests in spots on chest, with the greenish hue and large adults, with a terminal boss or knob chest spots disappearing in adults. The belly externally evident above supratympanic fold is conspicuously bright greenish yellow in in large females; dorsum smooth to tuber- juveniles, but white with brown mottling in culate between major ridges and tubercle se- adults. ries; scattered pale-tipped, pointed pustules We have accepted Cope's (1875) relatively of juveniles change ontogenetically to fewer fresh measurement of 103 mm for the gulo- and lower pustules in adults; ¯anks tuber- sus holotype; during a brief examination, culate to granular; paired suprascapular pli- Myers jotted down ``95 mm SVL'' a century cae forming a de®nite chalice- or hourglass- later; Lynch's (1975: 23) maximum size of shaped system; a discrete paired series of ``110 mm SVL'' for female E. biporcatus s.l. raised lateral and sacral tubercles, usually was based on the same specimen (personal forming a V-shaped ridge system (®g. 3D) or commun.). Even with the lesser ®gure, (in Panama) a pair of nonconverging parallel Eleutherodactylus gulosus attains the largest ridges (®g. 3E); venter smooth; snout outline size in the biporcatus group. The largest females of E. biporcatus s.s. and E. megace- from above truncate; nostrils not protuberant; phalus approach or slightly exceed the lower snout short, slightly obtuse in pro®le; eye end of the size range of E. gulosus (table 2). moderate about equal to loreal distance (E- N); upper eyelid with moderate numbers of widely spaced supraocular and superciliary Eleutherodactylus megacephalus (Cope) tubercles, width less than IOD; tympanum Figures 3D±E, 6, 9 (map), 15±16, 21B large, 80±90% of eye length in males, 40± Lithodytes megacephalus Cope, 1875: 110±111, 50% in females; vomerine odontophores tri- pl. 23, ®gs. 5, 11, 11a. Holotype: USNM 32579, angular, lying posterior and mediad to cho- an adult female from ``spur of the Pico Blan- anae; patches narrowly separated medially; co'', Costa Rica, ``at 6,000 feet elevation'' ϭ subarticular tubercles ovoid under ®ngers I± Cerro Utyum, CantoÂn de Talamanca, Provincia de LimoÂn, Costa Rica, 1829 m (see Savage, II, obtuse, ¯attened under ®ngers III±IV; the- 1970, 1974). nar tubercle low, oblong; palmar tubercle, Hylodes megacephalus: Brocchi, 1882: 57; Bou- larger, rounded, bi®d; no de®nite accessory lenger, 1882: 211. palmar tubercles; ulnar tubercles well devel- Hylodes biporcatus (part): Boulenger, 1882: 220. oped in juveniles, weak or absent in adults; Eleutherodactylus rugosus: Noble, 1918: 329. heel with 1 or 2 enlarged tubercles; sole Eleutherodactylus biporcatus: Dunn, 1931: 410; smooth; no de®nite lateral keel or fringe on Lynch, 1975: 23 (part). Jungfer, 1988: 16 (part toes; a prominent oval, sometimes com- [confused with E. gulosus specimens]). pressed inner metatarsal tubercle, outer low, DIAGNOSIS: Moderate-sized (males) to much smaller; no tarsal fold; several promi- 30 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 15. Eleutherodactylus megacephalus (Cope). An adult female from La Selva Biological Station, Heredia Prov., Costa Rica. [From a transparency by Robert Wayne Van Devender.] nent outer tarsal tubercles in juveniles, weak and light, or uniform; lower margin of orbit in adults. usually bordered by alternating series of COLORATION: Adult and juvenile Eleuth- small black and cream spots; supratympanic erodactylus megacephalus are similar in dor- fold usually black; usually a black spot on sal coloration, except that juveniles usually tympanum; no complex black and white have a lighter ground color so that the dark marking in groin, which is uniform or similar markings are more sharply de®ned. Upper to adjacent posterior venter in pattern and surfaces usually gray-tan to olive-brown, color; upper limb surfaces uniform or usually sometimes with a strong salmon cast,11 but marked with obscure dark bars; posterior large females may be dark brown; plicae out- thigh surface dark brown to black, uniform lined by black; distinct, small supra-axillary or variously spotted or mottled with light but and sacral black spots usually present in as- never with vivid alternating black bars and sociation with enlarged tubercles; vague light red or white interspaces; black seat patch interorbital bar or spot usually present, de- usually continuous with black background of marcated anteriorly and posteriorly by areas lower posterior thigh which extends some slightly darker than dorsal ground color; up- distance onto undersurface of thigh; no con- per lip with a dull light stripe, barred dark trasting pattern of black and white bars or white spots on inner surface of shank; throat 11 Jungfer (1988: 16) published a photograph of an apparently young male specimen (from CocleÂ Province, variable, light with small brown punctations Panama) that reproduced as a vivid light orange. to mostly brown with small light spots or 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 31 covered with large light spots or a reticulum with heavy black suffusion; one was greenish as on chest or venter; obscure, narrow light bronze with dense black venation. midgular stripe rarely present; series of light NATURAL HISTORY: Nothing has been pub- spots along edge of lower jaw in juveniles, lished on the breeding habits of Eleuthero- lost in large adults; throat and chest covered dactylus megacephalus and males are not anteriorly by a brown reticulum containing known to call. The usual habitat is well- large round light spots in juveniles; reticulum drained ground in tropical wet forest, includ- and round spots covering most of venter in ing secondary growth and cacao plantations, some examples but becoming irregular with but one was found in swamp forest near the increase in size; in some large adults, retic- RõÂo ConcepcioÂn on the north coast of Pana- ulum reduced to scattered dark spots; a sim- ma. ilar pattern may be present on undersurface Juveniles are found active on the leaf litter of thighs and shanks; Costa Rican juveniles both by day and night, although adults are have concealed yellow-orange to red-orange seldom seen. Dunn (1931: 410, as E. bipor- coloring12 underneath the legs and in the light catus) noted the scarcity of adults on Barro spots on the posterior venter, and on the Colorado Island in central Panama and haz- palms of hands and underside of ®ngers and arded a guess that they might ``live in bur- toes (sole of foot dark brown); anteriormost rows''. Based on a 13-month sampling pro- light ventral areas cream; with age, orange gram at La Selva in Costa Rica, Savage and hues replaced by cream so adults usually his students (unpublished) concluded that have at most a little orange in groin and on adults are nocturnal and hide in burrows dur- digital tips; iris noted as black ¯ecked with ing the day, sitting at the burrow entrance at gold in Costa Rica. night and ambushing passing prey. Presum- The yellow-orange to red-orange ventral ably they use burrows made by other animals coloring in Costa Rican juveniles has not inasmuch as frogs of this group are not known as excavators. been noted by Myers in Panama.13 His color Noble (1918: 330±331, as E. rugosus) ex- notes on juvenile and subadult specimens amined a few stomachs of Nicaraguan from Panama (several localities in Bocas del Eleutherodactylus megacephalus and found Toro and one on north coast of Veraguas) mostly beetles and a large ant (Neoponera). recorded one tiny juvenile in which all ven- Lieberman (1986: tables 7±10, as E. bipor- tral surfaces were bright greenish yellow. A catus) reported a wide variety of arthropods greenish color under the hind legs was re- and a ``vertebrate'' (presumably a frog or tained in some larger juveniles, but otherwise small squamate reptile) from 21 stomachs of the throats were gray with white dots, and frogs from La Selva, Costa Rica; the most the venters were gray to black with white important prey items by proportion of stom- spots, or marbled in black and white. Rear achs were beetles (30%), ants (23%), orthop- thigh surfaces in Panama were variable: terans (14%), isopods (9%), unidenti®ed lar- black with tan or yellowish tan spots, gray vae (9%), and spiders (5%). with white speckling, or silvery gray and DISTRIBUTION: Eleutherodactylus megace- black. The iris in Panama was described as phalus inhabits lowlands and premontane brown or (usually) bronze, sometimes lighter slopes on the Atlantic versant from south- below, with ®ne or dense black venation or western Honduras to central Panama (®g. 9), where it also occurs in uplands on the Paci®c 12 The bright juvenile colors in Costa Rica tend toward versant. It is found from near sea level to yellow in the smallest frogs of about 10±15 mm SVL, 1200 m elevation, perhaps rarely higher (as but are bright salmon to red-orange and most extensive in juveniles of about 25±30 mm SVL. already noted under Application of the Name 13 However, in their very useful survey of the am- Lithodytes megacephalus, the stated type lo- phibians of the Panama Canal region, IbaÂnÄez et al. cality at 6000 ft [1829 m] is unaccountably (1999: 41, 139) state that the venter, groin, and con- high). cealed surfaces of the hind legs are pale orange or yellow in juvenile ``biporcatus.'' The species referred to is REMARKS: As stated earlier (under Appli- uncertain, inasmuch as both E. megacephalus and E. op- cation of the Name Lithodytes megacephalus imus occur in the area of their study. Cope, 1875), we have had to conclude that 32 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 16. Eleutherodactylus megacephalus (Cope), juveniles (ϫ1.5) showing variation in the laterosacral ridge systemÐan instance of unusual intrapopulational variation, or of sympatry with an unnamed sibling species? Upper specimens left to right, AMNH 107276, 107279; lower specimens AMNH 107277, 107278, 107280; all from Quebrada El Guabo, 150±250 m, Bocas del Toro, Panama. The two uppermost frogs have posteriorly converging laterosacral plicae that meet in a V-shape, as is characteristic of Costa Rican populations of E. megacephalus. The three specimens in the lower row have a pair 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 33

Cope's holotype of megacephalus probably color pattern. None of the juveniles had red was collected at a lower elevation than stated or orange ventral coloring. The smallest fro- and that, in any case, it almost certainly is glet (12.5 mm SVL) in ®gure 16 had all ven- conspeci®c with some lowland populations tral surfaces uniformly bright greenish yel- of frogs from the Atlantic versant of lower low except for scattered white spots on the Central America. Following Dunn's Barro chest; the other juveniles had the throat gray Colorado Island paper (1931), authors have with irregular white dots, the venter pat- erroneously assigned the name Eleuthero- terned with larger white spots, and the rear dactylus biporcatus to lowland broad-headed of belly and undersides of the hind legs frogs from Honduras to northwestern Colom- greenish ¯esh. The ®ve juveniles shown in bia. But, in this sense, the misapplied name ®gure 16 are readily separated by laterosacral ``biporcatus'' proves to be a composite of at ridges and vividness of dorsal pattern into least three species, one of which (rugosus)is two kinds of frogsÐbut we are unable to resurrected and another (opimus) is described conclude whether this is an instance of sym- as new on following pages. patric ``sibling'' species or one of intrapo- The remaining part of the old ``biporca- pulational polymorphism. tus'' is here assigned to Cope's Eleuthero- Therefore, we are not prepared at this time dactylus megacephalus, with a distribution to give a de®nitive opinion on the status of on the Atlantic versant from northwestern frogs from the western half of the Isthmus of Honduras to central Panama, where there are Panama, being hindered by the paucity of some occurrences on the Paci®c side of the adult specimens and especially by the loss or low continental divide. However, even with temporary misplacement of several dozen this geographic restriction, there remains a critical specimens in the University of Kan- concern that we may still be dealing with a sas collections (see comment on page 43, composite species. This concern rises (1) Specimens Examined). Consequently, our as- from the conspicuous red or orange ventral signment of Panamanian specimens to E. me- coloring that is present in Costa Rican ju- gacephalus is tentative. veniles but lacking in Panamanian specimens Most Panamanian specimens assigned to (see last paragraph under Coloration above), Eleutherodactylus megacephalus are from and (2) from variation in the laterosacral tu- the lowlands and intermediate elevations of bercle or ridge system in populations in low- Bocas del Toro Province. Other material ex- land Bocas del Toro in extreme northwestern amined by us is scattered and sparse. Myers Panama. collected a single juvenile (KU 113676) from Concerning the laterosacral ridge system, the mouth of the RõÂo ConcepcioÂn, on the the most relevant material available at this north coast of Veraguas Province, Panama, time (see ®g. 16), is a small series from the about halfway between Bocas del Toro and foothills of the Cordillera de Talamanca in the Panama Canal at Barro Colorado Island, Atlantic-western Panama (AMNH 107276± which is our easternmost record for mega- 107282, Quebrada El Guabo, 150±250 m, cephalus. Between those localities, we know RõÂo Changuinola drainage). Included are an E. megacephalus from the low continental adult female (54 mm SVL) and a subadult divide north of El CopeÂ (CocleÂ Prov.), at El female (30 mm), both with V-shaped latero- Valle de AntoÂn (CocleÂ), and at Cerro Cam- sacral ridges. Of ®ve juveniles (®g. 16) in the pana (PanamaÂ). Available specimens from series, two have the V-shaped ridges and these sites are all small juveniles or sub- three have nonconverging, parallel laterosa- adults. They agree with similar-sized mega- cral ridges; they differ also in vividness of cephalus from Costa Rica in having the dor-

← of nonconverging, parallel plicae, which is the typical situation in Panamanian frogs currently assigned to this species. (The pale vertebral line in two frogs, although infrequent in the biporcatus group, is a common morph in many species of Eleutherodactylus and various other anurans; it probably lacks taxonomic signi®cance.) 34 AMERICAN MUSEUM NOVITATES NO. 3357 sum covered with widely-spaced whitish- a Latin adjective meaning rich or fruitful and tipped tubercles and evident suprascapular also, as in the present context, well-fed or fat. and laterosacral tubercle or ridge systems. (It DIAGNOSIS: Moderate-sized (males) to will be recalled that during ontogeny in E. large (females) frogs (®g. 17) resembling E. megacephalus, the tubercles not involved in megacephalus in having well-developed su- the ridging system become reduced so that prascapular plicae and sometimes a white- adults have a shagreened to nearly smooth spotted venter, but differing in lacking a def- dorsum, except for the suprascapular and la- inite laterosacral system of paired linear or terosacral ridges.) V-shaped tubercles or ridges (®g. 18). Frogs of this species group east of the Pan- Eleutherodactylus opimus is readily distin- ama Canal differ most conspicuously from guished from Costa Rican E. rugosus in be- Eleutherodactylus megacephalus in lacking ing much less tuberculate and in color pattern the laterosacral series of linearly arranged tu- (rugosus with posterior thigh brightly barred bercles or ridges, as described in the next black and red, and often with dorsolateral species account. stripes). Super®cially, E. opimus resembles the smaller Guatemalan E. aphanus, which Eleutherodactylus opimus, new species differs in having an immaculate venter lack- Figures 11 (map), 17±18, 21C ing any indication of dark markings or white spots; opimus also lacks the weak ®nger and Eleutherodactylus biporcatus (part): Dunn, 1931: toe keels of aphanus (Campbell, 1994: 298). 410; Lynch, 1975: 23; 1997: 354, ®g. 1 top MEASUREMENTS OF HOLOTYPE (in mm): (skull). Ruiz-Carranza et al., 1996: 395. Eleutherodactylus biporcatus: Lynch, 1998: 145. The holotype is a subadult female having tiny ova (to about 0.3 mm) and thin (0.2) HOLOTYPE: AMNH 87020 (®eld no. CWM nonconvoluted oviducts. SVL 41.9, tibia 10307), a subadult female from Quebrada length 21.5, foot length from proximal edge DocordoÂ, about 10 km above junction with inner metatarsal tubercle to tip of toe IV RõÂo San Juan, about 100 m elev., Depto. 18.5, head width 20.5, head length on the ChocoÂ, Colombia (approx. 4Њ34ЈN, 77Њ03Ј diagonal from angle of jaw to tip of snout W), collected February 5±6, 1971, by C. W. 17.2, upper eyelid width 4.9, interorbital dis- Myers and John Daly. tance 4.7, eye to posterior edge of nostril 5.0, PARATYPES: Colombia: ChocoÂ: AMNH eye length 5.6, tympanum length 2.9. 109473±109487, AMNH 109471±109472. MORPHOLOGY: Size of adult males un- Panama: ColoÂn: AMNH 89465, KU known (perhaps about 30±40 mm SVL?), 108430±108431, 113677±113678, 172226± one adult female 69 mm SVL; tibia 49±56% 172228. DarieÂn: KU 76221, 76226. Pana- of SVL (3 subadult and adult &); head width maÂ: AMNH 53704, 89464, KU 80326, 43±50% of SVL; cranial crests present (ab- 108432±108433, 108434±108435. San Blas: sent in smaller juveniles), posteriorly raised KU 113679. The species description is based or bossed in adult female; bony otic crest ev- primarily on the holotype and the specimens ident in larger specimens, terminating in an designated as paratypes. See the appendix externally expressed boss or knob; dorsum (Specimens Examined) for complete locality weakly to pronouncedly pustulate between data for paratypes. major ridges and in sacral region, with scat- REFERRED SPECIMENS: See also Specimens tered small tubercle; ¯anks tuberculate to Examined for additional material assigned to granular; paired suprascapular ridges form- E. opimus (specimens not at hand when the ing a de®nite chalice- or hourglass-shaped ®nal description was written). Specimens in- system; no discrete enlarged lateral and sa- dicated by ``*?'' were lost or misplaced be- cral tubercles or ridges forming either a par- fore we could con®rm identi®cation and allel paired or V-shaped laterosacral system; therefore can only tentatively be assigned to most specimens with 2±4 widely spaced pos- E. opimus, although that assignment for KU terior sacral tubercles forming a transverse 108934 is questionable (see footnote 15). series; smallest juveniles sometimes with ETYMOLOGY: The species name opimus is several small sacral tubercles aligned on each 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 35

Fig. 17. Eleutherodactylus opimus, new species. An adult female from Camp Summit, 300±400 m elev., San Blas-DarieÂn border, eastern Panama (KU 113679, 69 mm SVL). [Photograph by C. W. Myers.] side in a discontinuous longitudinal series in sory palmar tubercles except sometimes for anterior sacral region, these being ontoge- one between the large palmar tubercle and netically lost or reduced to a few widely base of ®nger IV; ulnar tubercles low and spaced tubercles; venter smooth; snout short, inconspicuous; heel with scattered small tu- its outline truncate from above, slightly ob- bercles, none markedly enlarged; sole tuse in pro®le; nostrils not protuberant; eye smooth; no de®nite lateral keel or fringe on moderate, roughly equal to distance from its toes; a prominent elongate, ovoidal inner anterior edge to nostril; upper eyelid includ- metatarsal tubercle, outer tubercle much low- ing supraciliary margin pustulate and tuber- er and smaller; no tarsal fold; outer tarsal tu- culate, with tubercles subequal or with sev- bercles weak or absent. eral supraocular tubercles larger than others; COLORATION: Small specimens of Eleuth- eyelid width more or less equal to interorbital distance; tympanum large, roughly half of erodactylus opimus are often lighter than eye length in females, larger in males; vo- larger ones, with more evident dark mark- merine odontophores triangular, lying poste- ings, and there may be a tendency for general rior and mediad to choanae, the patches nar- darkening during ontogenyÐthe largest rowly separated medially; subarticular tuber- specimen (®g. 17), an adult female, was al- cles ovoid and slightly protuberant under all most uniformly dark brown with a silvery ®ngers or barely ¯attened under III±IV; the- suffusion over the ¯anks and thighs. How- nar tubercle low, oblong; palmar tubercle ever, there is considerable variation in col- nearly the same size, bi®d; no de®nite acces- oration, and even equivalent-sized specimens 36 AMERICAN MUSEUM NOVITATES NO. 3357

thigh blackish (in life black, dark brown, or black with areas of gray or silvery gray), con¯uent with the dark seat patch; throat varying from light (white in life) with suffusion or punctations of light brown, to me- dium or dark brown with small white spots, usually with an ill-de®ned white midgular stripe; a series of light spots along edge of lower jaw at least in small specimens. Small to large white spots in brown reticulum across chest, this pattern tending to extend ventrolaterally, leaving belly nearly immaculate white. Iris in life described as bronzy brown in Colombian holotype; varying in Panamanian specimens from bronze dark- ened by brown or black suffusion, to black with bronze ¯ecking. NATURAL HISTORY: Nothing is known of breeding habits. The usual habitat is hilly or well-drained areas in tropical wet forest. My- Fig. 18. Eleutherodactylus opimus, new spe- ers found specimens (including the holotype) cies. A juvenile showing presence of well-de®ned mostly on leaf litter by day, but three speci- suprascapular plicae and absence of laterosacral mens from his ``Camp Summit'' locality, in- plicae, ϫ2.4. (KU 113677, 25 mm SVL, from ridge about 4 km SE Puerto PiloÂn, 190±240 m, cluding a large female (®g. 17), were found ColoÂn Prov., Panama.) on the forest ¯oor by night. Toft (1981: 141) found ants, orthropterans, and miscellaneous arthropod remains in three stomachs of Eleutherodactylus opimus (re- may differ greatly in the intensity of the ported as E. biporcatus). Toft concluded in ground color (®g. 19). summary that most (8 of 9 spp.) of the leaf- Dorsal colorations in life include light to litter Eleutherodactylus in her sample ``are very dark shades of brown, including gray- cryptic, sit-and-wait foragers which eat few ish, orangish, and reddish brown; plicae usu- large prey per day'' (Toft, 1981: 139); E. op- ally outlined by black; light-colored supra- imus was included in this category by impli- axillary and sacral tubercles usually set in cation. black spots; interorbital light bar present or DISTRIBUTION: Lowlands and premontane absent, when present usually vague and de- areas from central Panama (east of Panama marcated anteriorly and posteriorly by areas Canal) to northwestern Colombia on the Pa- slightly darker than dorsal ground color; ci®c versant (®g. 11). margin of upper lip pale, broken by two usu- REMARKS: Eleutherodactylus opimus dif- ally conspicuous dark bars radiating from fers from other members of the biporcatus eye to lip, and sometimes with vague dark group in having a distinctive karyotype, as blotching anteriorly on lip; lower margin of based on preparations made by Dr. Shyh- orbit usually bordered by alternating small Hwang Chen from three Panamanian speci- cream and black spots; the latter often su- mens collected at Nusagandi (®g. 21C, and perimposed on the two radiating dark bars; Chen, 2001). usually a black spot on upper side of tym- As in the case of Eleutherodactylus me- panum, con¯uent with black edging on su- gacephalus, available specimens of E. opi- pratympanic fold; groin pattern variable, mus consist mainly of juvenile and subadult from uniformly pale to mottled black and frogs. We have seen no adult males and only white; upper limb surfaces variable, from vir- one adult female (shown in ®g. 17), which tually uniform to black barred; anterior thigh was collected by Myers at ``Camp Summit'' surface barred or not; lower side of posterior on the low continental divide between San 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 37

Fig. 19. Light and dark variants of Eleutherodactylus opimus, new species, in life. Both are subadult females of equivalent size (42±43 mm SVL) and with uniformly small ovarian ova. Upper: AMNH 87020 (holotype) from Quebrada DocordoÂ in western Colombia. Lower: AMNH 89464 from El Llano- CartõÂ Road in central Panama. [Photographs by C. W. Myers.] 38 AMERICAN MUSEUM NOVITATES NO. 3357

Fig. 20. Eleutherodactylus rugosus (W. Peters). An adult male from vicinity of RincoÂn de Osa, Puntarenas Prov., Costa Rica. [From a transparency by Roy W. McDiarmid.]

Blas and DarieÂn in eastern Panama (locality Eleutherodactylus ¯orulentus: Taylor, 1952: 765. discussed in Myers, 1969: 26). This individ- Eleutherodactylus biporcatus (part): Lynch, 1975. ual (identi®ed in the ®eld as ``E. biporca- DIAGNOSIS: An extremely rugose, moder- tus?'') would have been designated as ho- ate-sized frog (®g. 20) trenchantly distin- lotype, except that the skull (not seen) was guished from Eleutherodactylus megace- removed before our reexamination of the phalus and from all other species of the bi- specimen, and four other specimens (KU porcatus group in having (except in smallest 113680±113681, 113693±113694) collected juveniles) black vertical bars alternating with at the same locality were lost or misplaced scarlet interspaces on the posterior thigh sur- before their identity could be con®rmed. A face, and contrasting light (usually red) and lateral outline view of the skull was given black markings in the groin and under sur- by Lynch (1997: ®g. 1 top) as E. biporcatus. faces of the hind limbs. MORPHOLOGY: Adult males 27±44 mm Eleutherodactylus rugosus (W. Peters) SVL, adult females 35±65 mm (mean SVL Figures 3C, 7, 14 (map), 20, 21D ϭ 36.1 mm, 10 (, 50.9 mm, 10 &); tibia Hylodes rugosus W. Peters, 1873: 610±611. Ho- 47±57% of SVL; head width 41±51% of lotype: ZMB 7812, a juvenile from ``Chiriqui'' SVL; large bosses or knobs externally ex- ϭ Provincia de ChiriquõÂ, Panama. pressed at posterior ends of frontoparietal Lithodytes pelviculus Cope, 1877: 89. Holotype: crests and at terminus of otic crest above USNM 32326, west coast of Central America. supratympanic fold in large adults; dorsum Lithodytes ¯orulentus Cope, 1893: 336±337. Ho- strongly tuberculate, with numerous large lotype: No. 327 G. K. Cherrie collection (now lost) from Boruca, CantoÂn de Buenos Aires, warts and ridges, and with many smooth to Provincia de Puntarenas, Costa Rica. pointed tubercles that are sometimes white Hylodes ¯orulentus:GuÈnther, 1900 (1885±1902): tipped; a chalice- or hourglass-shaped sys- 234. tem of suprascapular plicae, often with a Eleutherodactylus rugosus: Taylor, 1952: 730. medially directed branch from each plica; 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 39

¯anks tuberculate; venter smooth; snout narrow irregular midgular light stripe usu- short, slightly obtuse in pro®le; nostrils not ally indicated; series of light spots along protuberant; upper eyelid with numerous edge of lower jaw; chest and anterior venter warts, several enlarged, width less than or entire venter brown with large light spots; IOD; tympanum large, 80±90% of eye a similar pattern may be present on under- length in males, 40±50% in females; vo- surfaces of thigh and shank; sole of foot merine odontophores triangular, lying pos- dark; in juveniles, palms, toes posterior ven- terior and mediad to large choanae; patches ter, undersurfaces of hind limbs bright or- narrowly separated medially; subarticular ange to red; contrastingly marked light spac- tubercles ovoid, obtuse under ®ngers I±II, es on posterior thigh may be white in small- ¯attened under ®ngers III±IV, slightly pro- est examples. jecting under toes; ulnar tubercles well de- NATURAL HISTORY: On the Osa Peninsula, veloped; thenar tubercle low, oblong; pal- Roy McDiarmid (unpubl.) observed that ju- mar tubercle larger, bi®d; no de®nite acces- veniles of Eleutherodactylus rugosus were sory palmar tubercles; heel with 1 or 2 en- diurnal and adults nocturnal. Adults were larged pointed tubercles; sole smooth; no found in shallow depressions in leaf litter. de®nite lateral ridge or fringe on toes; an The diet was primarily insectivorous, pri- inner enlarged, oval metatarsal tubercle, marily coleopterans, but lizards and other outer tubercle much smaller, round; no tarsal frogs were occasionally eaten. fold; several prominent outer tarsal tuber- DISTRIBUTION: Lowlands and premontane cles in juveniles, weak to well developed in slopes of the Paci®c versant from the RõÂo adults. Carara, Puntarenas Province, Costa Rica COLORATION: Adult and juvenile Eleuth- (9Њ47ЈN) to extreme western Panama (10± erodactylus rugosus are similar in dorsal 1220 m; ®g. 14). coloration except that juveniles are lighter REMARKS: Small juveniles of Eleuthero- and have the dark markings more vivid. Up- dactylus rugosus are covered dorsally by per surfaces various shades of brown, but large, scattered white-tipped tubercles. Dur- usually dark reddish brown with a pair of ing ontogeny, the tubercles become dark- lighter (tan to reddish brown) broad dorsal ened and tend to fuse into ridges. Adults re- stripes extending posteriorly from shoulders tain tubercles and ridges to a much greater for about three-fourths of distance to groin; plicae and enlarged supra-axillary tubercles extent than in other species of the group. bordered by black; usually a light interor- Most individuals of all ages have a pair of bital bar bordered by black posteriorly; up- relatively broad, light tan to reddish brown per lip barred black and tan to dull cream dorsolateral stripes lateral to the suprascap- or no obvious pattern; lower margin of orbit ular plicae and extending farther posteriad. often bordered by alternating series of small The dorsolateral stripes are prominent in the black and cream spots; supratympanic fold holotype of this species, but do not occur in usually black; usually a black spot on tym- the variation of any other member of the panum; groin with white and black complex biporcatus group. marking that is continuous with similar pat- Taylor (1952, 1954) anticipated our con- tern on anterior and usually underside of clusion that this form is a valid species, al- thigh; arms obscurely barred with black though he applied the name Eleutherodac- above; hind limbs brown, barred with black tylus ¯orulentus to it. He incorrectly used on upper surfaces; black bars continue onto the name Eleutherodactylus rugosus for the posterior thigh surface and are separated by populations recognized as E. megacephalus bright scarlet interspaces; black lower third in the present paper. Although the main por- of posterior thigh surface and seat patch tion of the range of this species is around continuous; inner and often ventral surface the Golfo Dulce, it also occurs a consider- of shank barred like posterior thigh or able distance northwestward along the low- marked with large white spots on a dark lands of the Paci®c coast, and into the in- brown ®eld; throat suffused by dark brown land El General and Coto Brus Valleys of pigment that may enclose tiny light spots; a Costa Rica. 40 AMERICAN MUSEUM NOVITATES NO. 3357

IS THE ELEUTHERODACTYLUS Then, Dr. Chen called our attention to a BIPORCATUS GROUP paper by Schmid et al. (1992) describing the MONOPHYLETIC? karyology of Eleutherodactylus biporcatus (reported as E. maussi). In that study, 70 fe- That frogs of this group are unusual with- males and one male were reported to have a in the genus in having broad heads and cra- 2N ϭ 36 (considered as the ancestral con- nial crests and lacking expanded digital dition), whereas 15 additional males had 2N disks was early recognized by Peters (1863), ϭ 35. N.F.s for these karyotypes are 42. who erected a new genus (Strabomantis) for Dr. Chen very graciously prepared idio- its ®rst discovered member. The taxonomic grams of the karyotypes of members of the history of the group is reviewed earlier in biporcatus group for us (®g. 21). The differ- the present paper, where it is pointed out ences in morphology are striking and leave that Venezuelan Eleutherodactylus bipor- open to question whether there is a close re- catus sensu stricto and its allies belong to lationship between the Central American the Middle American clade (subgenus Crau- species and Venezuelan E. biporcatus s.s. gaster) characterized by the E condition of Eleutherodactylus gulosus, E. megacephalus, the adductor mandibulae muscle. Most and E. opimus have karyotypes similar to South American forms of Eleutherodacty- that of E. rugosus. But the karyotype of lus, including the broad-headed members of Eleutherodactylus biporcatus is much more the E. sulcatus group have the S condition similar to those of species having the S con- of the adductor. dition of the adductor (e.g., E. ridens,2Nϭ Based on Bogart (1973, 1981) and De- 34, N.F. ϭ 36) than to any other species of Weese (1976), Savage (1987) pointed out that Eleutherodactylus having the E condition.14 members of the Middle American clade, in- The possibility then occurred to us that cluding Eleutherodactylus rugosus (``bipor- Lynch (1986) might have been mistaken in catus''), were characterized by a 2N karyo- reporting the E condition in Eleutherodac- type of 18, 20, or 22 chromosomes equal to tylus biporcatus (as E. maussi). But dissec- a nombre fundamental (N.F.) of 36, 38, or 44. tion of specimens of this species (AMNH Much greater variation was found in those 70545, 70556) con®rmed Lynch's observa- species having the S condition of the adductor tions that the E condition characterizes this mandibulae, but most South American forms taxon. We also have con®rmed that the E have high 2N (32, 34, 36) and N.F. values (32, state of the adductor is found in all other spe- 36, 38, 44, 46). However, Eleutherodactylus cies assigned to the biporcatus group in this sulcatus, a broad-headed Amazonian species paper as previously noted by Lynch (1986) (S condition), has a low 2N ϭ 22, with N.F. and Savage (1987) and including Eleuther- ϭ 38 (DeWeese, personal commun.). odactylus gulosus (AMNH 95037). DeWeese's (1976) examination of the kar- Contrary to Miyamoto and Tennant (1984) yotypes of Eleutherodactylus rugosus (re- and Lynch (1986), we regard both the E and ported as E. biporcatus) showed a 2N ϭ 20 S states of the adductor muscles as derived and N.F. ϭ 38. Shyh-Hwang Chen of Taiwan from the plesiomorphic condition where both Normal National University has recently the posterior subexternus and externus super- completed a review of chromosome morphology in the Central American clade 14 Added in press: Dr. Chen is now inclined to believe (Craugaster). His studies showed that that the frogs used by Schmid et al. (1992) for karyo- Eleutherodactylus gulosus has a karyotype typic study may have been another species somehow similar to that of E. rugosus. Eleutherodac- misidenti®ed as Eleutherodactylus maussi (E. biporcatus s.s.). Eight other named species of Eleutherodactylus oc- tylus megacephalus and E. opimus have sim- cur in the Cordillera de la Costa (Lynch and La Marca, ilar karyotypes, with 2N ϭ 20, N.F. ϭ 36. 1993: 144); all except biporcatus are known or thought Although differing in details of chromosome to have the S condition of the adductor (based on the morphology, the similarities in karyotypes subgeneric assignments in Lynch and Duellman, 1997: 219±234). Because there are no preserved voucher spec- among these four species within the Central imens (Schmid, personal commun.), we could not verify American clade seemed to con®rm the rec- the identi®cation of the species studied by Schmid et al., ognition of the biporcatus group. whose work needs corroboration. 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 41

Fig. 21. Idiograms of haploid chromosome complements for ®ve of the six species of the Eleuth- erodactylus biporcatus group. A: E. gulosus. B: E. megacephalus. C: E. opimus. D: E. rugosus. E: E. biporcatusÐfemales and occasional males (2N ϭ 36). F: E. biporcatusÐmost males (2N ϭ 35), with the last three chromosomes being unpaired in diploid cells; the male heteromorphism was produced by a centric fusion between single chromosomes from pairs 17 and 18, with retention of the two other members as single chromosomes. See footnote 14. [Courtesy of Shyh-Hwang Chen.] 42 AMERICAN MUSEUM NOVITATES NO. 3357

®cialis adductors are present (S ϩ E condi- ACKNOWLEDGMENTS tion). We do so because the presence of both a super®cal adductor mandibulae muscle lying We are immensely grateful to Dr. Rainer wholly laterally to the mandibular branch of GuÈnther for lending the type material of Hy- the mandibular nerve (V3), and a deeper muscle lying mesially to that nerve, seems to be a lodes rugosus and Strabomantis biporcatus general pattern found in caecilians (Edgeworth, from the Zoologisches Museum in Berlin 1935; Bemis et al., 1983) and most (nonpae- (ZMB)Ðthe present paper could not have domorphic) salamanders (Luther, 1914; Fran- been written without access to these critical cis, 1934; Edgeworth, 1935; Baird, 1951; Car- specimens. roll and Holmes, 1980), as well as in some Other important museum material and data groups of anurans. This pattern (S ϩ E) there- were made available by the following col- fore appears primitive for tetrapods. leagues (institutional abbreviations as listed We agree with Lynch (1986) that although in the appendix following): Federico BolanÄos the E condition has originated independently (UCR), John E. Cadle and JoseÂ Rosado in several different anuran families, it may (MCZ), Jonathan A. Campbell (UTA), Theo- be regarded as a synapomorphy for the Mid- dore Daeschler and Ned Gilmore (ANSP), dle American clade (subgenus Craugaster) William E. Duellman, Linda Trueb, and John within the Leptodactylidae. If our interpre- E. Simmons (KU), Arnold G. Kluge and tation of jaw muscle evolution is correct, the Greg Schneider (UMMZ), Alan Resetar S condition cannot be considered plesiom- (FMNH), and Roy W. McDiarmid (USNM). orphic to the E character state but has an in- Photographs of several taxa were kindly pro- dependent origin from an ancestral S ϩ E vided by Jonathan A. Campbell, William E. condition. This in turn suggests a relatively Duellman, Roy W. McDiarmid, and Robert ancient separation of two major lineages Wayne Van Devender. within Eleutherodactylus s.l. Shyh-Hwang Chen, National Normal Uni- Although the condition of the adductor versity, Taiwan, prepared the idiograms of muscle clearly aligns the Venezuelan Eleuth- the karyotypes based on material from his erodactylus biporcatus s.s. with the Middle American clade, its karyotype (unfortunately doctoral dissertation research at the Univer- uncorroborated, see footnote 14) and differ- sity of Miami. We also thank Roberto IbaÂnÄez, ent morphology (coarsely areolate venter, in- Smithsonian Tropical Research Institute, for ner tarsal fold, accessory palmar and plantar aiding Dr. Chen in the ®eld in Panama. James tubercles, and male vocal slits) cast doubt on E. DeWeese, Napa Valley College, provided its putative relationship to the Central Amer- data from his unpublished analysis of the ican species of the ``biporcatus group'' as karyotype of Eleutherodactylus sulcatus. treated in the present paper. Unsatisfying as We thank Jonathan A. Campbell and W. it is, we must leave ®nal solution of this re- Ronald Heyer for reading and commenting maining puzzle to future workers. on the manuscript.

APPENDIX: SPECIMENS EXAMINED Collection Abbreviations: The following ab- FMNH Field Museum of Natural History breviations of institutional collections are used KU Museum of Natural History, University in the text and in the list of specimens exam- of Kansas ined: MCZ Museum of Comparative Zoology, Har- AMNH American Museum of Natural History vard ANSP Academy of Natural Sciences of Phil- SMF Natur-Museum und Forschungs-Institut adelphia Senckenberg BMNH British Museum (Natural History) UCR Universidad de Costa Rica, Museo de CRE, Costa Rican Expeditions (collection ZoologõÂa now at Los Angeles County Museum UMMZ University of Michigan Museum of Zo- of Natural History) ology 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 43

USNM National Museum of Natural History, UCR 10185. Pitilla Biol. Station, 600±825 m, Smithsonian Institution CRE 10435. 8 km W Quesada, 250 m, CRE 7156 UTA University of Texas at Arlington (2). Reserva San RamoÂn, UCR 10019±10026, ZMB Zoologisches Museum an der Hum- 11171, 11417, 11448, 11450, 11453, 11456. Ta- boldt-UniversitaÈt zu Berlin bacoÂn, UCR 3794±3795. Cartago: El Silencio de Sitio Mata, 1200 m, CRE 232, 235 (6), 237. Note: Some 40 specimens indicated by ``*?'' Morehouse Finca, KU 53546. Turrialba, KU after a KU number are assigned to E. megace- 28176±28182, 30771, 34992. Guanacaste: be- phalus or E. opimus on the basis of geographic tween CanÄas and TilaraÂn, KU 53595. San Bosco, probability (see page 33), although specimens 700 m, CRE 6277 (2). El Silencio de TilaraÂn, 880 from one locality (RõÂo Changena, B. del T.) con- m, CRE 7066 (2). El Silencio de TilaraÂn, 910 m, ceivably might instead represent E. gulosus. CRE 6236. 3.5 km NNE TilaraÂn, 700±760 m, Mostly collected by Myers, and by Duellman and CRE 3698±3701. 5 km NE TilaraÂn, KU 36973± Myers, these specimens were lost or misplaced in 36977. VolcaÂn Cacao, 1157 m, MVZ 207243± the University of Kansas collection before we 207244. VolcaÂn Tenorio, UCR 10672. Heredia: E could con®rm identi®cations. crest Barranca RõÂo SarapiquõÂ, opposite Isla Boni- ta, 925 m, KU 65893. La Selva, 60 m, CRE 65, Eleutherodactylus aphanus 67±69, 512, 699, 2970 (2), 3190 (3), 3385±3386, Guatemala: Izabal: 12.0 km WSW Puerto 3513, 3520±3521, 3523±3524, 3630, 4025, 4027, Santo TomaÂs, 786 m, UTA A-33405 (ad. ( ho- 6300, 6370, 6575, 8291, 8293, 8295±8296, 8361, lotype); also the paratypic series designated in 8339, 9701±9702, 9706, KU 157695±157700, original description. 157704, UCR 705, 707. Puerto Viejo, KU 36952, UCR 363. 3 km E Puerto Viejo, 50 m, CRE 9731. Eleutherodactylus biporcatus (E. maussi 4 km E Puerto Viejo, UCR 316, 324. 4±6 km E auctorum) Puerto Viejo, 50 m, CRE 3404±3406. 5 km E Puerto Viejo, UCR 3779±3780. Rara Avis, UCR Venezuela: Aragua: Rancho Grande [ϭ 910± 10611. VolcaÂn Poas, KU 28174±28175. Zona Pro- 1170 m], AMNH 70533±70552, 70554±70558, tectora La Selva, 320±400 m, CRE 7299, UCR KU 166369±166371, 166373±166374, 185671. 13275. LimoÂn: 8±13 km SW Amubre, 300±500 Miranda: Guatopo, KU 185672. m, CRE 7175. BambuÂ, 49 m, CRE 7183±7184. Cahuita, UCR 848. 8 km S Cahuita, UCR 381± Eleutherodactylus gulosus 382. Camadre, UCR 7903. Cerro Nimaso, UCR 8458, 8462±8463. El Tigre, 680 m, CRE 290 (3). Costa Rica: Limon: ``Spur of the Pico Blanco'' Sendero a Gandoco de Manzanillo, UCR 9867± [ϭ Cerro Utyum], 6,000 ft., USNM 32590 (ad. & 9869. 4 km W Guapiles, RõÂo Toro Amarillo, 260 holotype, Lithodytes gulosus). Panama: Bocas m, CRE 850. Katsi, UCR 3202. La Lola, 36±39 del Toro: GutieÂrrez, MCZ 9910±9914 (juvs). RõÂo m, CRE 206±207, 500, 8068 (3), KU 34985± Claro near junction with RõÂo Changena, 910 m, 34990, UCR 3446. Los Diamantes, UCR 726. KU 115221 (juv.). ChiriquõÂ: upper RõÂo ChiriquõÂ, Pandora, 50 m, CRE 7191±7193. Penshurst, 4 m, Fortuna Dam Site, 1000 m, AMNH 95035±95046 CRE 3474. Reserva BioloÂgica Hitoy-Cerre, UCR (juvs.). South slope Quebrada de Arena, 1120 m 11201, 11365. RõÂo Toro Amarillo, 260 m, CRE (upper RõÂo ChiriquõÂ drainage, 82Њ12Ј43ЉW, 850 (2). About 18 km E. Siquirres, 300 m, 8Њ46Ј32ЉN), AMNH 114676, 124372, CRE 10365±10366. Continental divide above upper AMNH 136244. Suretka, KU 36953±36955, UCR Quebrada de Arena, 1160±1220 m (at longitude 3206. Tortugero N. P., RõÂo Agua Fria Sta., 20 m 82Њ13ЈW), AMNH 124365±124369 (juvs.), (about 83Њ35ЈW, 8 Њ27ЈN), CRE 8841±8844. Hon- 124370, 124371. 12.6 km N Los Planes on Gual- duras: El Paraiso: Arenal, LACM 16847. Nica- aca±ChiriquõÂ Grande Road at continental divide, ragua: AtlaÂntico Norte (formerly Zelaya): Backas 1180 m, USNM 339792 (juv.). Creek, AMNH 7268±7269. Cupitna Camp, AMNH 7270±7271, 7273±7274, 7300, 7302. Eden Mine, AMNH 7276±7281. AtlaÂntico Sur Eleutherodactylus megacephalus (formerly Zelaya): Camp Corozo, RõÂo Huahu- Costa Rica: No other data, KU 34991 (11 ashan, AMNH 54970. Camp Santa Ana, RõÂo Hu- juv.). Alajuela: Boca Tapada, UCR 12595. 4 km ahuanshan, AMNH 54982±54984, 54996. Kana- S Canelete, UCR 6746. Nr. Cariblanco, 853 m, wa, AMNH 7267. RõÂo Escondido, MCZ 15022, CRE 6424. Colonia Libertad, UCR 10529. 4.8 km USNM 19909±19910. RõÂo Patch, AMNH 55026. SW El Tanque, 80 m, CRE 7154 (8). 3 km W La RõÂo Pichinga, back of Pearl Lagoon, AMNH Fortuna, 350 m, CRE 8078. 4 km W La Fortuna, 55017±55018. Matagalpa: JericoÂ, BMNH 396 m, CRE 7152 (2). La Palma, nr. Dos Rios, 94.7.26.50. 19 km N Matagalpa, UMMZ 116453. 44 AMERICAN MUSEUM NOVITATES NO. 3357

RõÂo San Juan: Poderoso, AMNH 7293. Tuli [Tule] Hydrographic Station, 150 m, AMNH 89465 (par- Creek [SE of San Miguelito], AMNH 7283, atype). ``RõÂo Chenillo'' nr. Monte Lirio [probably 7285±7286, 7289, 7291±7292, MCZ 7330±7331. Quebrada La Chinilla, inlet on Gatun Lake E of Panama: Bocas del Toro: Almirante, MCZ 9893. Monte Lirio, 9Њ14ЈN, 79Њ49ЈW], MCZ 11846 [ten- Ca. 4.8 km W Almirante, 40 m, KU 108406*?. tative identi®cation of juv. in poor condition]. Ca. 12.8 km W Almirante, 270 m, KU 108407± Ridgetop trail SW Cerro Bruja, 240±370 m, KU 108410*?. 10 km NW Almirante, FMNH 67713. 108430±108431 (paratypes). About 4 km SE 11 km NW Almirante, FMNH 67714±67716. 7.1 Puerto PiloÂn, 190±240 m, KU 113677±113678 km airline WSW ChiriquõÂ Grande, 70±100 m, (paratypes). RõÂo Guanche, 15 m, KU 172226± AMNH 113983. Isla Bastimentos, ca. 0.8 km S 172228 (paratypes). DarieÂn: Tarcarcuna, 550 m, mouth Alvarez (Alberry) Cr., USNM 297859. Isla KU 76221 (paratype). Laguna, 820 m, KU 76226 Bocas del Toro [ϭ I. ColoÂn], FMNH 67712. Isla (paratype). RõÂo Tuira at RõÂo Mono, 130 m, KU ColoÂn, La Gruta, 20±80 m, KU 108411± 108934*?.15 PanamaÂ: near community of Altos de 108414*?. Isla ColoÂn, near La Gruta, AMNH Pacora (east of Cerro Jefe), 700±800 m, KU 102239, USNM 338057±338067. Isla ColoÂn, 108432±108433 (paratypes). 4.8 km N commu- about 2.5 km N La Gruta, 40 m, AMNH 124395. nity of Altos de Pacora, on road to Mandinga, 740 East end Isla Escudo de Veraguas, 1 m, KU m, KU 108434±108435 (paratypes). Candelaria 113687±113689*?. Isla Popa, 1 km SE Deer Is- and Peluca [hydrographic] stations, near Boquer- land Channel, USNM 298035±298042. Isla Popa, oÂn, AMNH 53704 (paratype). Cerro La Victoria, 1 km E Sumwood Channel, USNM 347005± FMNH 67711. Cerro La Victoria, Quebrada 347009. Laguna de Tierra Oscura, 3.7 km S Tiger Buenos Aires, FMNH 163863±163866. Km 11.7 Key, USNM 348393±348394. La Loma (Buena- on El Llano±CartõÂ Road, KU 172229*?. Km 12.8 vista), MCZ 9955±9961. Peninsula Valiente, on El Llano±CartõÂ Road, 290 m, AMNH 89464 Blue®elds, 10 m, KU 108415±108424*?. Penin- (paratype). East slope Cerro Jefe, 650 m, KU sula Valiente, Punta Alegre, USNM 338604. 80326 (paratype). Nusagandi, nr. EstacioÂn Ibe Mouth of RõÂo Cahuita, 1 m, KU 113690± Igar, 350 m, CRE 10585, 10588. Nusagandi, 113692*?. RõÂo Claro nr. junc. with RõÂo Changena, about 2 km S EstacioÂn Ibe Igar, 300±350 m, CRE KU 113682±113683*?. RõÂo Changena, 830 m, 10604±10605. RõÂo Indio, trib. RõÂo Chagres, KU 108425±108429*?, 113684±113686*?, USNM 102852. San Blas: Camp Summit, 300 m, 115046*?. RõÂo Changuinola nr. Quebrada El Gua- KU 113679 (paratype), KU 113680±113681*?, bo (16 km airline W Almirante), 150±250 m, 113693±113694*?. AMNH 107276±107282. Southwestern headwa- ters RõÂo Guabo, 780 m (82Њ11ЈW, 8 Њ47ЈN), Eleutherodactylus rugosus AMNH 124392±124394. Southwestern headwa- ters RõÂo Guabo, 530 m (82Њ11Ј30ЉW, 8 Њ47Ј33ЉN), No speci®c locality: ``west coast of Central AMNH 124394. CocleÂ: Continental divide N El America'', USNM 32326 (holotype, Lithodytes CopeÂ, 600 m (longitude 80Њ36ЈW), AMNH pelviculus). Costa Rica: Puntarenas: Aguabuena, 98367±98369. El Valle de AntoÂn, ANSP 21750± KU 36978±36984. Quebrada Aguabuena, nr. Rin- 21751, 23500, FMNH 67710. ColoÂn: GatuÂn, coÂn de Osa, UCR 889±890, 909, 4575. 0.5 km ANSP 24855. PanamaÂ: Barro Colorado Island, NE Alfombra, 884 m, CRE 7097. Boruca, FMNH MCZ 15285±15287. Cerro Campana, FMNH 3028. RõÂo Carara, UCR 7723±7725. CajoÂn, UCR 153694±153695. Veraguas: [Atlantic coast] 7896. Cerro Puntado, 200 m, CRE 7139. El He- lechales, 1050±1150 m, CRE 8268, 8272. Esqui- mouth of RõÂo ConcepcioÂn, 1 m, KU 113676. [Pa- nas Forest Preserve, KU 34920±34922. Gol®to, ci®c versant] 5±6 mi NW (via road) Santa Fe, KU 34923. Las Cruces, 1100±1200 m, CRE 3182 AMNH 108344±108345. (6 spec.), 3433, 3485, 3487±3488, 3889, 6583, 8435, 8618±8624, 8658±8662, 8667, 8691, UCR Eleutherodactylus opimus 1680, 6175±6176. Marenco Biol. Station, UCR 13878. Palmar, KU 34910±34919. 3.2±4.8 km W Colombia: ChocoÂ: Quebrada DocordoÂ, about Palmar Norte, about 10 m, CRE 7101. 8 km NE 10 km above junction with RõÂo San Juan, about Potrero Grande, 600 m, CRE 8279. 15±25 km E 100 m, AMNH 87020 (subad. & holotype). Que- Potrero Grande, trail to 2nd savanna, 1000±1640 brada Pangala, lower RõÂo San Juan (about 17 km m, CRE 8276. RincoÂn de Osa, UCR 2651±2653, airline NE Palestina, nr. 4Њ15ЈN, 77Њ00ЈW), AMNH 109473±109487 (paratypes). Quebrada 15 Based on color notes made by W. E. Duellman dur- Taparal, lower RõÂo San Juan (about 7 km airline ing his and Myers' 1965 DarieÂn expedition into the up- NE Palestina, nr. 4Њ12ЈN, 77Њ07ЈW), AMNH per RõÂo Tuira region, the identi®cation of KU 108934 109471±109472 (paratypes). Panama: ColoÂn: as a member of the biporcatus group is open to question RõÂo BoqueroÂn, about 2.5 km (airline) N Peluca until the specimen can be found and reexamined. 2002 SAVAGE AND MYERS: ELEUTHERODACTYLUS BIPORCATUS GROUP 45

UMMZ 123621. 2 km W RincoÂn de Osa, UCR Brocchi, Paul 2066±2067, 4563, 4566. 3 km SW RincoÂn de Osa, 1882. EÂ tude des batraciens de l'AmeÂrique 40±60 m, CRE 3506, 9202, 9249±9250. 3 km W Centrale. In Mission Scienti®que au RincoÂn de Osa, 25±300 m, CRE 3112, 9614± Mexique et dans l'AmeÂrique Centrale. 9619, 9639. 3 km WSW RincoÂn de Osa, 20±40 Recherches zoologiques, pt. 3, sec. 2, m, CRE 3429±3432, 9328. 3 km WSW RincoÂn pp. 1±122 ϩ 24 pls. [pls. 1±17, 17bis., de Osa, Camp Seattle, 20±30 m, CRE 705 (5 18, 18bis., 19±20, 20bis., 21]. Paris. spec.), 752 (2), 754 (3). 3±5 km WSW RincoÂn de Campbell, Jonathan A. Osa, 20±30 m, CRE 3227. 5.5 km SW RincoÂn de 1994. A new species of Eleutherodactylus Osa, 10±30 m, CRE 3495±3497, 7235, 9387± (Anura: Leptodactylidae) of the bipor- 9388, 9665. 6 km SSW RincoÂn de Osa, 20 m, catus group from eastern Guatemala. CRE 3148. RõÂo RincoÂn, UCR 7607 12±15 mi Herpetologica 50(3): 296±302. WSW San Isidro de El General, KU 53567. San Carroll, Robert L., and Robert Holmes Pedrillo, UCR 11289. 2 km S San Vito, 1200 m, 1980. The skull and jaw musculature as CRE 916 (2 spec.). Sirena, 5 m, UCR 11290± guides to the ancestry of salamanders. 11291. Villa Neily, CRE 179. San JoseÂ: Pozo Zool. J. Linn. Soc. 68(1): 1±40. Azul, BMNH 1907.6.28.24. San Isidro del Gen- Chen, Shyh-Hwang eral, KU 34908±34909. Panama: ChiriquõÂ: No 2001. Cytogenetic study of the Lower Central other data, ZMB 7812 (juv. holotype, Hylodes ru- American frogs of the subgenus Crau- gosus). Boquete trail, MCZ 11845. Progreso, gaster (Anura: Leptodactylidae: Eleu- UMMZ 58447±58449. therodactylus). Unpublished Ph.D. diss., Univ. Miami, xiii ϩ 369 pp. REFERENCES Cochran, Doris M., and Coleman J. Goin 1961. A new genus and species of frog (Lep- Baird, Irwin L. todactylidae) from Colombia. Fieldiana 1951. An anatomical study of certain sala- Zool. 39(48): 543±546. manders of the genus Pseudoeurycea. Cope, Edward Drinker Univ. Kansas Sci. Bull. 34, pt. 1(6): 1862. On some new and little known Ameri- 221±265. can Anura. Proc. Acad. Nat. Sci. Phil- Beebe, William, and Jocelyn Crane adelphia 14: 151±159. 1947. Ecology of Rancho Grande, a subtrop- 1875. (November 26) On the Batrachia and ical cloud forest in northern Venezuela. Reptilia of Costa Rica. J. Acad. Nat. Zoologica 32(1): 43±59 ϩ pls. 1±5. Sci. Philadelphia. ser. 2, vol. 8, pt. 2: Bemis, William E., Kurt Schwenk, andMarvalee 93±154 ϩ pls. 23±28. [Publication date H. Wake is that of author's separates, issued in 1983. Morphology and function of the feed- advance] ing apparatus in Dermophis mexicanus 1877. (August 15) Tenth contribution to the (Amphibia: Gymnophiona). Zool. J. herpetology of tropical America. Proc. Linn. Soc. 77(1): 75±96. Am. Philos. Soc. 17: 85±98. [Publica- Boettger, Oskar tion date is that of author's separates, 1893. Reptilien und Batrachier aus Venezue- issued in advance] la. Ber. Senckenb. Naturforsch. Ges. 1893. (December 30) Second addition to a 1893: 35±42. knowledge of the Batrachia and Repti- Bogart, James P. lia of Costa Rica. Proc. Am. Philos. ``1970'' [1973]. Los cromosomas de an®bios Soc. 31: 333±347. [Publication date is anuros del geÂnero Eleutherodactylus. that of author's separates, issued in ad- Actas IV Congreso Latinamer. Zool. 1: vance] 65±78. DeWeese, James E. 1981. Chromosome studies in Sminthillus 1976. The karyotypes of Middle American from Cuba and Eleutherodactylus from frogs of the genus Eleutherodactylus Cuba and Puerto Rico (Anura: Lepto- (Anura: Leptodactylidae): case study of dactylidae). R. Ont. Mus. Life Sci. the signi®cance of the karyologic meth- Contrib. 129: 22 pp. od. Unpublished Ph.D. diss., Univ. Boulenger, George Albert South. California, Los Angeles, x ϩ 1882. Catalogue of the Batrachia Salientia s. 210 pp. Ecaudata in the collections of the Brit- DumeÂril, A. M. Constant, and Gabriel Bibron ish Museum. London: British Museum, 1841. ErpeÂtologie geÂneÂral ou histoire naturel- xvi ϩ 503 pp. ϩ 30 pls. le compleÁte des reptiles. Paris, Librairie 46 AMERICAN MUSEUM NOVITATES NO. 3357

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