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Two New Species Of The Eleutherodactylus Rugulosus Group (Amphibia : Anura : Leptodactylidae) From Honduras PDF

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Preview Two New Species Of The Eleutherodactylus Rugulosus Group (Amphibia : Anura : Leptodactylidae) From Honduras

1 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 112(3):515-522. 1999. Two new species of the Eleutherodactylus rugulosus group (Amphibia: Anura: Leptodactylidae) from Honduras James R. McCranie and Larry David Wilson (JRM) 10770 SW 164th Street, Miami, Florida 33157-2933, U.S.A.; (LDW) Department of Biology, Miami-Dade Community College, Kendall Campus, Miami, Florida 33176, U.S.A. — Abstract. Two new species ofEleutherodactylus, E. pechorum and E, olan- chano, of the E. rugulosus group are described from Honduras. The following combination of characters will distinguish E. pechorum from the remaining E. rugulosus group members: male vocal slits present; male nuptial thumb pad < absent; finger II I; female loreal length/eye length ratio <1.00; keels present on unwebbed portions of toes; toes with basal webbing; canthus rounded; pos- terior surface of thighs marked with numerous pale spots and blotches; antero- ventral and posteroventral surfaces of tibial segments barred; ventral surfaces pale yellow; and size moderate. Eleutherodactylus olanchano is distinguished from all other E. rugulosus group species by the following combination of characters: male vocal slits and nuptial thumb pads absent; finger II > I or II « I; tympanum length/eye length ratio >1.00; and male size very small (fe- males unknown). Recent fieldwork in Honduras has result- Savage will likely severely restrict the geo- ed in the discovery of two new species of graphic distribution of E. rugulosus. Thus, the Eleutherodactylus rugulosus group. the specific identity of the Honduran and One of these species is a streamside dwell- Nicaraguan material of E. ''rugulosus'' ex- er, as are most members of the group. How- amined is unresolved at this time. Abbre- ever, the other species occurs on forested viations used in each species description are hillsides, well above streams that are inhab- EL (eye length), E-N (anterior border of ited by E. aurilegulus, another rugulosus eye to posterior edge of nostril; equals lo- HL group member. The forest dwelling species real length of Savage 1975), (head HW represents only the third known member of length; tip of snout to angle of jaw), the rugulosus group occurring in that type (greatest width of head), LL (leg length), of habitat (Campbell et al. 1994, Johnson SL (snout length; anterior border of eye to & SVL TL Savage 1995). tip of snout), (snout-vent length), TM (tibia length), and (tympanum length). Materials and Methods Measurements were made to the nearest 0. millimeter under a dissecting microscope. All morphological and color pattern traits Museum acronyms follow those of Leviton discussed herein follow the character states et al. (1985) and color codes are those of and terminology defined by Savage (1975). Smithe (1975-1981). Comparative data for E. azueroensis, E. Systematics taurus, and E. vocalis were taken from Sav- age (1975). Comparative material examined Eleutherodactylus pechorum, new species is listed in Appendix I. The name rugulosus — Fig. 1 is in quotes in this appendix because work Holotype. National Museum of Natural in progress by J. A. Campbell and J. M. History (USNM) 530000, an adult male, 516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Adult male holotype of Eleutherodactylus pechorum (USNM 530000), SVL 33.5 mm. from near a small dam along a small trib- >1.000, n = 4), and larger female size (to utary of the Quebrada de Las Marias 59 mm SVL versus to 41 mm, w = 4). In (15°17.64'N, 85°21.29'W), about 12 airUne addition, E. anciano is known only from km NNE La Colonia, Departamento de remnants of broadleaf cloud forest (Lower m Olancho, Honduras, 680 elev., collected Montane Moist Forest formation of Hold- m 2 Aug 1998 by J. R. McCranie, K. L. Wil- ridge 1967) at 1770 to 1840 elevation in liams, and L. D. Wilson. Original number southwestern Honduras. The new species LDW 11375. differs from E. azueroensis of the Peninsula Paratype.—\5SNM 529999, and adult fe- de Azuero, Panama, in having the posterior male with the same locality data as the ho- surface of the thighs marked with numerous lotype, except collect—ed 1 Aug 1998. pale spots and blotches (a few small pale BMNH Referred specimen. 1985.1454, spots in E. azueroensis), the anteroventral an adult female from along the Rio Cuya- and posteroventral surfaces of the tibial mel, Departamento de Colon, Honduras. segments barred (uniformly brown), the — Diagnosis. The combination of the canthus rounded (moderately sharp), and presence of vocal sUts and the absence of basal toe webbing (moderately webbed; nuptial thumb pads in males will distin- webbing formula H 2-3 1/2 III 3-4 1/3 IV guish E. pechorum from all other described 4 1/3-3" V in E. pechorum versus II 2"-3 members of the E. rugulosus group, except 1/4 III 2 2/3-4 IV 4-2 1/2 V in E. azue- for E. anciano, E. azueroensis, E. taurus, roensis). Eleutherodactylus pechorum dif- and E. vocalis. Eleutherodactyluspechorum fers from E. taurus of southwestern Costa is distinguished from E. anciano by having Rica and adjacent Panama in having toe finger II shorter than finger I (finger II lon- keels on unwebbed portions of toes (well ger than or equal in length to finger I in E. developed fleshy toe fringes in E. taurus), anciano), toe keels on toe IV of females four and one-third segments on toe IV free (weakly infolded fringes on at least one side of webbing (<4), definite toe discs (barely of toe rV in all four females), shorter fe- expanded), and females to 59 mm SVL (fe- male snouts (E-N/EL < 1.000 versus males to 80 mm). The new species can be VOLUME NUMBER 112, 3 517 distinguished from E. vocalis of western to median portion of orbit; supratympanic Mexico by having finger II shorter than fin- fold slightly darker brown than adjacent ger I (finger II longer than or equal in area; groin mottled with dark brown and length to finger I in E. vocalis), ventral sur- white; posterior surface of thighs dark faces pale yellow (white), and the antero- brown with numerous cream spots and ventral and posteroventral surfaces of the blotches; anterior surface of thighs pale tibial segments barred (suffused). brown with dark brown vertical bars; un- Descript—ion of holotype (alcohol after derside of thighs and tibial segments cream formalin). An adult male with the follow- with very sparse dark brown flecks, barred ing measurements (percentages of SVL in with dark brown along margins. parentheses): SVL 33.5 nmi; HL 13.1 mm Color in life (based on a Kodachrome® HW mm mm (39.1); 12.5 (37.3); EL 3.9 slide, except for ventral coloration, which (11.6); SL 5.6 mm (16.7); E-N 3.4 mm was recorded in life): dorsal surfaces of (10.1); TM 3.5 mm (10.4); LL 56.5 mm head and body Verona Brown (223B), with mm (168.7); and TL 19.6 (58.5). Dorsum slightly darker brown mottling and slightly smooth with several enlarged tubercles on paler brown, scattered spots; postorbital upper eyelids, short postorbital ridges also ridges mottled Verona Brown (223B) and present; snout nearly rounded in dorsal as- slightly darker brown; dorsal surfaces of pect, rounded in profile; canthus rostralis limbs Raw Umber (23), with darker brown rounded; tympanum prominent; first finger crossbands; interorbital region mottled pale longer than second; finger discs definite, a brown, medium brown, and dark brown, little over twice width of digitjust proximal suggestive of pale interorbital bar outlined to discs on fingers III-IV; well developed with darker brown; dark brown upper lip inner tarsal fold; heels rugose; plantar tu- bars separated by pale brown; tympanum bercles absent; inner metatarsal tubercle intermediate between Verona Brown elongated, elevated, visible from above; (223B) and darker brown mottling; iris cop- outer metatarsal tubercle small, round, bare- pery brown, reticulated with black; belly ly elevated; toe discs definite, that on toe and ventral surfaces of thighs pale yellow. IV about 1.6 times width of digit just prox- Varia—tion in paratype (alcohol after for- imal to disc; toes with marginal ridge; toes malin). Color is described as follows: bel- basally webbed, webbing formula I 2-2 1/ ly, chest, and throat cream, lightly flecked 2 II 2-3 1/2 III 3-4 1/3 IV 4 1/3-3" V; with grayish-brown; dorsum grayish-brown vomerine tooth patches on elevated, nearly with distinct dark brown mottling and elon- triangular-shaped ridges located postero- gated, narrow spots; several dirty white medially to ovoid choanae, tooth patches small spots also present on lower back; separated by distance less than width of ei- postorbital ridges distinctly darker brown ther patch; paired vocal slits present; nuptial than adjacent dorsum; dark line absent pads absent. Belly cream, lightly flecked along canthal ridges, although several dark with dark brown; throat and chest cream, flecks present just anterior to eye; loreal re- moderately flecked with dark brown; dor- gion pale brown; upper and lower lips with sum brown with indistinct dark brown mot- medium brown bars separated by dirty tling, several white spots scattered on back; white lines, dark lower lip bars not extend- postorbital ridges slightly darker brown ing onto chin; top of head grayish-brown than adjacent dorsum; dark line absent with slightly paler brown interorbital bar, along canthal ridges; loreal region brown; pale bar bordered anteriorly and posteriorly upper and lower lips with dark brown bars by dark brown; supratympanic fold darker separated by white lines, dark lower lip bars brown than adjacent area; groin mottled not extending onto chin; top of head brown with grayish-brown and pale brown, several with slightly paler interorbital bar anterior dark brown spots also present; posterior 518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON surface of thighs grayish-brown with nu- long, but sparsely populated by the indig- merous cream spots and blotches; anterior enous Pech tribe. surface of thighs pale brown with grayish- brown vertical bars; underside ofthighs and Eleutherodactylus olanchano, new species tibial segments cream, barred with grayish- Fig. 2 brown along margins. — The following measurements (percent- Holotype. National Museum of Natural ages of SVL in parentheses) were recorded: History (USNM) 529998, an adult male, SVL 41.2 mm; HL 17.0 mm (41.3); HW from a hillside above the Quebrada El Pinol 16.0 mm (38.8); LL 70.7 mm (171.6); and (15°07'N, 86°44'W), Parque Nacional La TL 24.8 mm (60.2). — Muralla, Depmartamento de Olancho, Hon- Comments on referred specimen. A sin- duras, 1200 elev., collected 14 Aug 1994 gle adult female (BMNH 1985.1454; SVL by J. R. McCranie and L. D. Wilson. Orig- LDW 59.1 mm) from about 50 airline km NNE inal number 10319. Paratypes.—US^M 529991-97, of the type locality of E. pechorum is re- all ferred to this species. This specimen is very adult males,mall from the type locality, similar in color pattern and morphology to 1180-1200 elev., coUected 21 Jul 1993 (USNM 529996-97) or 12-14 Aug 1994 the female paratype of E. pechorum. Also, although the type locality for E. pechorum (the remaining specimens). USNM is in the headwaters of the Rio Wampu, the Referred specimens.— 529989- locality is only some 15 km from the head- 90, both adult males, from near Rio de En- medio (14°52'N, 86°48'W), Montana El Ar- waters of the Rio Paulaya. The locality for BMNH mado, Departamento de Olancho, Hondu- the specimen is along a tributary of m ras, 1350 e—lev., coUected 13 Jun 1993. the latter river. Additionally, E. epochthi- Diagnosis. ^The combination of small dius, another streamside Eleutherodactylus adult male size (SVL 18.7-29.8, X = 24.8; (E. milesi group), is known from both the females unknown), large male tympanum E. pechorum type localiBtyMNanHd from a lo- (TM/EL 1.00-1.14, X = 1.06), finger H cality very near that for 1985.1454. — longer than or equal in length to finger I, Natural history notes. Both type spec- and males lacking vocal sUts and nuptial imens were collected at night alongside a thumb pads will distinguish E. olanchano small stream flowing through nearly pris- m from all other described species of the E. tine forest at 680 elevation. However, at m rugulosus group. Eleutherodactylus olan- 660 elevation, the former forest around chano is the smallest known member of the this stream has been cleared, as have most E. rugulosus group. Males of all other spe- of the hillsides surrounding the forest on mm SVL cies reach at least 33 (the single both sides of the stream. The forest along- known adult males of E. anciano and E. side this portion of the stream remains in- pechorum), 38 mm SVL {E. vocalis), or 40 tact only because water from that portion of mm or longer (the remaining E. rugulosus the stream is piped to several villages below group species: Savage 1975, Savage et al. the stream. This extensive deforestation is 1988, Campbell et al. 1994, Johnson & taking place even though the region is part Savage 1995). of the Rio Platano Biosphere Reserve, es- Description of holotype (alcohol after — tabhshed as a World Heritage Site in 1980. formalin). An adult male with the follow- The type locaHty is in the Premontane Wet ing measurements (percentages of SVL in Forest formati—on of Holdridge (1967). parentheses): SVL 29.8 mm; HL 13.1 mm HW mm mm Etymology. The name pechorum means (44.0); 11.7 (39.3); EL 3.5 mm mm "belonging to or pertaining to" the Pech, (11.7); SL 4.8 (16.1); E-N 3.0 in reference to this frog inhabiting an area (10.1); TM 4.0 mm (13.4); LL 55.6 mm VOLUME NUMBER 112, 3 519 Fig. 2. Adult male holotype of Eleutherodactylus olanchano (USNM 529998), SVL 29.8 mm. mm (186.6); and TL 17.6 (59.1). Dorsum brown bars separated by cream lines, dark smooth to weakly granular, with several lower lip bars not extending onto chin; top distinct tubercles on lower back and upper of head pale brown with a few dark brown eyelids; postorbital ridges absent; canthus small spots in interocular region near mid rostralis rounded; tympanum prominent; level of orbit; supratympanic fold very dark first finger equal in length to second; finger brown; groin cream, heavily flecked with discs definite, about 1.8 times width ofdigit dark brown; posterior surface of thighs dark just proximal to discs on fingers III-IV; brown, with indistinct pale brown blotches; well developed inner tarsal fold; toe discs anterior surface of thighs pale brown, with definite, that on toe IV about 2.0 times slightly darker, incomplete vertical bars; un- width of digit; toes with marginal ridge; derside of thighs and tibial segments cream, toes moderately webbed, webbing formula with very sparse dark brown flecks, barred I 2-2 1/2 II 2-3 1/3 III 3-4 IV 4-2 3/4 V; with dark brown along margins. vocal slits absent; nuptial pads absent. Belly Color in life (based on a Kodachrome® cream, very lightly flecked with dark slide, except for ventral coloration, which brown; chest cream, heavily flecked with was recorded in life): dorsal surfaces of dark brown; throat heavily flecked with head and body Cinnamon-Rufous (40), with dark brown; dorsum dark brown with sev- several dark brown spots in postorbital and eral very dark brown markings in postocu- interorbital regions and slightly darker lar region; dark fine absent along canthal brown mottling on lower back; dorsal sur- ridges; loreal region mottled pale and dark faces of limbs Cinnamon-Rufous (40), with brown; upper and lower lips with dark darker brown, somewhat indistinct, cross- 520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bands; dark brown upper lip bars separated adult males (USNM 529989-90; 26.5 and by pale brown; supratympanic folds dark 24.0 nmi SVL, respectively) from about 30 brown; tympanum pale brown; iris reddish- airline km SSW of the type locality agree brown, reticulated with dark brown on up- in all diagnostic features with those of the per half, grayish-brown, reticulated with type series. These specimens were collected dark brown on lower half, both halves not by M. R. Espinal. His collection from the separated by dark band; belly and ventral Rio de Enmedio region also contains two (USNM surfaces of thighs pale yellow. adult females of E. aurilegulus mm Variatio—n in type series (alcohol after 529987-88; 53.5 and 53.6 SVL, re- formalin). Belly coloration of all para- spectively). Espinal did not record any hab- types is similar to that of holotype; most itat information for this collection, but it specimens have less brown flecking on seems likely that the E. olanchano were chest and throat than does holotype; how- taken from forested hillsides above the ever, one specimen (USNM 529992) has streamside locality where E. aurilegulus throat and chest more intensely punctated would be expected to occur. — with dark brown; dorsal surfaces of head Natural history notes. The type series and body vary from grayish-brown to dark was collected both during the day and at brown, with varying amounts of dark night while active among leaves on the for- brown spots or lines present in postorbital est floor on a single hillside above the Que- and interocular regions, these dark marks brada El Pinol. Specimens were collected indistinct in darker specimens; one speci- in July and August. Two species of stream- men (USNM 529992) has a rather broad, side Eleutherodactylus (E. aurilegulus and pale brown middorsal stripe extending from E. stadelmani; E. rugulosus and E. milesi tip of snout to just above vent; most spec- groups, respectively) were common along- imens have indistinct pale brown blotches side the Quebrada El Pinol at the base of on posterior surface of thighs; however, the hillside, while E. lauraster (E. rhodopis these blotches can be rather distinct in a few group) was found in the same habitat as the USNM specimens (e.g., 529991-92); un- E. olanchano. This locality is in the Pre- derside of thigh and tibial segments similar montane Wet Forest formation ofHoldridge to that of holotype in all specimens, except (1967), as is the locality for the referred that one specimen (USNM 529992) has specimens. The known elevational range slightly more flecking medially on thighs for this species—is 1180 to 1350 m. near knees than does the holotype. Etymology. The name olanchano is Morphological measurements of the en- Spanish, meaning "native ofOlancho," and tire type series include SVL given in mil- is used in reference to this Honduran de- limeters and other measurements as per- partment, in which this species is apparent- centages of SVL, range followed by mean ly endemic. The name is a noun used in SVL HL in parentheses: 18.7-29.8 (24.8); apposition to the generic name. HW 40.3-44.6 (43.5); 37.1-40.1 (38.6); LL 168.8-208.6 (191.0); and TL 56.4-62.9 Discussion (59.4). Additionally TM/EL ranges and means are 1.00-1.14 (1.06) in the entire Members of the Eleutherodactylus ru- type series, and finger II varies from being gulosus group occur from southern San longer than finger I to being equal in length Luis Potosi, Mexico on the Atlantic versant to finger I. Examination of the testes of two and from northern Sinaloa, Mexico on the specimens (USNM 529993-94; 28.1 and Pacific versant southward to central Panama mm 23.1 SVL, respectively) confirmed that (Savage 1975). The two new species de- they are adults. scribed herein appear to have relatively — Comments on referred specimens. Two small geographical distributions as do many VOLUME NUMBER 112, 3 521 Other Central American members of this G. Schneider (UMMZ), S. W. Gotte and R. group (Savage, pers. comm.). W. McDiarmid (USNM), and R. Gunther Among the described species of the E. (ZMB). An early draft of this manuscript rugulosus group, E. pechorum appears to be was reviewed by J. M. Savage, who also most similar morphologically to E. azue- provided many helpful comments concern- roensis of low and moderate elevations ing the systematics of the E. rugulosus (60-940 m) of the Peninsula de Azuero, group. Panama. However, the intervening lowlands (with the exception of the Golfo Dulce area Literature Cited of the Pacific versant of southern Costa Rica and adjacent Panama where E. taurus CampbelAl,nJe.wA.,speJ.ciMe.s SoafvaEglee,ut&herJo.dRa.ctMyleuyser.(An1u9r9a4:. occurs) are inhabited by one or more mod- Leptodactylidae) of the rugulosus group from — erately webbed species of the E. rugulosus Guatemala and Belize. Herpetologica 50:412- group in which the males lack vocal slits 419. and nuptial thumb pads. No specimens from Holdridge, L. R. 1967. Life zone ecology. Revised edi- tion. Tropical Science Center, San Jose, Costa the intervening territory resemble either E. Rica, 206 pp. pechorum or E. azueroensis in the subtle Johnson, J. D., & J. M. Savage. 1995. A new species features that distinguish species in this of the Eleutherodactylus rugulosus group (Lep- — group. The localities for E. pechorum lie in todactylidae) from Chiapas, Mexico. Journal excess of 900 km NNW of the nearest of Herpetology 29:501-506. & known locality for E. azueroensis. Leviton, A. E., R. H. Gibbs, Jr., E. Heal, C. E. Dawson. 1985. Standards in herpetology and The second new species described herein ichthyology: part I. Standard symbolic codes for {E. olanchano) possesses a combination of institutional resource collections in herpetology — male characters (very small size, very large and ichthyology. Copeia 1985:802-832. tympanum, finger II longer than or equal in Savage, J. M. 1975. Systematics and distribution ofthe length to finger I, and the absence of vocal Mexican and Central American stream—frogs re- lated to Eleutherodactylus rugulosus. Copeia slits and nuptial thumb pads) not shared 1975:254-306. with any other described member of the , J. R. McCranie, & L. D. Wilson. 1988. New group. upland stream frogs of the Eleutherodactylus rugulosus group (Amphibia: Anura: Leptodac- — tylidae) from Honduras. Bulletin Southern Acknowledgments California Academy of Sciences 87:50-56. Smithe, F. B. 1975-1981. Naturahst's color guide, part Collecting and exportation permits were I. Color guide. The American Museum of Nat- provided by M. Acosta, A. Barahona, E. ural History, New York, 182 color swatches. Munoz, G. Rodriguez, and C. Romero of COHDEFOR, Tegucigalpa. Field assistance Appendix I was provided by D. Almendarez, M. R. Es- Comparative material examined — pinal, G. A. Flores, and K. L. Williams. Es- Eleutherodactylus anciano. Honduras Ocote- pinal also donated the specimens from his peque: El Chaguiton, KU 208999-9001, ROM 18076- Rio de Enmedio collection and provided 80. — valuable help in obtaining the COHDEFOR Eleutherodactylus aurilegulus. Honduras Atlanti- AMNH MCZ da: Lancetilla, 54792-96, 16184-88, permits. Comparative material was provid- 16280, 16284-85, 17448-50, 21270; 2.0 km SE Lan- ed by C. J. Cole (AMNH), B. T. Clarke cetilla, TCWC 30129; mountains S ofLancetilla. MCZ (BMNH), W. W. Tanner (BYU), E. J. Cen- 16191; 7 km NW Las Mangas. UF 90216; Quebrada sky and J. J. Wiens (CM), A. Resetar La Muralla. SMF 77636-39; Quebrada de Oro, KU LACM (FMNH), J. E. Cadle and J. Rosado (MCZ), U20M90M0Z2-32, 209033 (3), 13A72M8N6-H97; Tela, A. Holman (MSUM), R. W. Murphy 70329 (7). CoClMon: Balfate. 45696- 703, 45718; Trujillo, 63920-21. Olancho: Que- (ROM), G. Kohler (SMF), K. R. Vaughan brada La Calentura, USNM 343704; Quebrada Las (TCWC), D. L. Auth (UF), A. G. Kluge and Cantinas, USNM 343710-14; between El Dictamo and 522 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON AMNH Parque Nacional La Muralla Centre de Visitantes, mayagua: between La Mision and Trincheras, USNM 343625-36; Rio de Enmedio. USNM 529987- 54752; Siguatepeque, MCZ 26421. Cortes: AguaAzul, 88: Quebrada La Habana. USNM 343705-09: Quebra- AMNH 54951: Rio Guayabal, AMNH 54751; Haci- USNM FMNH da de Las Mesetas, 343695-702: Quebrada del enda Santa Ana, 4661 (3), 4663, 4665-69, Monte Escondido. USNM 343703: near Parque Na- 4691-93, 4695-97, MCZ 17433-34, 21282, SMF cional La Muralla Centro de Visitantes. USNM 29865; Caiion Santa Ana, FMNH 4671, 4674-75; 343622-24: Quebrada El Pinol, USNM 343657-94: Lago de Yojoa, MSUM 4540, 4637. El Paraiso: ca. 6 confluence of quebradas El Pinol and Las Cantinas, km E Danli, TCWC 23811; Danli, BYU 18209; Mon- USNM USNM AMNH MCZ 343638-56: near Los Planes, 343637: serrat, 54814; Montafia del Volcan, Quebrada Salitre Lajas, USNM 343715-16: Sendero 26433-35. Francisco Morazan: Agua Amarilla, USNM AMNH AMNH El Pizote. 343717-19. Yoro: Monte Mataderos, 54883; San Francisco, 54750; Cerro MCZ FMNH MCZ MCZ 21287: Portillo Grande, 21859-60, Uyuca, 26440; Rio Yeguare Valley, MCZ UMMZ UMMZ MCZ 34696-97. 21273-75, 21289, 77852 (3): 25954-56, 94055 (6); El Zamorano, Santa Marta, FMNH 21857-58, MCZ 21271-72, 26376-77, 26469. Intibuca: 5 km NE Jesus de Otoro, UMMZ FMNH TCWC 21288, 77851 (3): Subirana Valley. 23810. Santa Barbara: Montafia de Santa Bar- 21782, MCZ 21283-86; 6.6 km S Yoro, MVZ bara, AMNH 55309-12. '^HONDURAS,*' ZMB W T17C1W37C5-7263,619U:SNcaM. 32217k58m2-W83;Yocrao.,3MkVmZ 175Y8o0r0o., c1a3r2a0g2ua(—hoAltoltaynpteicoof HSuyrl:odReisolaCehviiqsusiitmou,s SWeMrFner7)7.82N5i.- USNM 217584-85. Granada: Volcan Mombacho, SMF 78222-28. Mata- — Eleutherodactylus ''rugulosus.'' Honduras Co- galpa: Selva Negra, SMF 78207-21.

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