NORTHERN ECUADOR URL: ...multimedia20stg.blob.core.windows.net/...alcohol-preserved specimens of frogs from the herpetological collections at Museo de Zoologı´a of the Universidad

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TWO NEW SPECIES OF FROGS (LEPTODACTYLIDAEELEUTHERODACTYLUS) FROM THE HIGH ANDES OFNORTHERN ECUADORAuthor(s) Juan M Guayasamin Diego Almeida-Reinoso Fernando Nogales-SornosaSource Herpetological Monographs 18(1)127-141Published By The Herpetologists LeagueDOI httpdxdoiorg1016550733-1347(2004)018[0127TNSOFL]20CO2URL httpwwwbiooneorgdoifull1016550733-1347282004290185B01273ATNSOFL5D20CO3B2

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Herpetological Monographs 18 2004 127ndash141 2004 by The Herpetologistsrsquo League Inc

TWO NEW SPECIES OF FROGS (LEPTODACTYLIDAEELEUTHERODACTYLUS) FROM THE HIGH ANDES OF

NORTHERN ECUADOR

JUAN M GUAYASAMIN14 DIEGO ALMEIDA-REINOSO

2 AND FERNANDO NOGALES-SORNOSA3

1Natural History Museum amp Biodiversity Research Center Department of Ecology and Evolutionary BiologyThe University of Kansas Lawrence KS 66045-7561 USA

2Universidad Central del Ecuador Facultad de Filosofıa Letras y Ciencias de la EducacionEscuela de Biologıa y Quımica Av America Quito Ecuador

3Fundacion Ecologica Arcoiris Segundo Cueva Celi 03-15 y Clodoveo CarrionCiudadela Zamora Loja Ecuador

ABSTRACT We describe two new species of Eleutherodactylus that are hypothesized to belong to the Eorcesi Group Both species are found in Andean habitats of northern Ecuador at elevations above 3000 m Thepresence of an anteriorly exposed frontoparietal fontanelle distinguishes the two new species from most othermembers of the genus in which the frontoparietal fontanelle is covered by frontoparietal bones Additionallyboth species have fingers and toes with fleshy lateral fringes and vomerine teeth reduced or absent The newspecies differ from one another mostly by tuberculation and coloration patterns

Key words Andes Ecuador Eleutherodactylus orcesi Leptodactylidae New species

THE ELEUTHERODACTYLUS ORCESI GROUP wasrecognized by Lynch (1981) for an assemblageof species distributed in the high Andes(gt3000 m) of Colombia and Ecuador Theonly diagnostic feature cited for the group wasthe presence of an anteriorly exposed fronto-parietal fontanelle (Lynch et al 1996) Cur-rently the Eleutherodactylus orcesi Groupcontains six species E obmutescens Lynch1980 E orcesi Lynch 1972 E racemus Lynch1980 E simoteriscus Lynch et al 1996 Esimoterus Lynch 1980 and E thymelensisLynch 1972 All of these species live inparamos at elevations of 3000ndash4150 m andreplace one another along the geographic axisof the Andes of Colombia and Ecuador(Lynch 1981 Lynch et al 1997) If the groupis monophyletic its pattern of distribution iscongruent with an allopatric mode of specia-tion (Lynch et al 1997)

It is not surprising that fieldwork in poorlycollected Andean localities results in thediscovery of undescribed species Herpetolog-ical expeditions by Oxford University of UK incooperation with the Pontificia UniversidadCatolica del Ecuador and the UniversidadCentral del Ecuador to localities in montaneforests and paramo in the Andes of northernEcuador have revealed two more species of the

Eleutherodactylus orcesi Group Herein wedescribe these newly discovered species

MATERIALS AND METHODS

Specimens were euthanized in the field byimmersion in chloretone fixed in 10 forma-lin and preserved in 70 ethanol as sug-gested by Simmons (2002) We examinedalcohol-preserved specimens of frogs fromthe herpetological collections at Museo deZoologıa of the Universidad Catolica delEcuador (QCAZ) Natural History Museumof The University of Kansas (KU) andInstituto de Ciencias Naturales of the Uni-versidad Nacional de Colombia (ICN) Inaddition to the type series of the two newspecies specimens examined are listed inAppendix I Morphological measurementswere taken as described in Guayasamin(2004) and are (1) snoutndashvent length (SVL)(2) tibia length (3) foot length (4) head length(5) head width (6) interorbital distance (7)upper eyelid width (8) internarial distance (9)eye-to-nostril distance (10) snout-to-eye dis-tance (11) eye diameter (12) tympanumdiameter (13) eye-to-tympanum distance(14) radioulna length (15) hand length (16)Finger-I length Sexual maturity was deter-mined by the presence of eggs or convolutedoviducts in females and by the presence ofvocal slits in males For ease of comparison4 CORRESPONDENCE e-mail juanmkuedu

127

the diagnosis and description of the newspecies follow those of Lynch and Duellman(1997) Terminology for digital disc features isthat described by Savage (1987 1997)

SYSTEMATICS

Eleutherodactylus huicundo sp nov

Figs 1ndash4 8

HolotypemdashQCAZ 14753 an adult femalecollected by Diego Almeida near El Playon deSan Francisco (approx 778409 W 008389 Nelevation 3400 m) Provincia SucumbıosEcuador on 26 August 2000

ParatypesmdashAll paratypes were collectednear El Playon de San Francisco and havethe same data as holotype except as followsQCAZ 14751 an adult male collected byFernando Nogales at an elevation of 3600 mQCAZ 14752 an adult female collected byFernando Nogales QCAZ 14745ndash46 adultfemales collected by Diego Almeida at anelevation of 3350 m QCAZ 14747 an adultmale collected by Fernando Nogales at anelevation of 3700 m QCAZ 14754 an adultmale collected by Nora Schultz QCAZ 14768an adult female collected by Diego AlmeidaQCAZ 15391 an adult female collected byJuan M Guayasamin Elisa A Bonaccorso andDiego Almeida on 5 June 2001 QCAZ 15392QCAZ 15394 adult males collected by Elisa ABonaccorso Juan M Guayasamin and DiegoAlmeida at an elevation of 3400ndash3600 m on 5June 2001 QCAZ 14748ndash50 juveniles col-

lected by Nora Schultz at an elevation of 3229ndash3240 m on August 2000 and QCAZ 15395juvenile collected by Diego Almeida Juan MGuayasamin and Elisa A Bonaccorso at anelevation of 3400ndash3600 m on 5 June 2001

DiagnosismdashEleutherodactylus huicundo asother species in the Eleutherodactylus (Eleu-therodactylus) orcesi Group has an anteriorlyexposed frontoparietal fontanelle AdditionallyE huicundo is diagnosed by having (1) skin ofdorsum shagreen to warty that of flanks withsome flat warts venter areolate dorsolateralfolds absent (2) tympanic annulus evidentround 364ndash480 of eye length (xx 5 428 631 n 5 11) tympanic membrane poorlydefined (3) eye-to-nostril distance 647ndash862eye length (xx 5 745 6 67 n 5 11) snoutrounded in dorsal view angular in lateral viewusually with papilla at tip (Figs 3 8) (4) uppereyelid narrower than interorbital space withsome low tubercles and one enlarged non-conical tubercle cranial crests absent (5)vomerine teeth absent or reduced and difficultto see (6) males with vocal slits and mediansubgular vocal sac nuptial pads absent (7)Finger I shorter than Finger II fingers withexpanded discs (Fig 4) (8) fingers with fleshylateral fringes (Fig 4) (9) ulnar tuberclessmall two or three in number (10) heel withone enlarged non-conical tubercle tarsus withtwo or three tubercles along outer edge (11)two metatarsal tubercles prominent inneroval two to three times size of subconicalouter tubercle supernumerary plantar tu-bercles numerous (Fig 4) (12) toes with

FIG 1mdashDorsal (A) and ventral (B) views of paratype of Eleutherodactylus huicundo QCAZ 15394 SVL 5 198 mmmale

128 HERPETOLOGICAL MONOGRAPHS [No 18

fleshy lateral fringes (Fig 4) (13) in ethanoldorsum usually gray to dark gray with darkermarkings outlined with thin black and whitelines throat and venter gray to dark gray ordark brown with small darker spots palmarand plantar surfaces dark gray with Fingers Iand II being paler (Figs 2 4) and (14) adultssmall males 198ndash213 mm SVL (xx 5 206 606 n 5 5) females 232ndash287 mm SVL (xx 5257 6 23 n 5 6)

Comparison with similar speciesmdashThepresence of an anteriorly exposed frontopa-rietal fontanelle distinguishes the species ofthe Eleutherodactylus orcesi Group from mostother members of the genus in which thefrontoparietal fontanelle is covered by fronto-parietal bones (Lynch 1971 Lynch et al

1996) Differences among species in the Eorcesi Group are in Table 1 Additionally Ehuicundo is distinguished from E ortizi spnov by having tarsal tubercles (usually absentin E ortizi) males of E huicundo haveshagreen to warty flanks (areolate in malesand females of E ortizi) and pronouncedtubercles on the upper eyelid forearm heeland tarsus (tubercles small or absent in malesand females of E ortizi) Eleutherodactylushuicundo differs from E orcesi E simoter-iscus and E simoterus by having tubercles onupper eyelid forearm and tarsus (absent in Eorcesi E simoteriscus and E simoterusLynch 1980 Lynch et al 1996) from Eracemus by the presence of canthal andsupratympanic stripes (absent in E racemus)

FIG 2mdashDorsal and ventral views of holotype (A B) of Eleutherodactylus huicundo QCAZ 14753 SVL 5 284 mmfemale and of paratype (C D) QCAZ 15394 SVL 5 198 mm male

2004] HERPETOLOGICAL MONOGRAPHS 129

and by having ulnar and tarsal tubercles(absent in E racemus) and from E thyme-lensis by lacking paravertebral ridges (presentin E thymelensis)

Description of holotypemdashAdult female(QCAZ 14753) with head narrower thanwidest part of body head wider than long(head length 881 head width) snout roundedin dorsal view and angular in lateral viewrelatively short (snout-to-eye distance 161SVL 20 SVL) with small papilla at tip(Fig 3) in lateral view canthus rostralisdistinct (Fig 3) loreal region concave nostrilsslightly protuberant directed laterally inter-orbital area flat broader than upper eyelid(upper eyelid width 719 interorbital dis-tance) cranial crests absent upper eyelidbearing one enlarged nonconical tubercletympanic membrane ill-defined paler thansurrounding skin tympanic annulus distinct

round supratympanic fold obscuring upperand posterodorsal edges of annulus (Fig 3)tympanum diameter 429 eye length oneenlarged nonconical postrictal tubercleChoanae small nearly elliptical not con-cealed by palatal shelf of maxillary dentiger-ous process of the vomer oblique widelyseparated posteromedial to choanae eachbearing 0ndash3 small teeth shortest distancebetween dentigerous processes 324 dis-tance between choanae tongue longer thanwide granular with small notch in posteriorborder

Skin of head shagreen with one smallinterorbital tubercle dorsum shagreen withsmall tubercles in scapular and sacral regionsand posterior to eye dorsolateral folds absentflanks with some large flat warts venterareolate with warty pelvic patch discoidalfold not well-defined anal sheath absentForearm slender radio-ulna length 254SVL white flecks denote three low ulnartubercles ulnar fold absent hand lengthlonger than radio-ulna length (hand length342 SVL) fingers with fleshy lateral fringesthat join at base to form basal web fingerlengths I II IV III palmar tuberclebifid thenar tubercle oval (Fig 4) subarticulartubercles round prominent supernumerary

FIG 3mdashDorsal (A) and lateral (B) views of head ofholotype of Eleutherodactylus huicundo QCAZ 14753SVL 5 284 mm female

FIG 4mdashVentral view of right hand (A) and foot (B) ofholotype of Eleutherodactylus huicundo QCAZ 14753SVL 5 284 mm female


palmar tubercles numerous and conspicuous(Fig 4) disc cover of Finger I slightlyexpanded those of Fingers IIndashIV expanded(Fig 4) outer discs of fingers larger than thoseof toes all disc covers with elliptical ventralpads defined by grooves

Hind limbs relatively slender tibia length500 SVL foot length about same as tibialength white flecks on three low tubercles ontarsus heel with enlarged nonconical tubercletoes with fleshy lateral fringes that join at baseto form basal web (Fig 4) subarticulartubercles round prominent inner metatarsaltubercle oval two to three times size ofsubconical outer tubercle supernumeraryplantar tubercles prominent and numerous(Fig 4) all disc covers expanded toes withdefined pads disc pads nearly elliptical toelengths I II III V IV (Fig 4) tip ofToe V reaching proximal border of distalsubarticular tubercle of Toe IV tip of Toe IIIreaching distal border of medial subarticulartubercle of Toe IV

Color in ethanol of holotype (Figs 2A B3)mdashDorsum and flanks dark gray with darkergray markings narrowly outlined with blackinner and outer white lines Interorbitalcanthal and supratympanic stripes blackanterior border of black interorbital stripeoutlined by narrower white stripe Outer edgeof upper eyelid with white stripe that extendsalong canthus rostralis to tip of snout Arealimited by interorbital and canthal stripes palerthan rest of dorsum Two weakly defined thingrayish white labial bars below eye Limbs withfour transverse dark gray bars outlined bynarrow white lines Groin dark brownish gray

with minute white spots Posterior surfaces ofthighs dark gray with minute cream spotsThroat and venter grayish brown with smalldark brown spots pelvic patch gray Palmar andplantar surfaces dark gray Dorsal and ventralsurfaces of Fingers I and II cream and FingersIII and IV dark gray Dorsal and ventralsurfaces of Toes IndashIII cream and Toes IV andV dark gray

Measurements of holotype (mm)mdashSVL 5261 tibia length 5 142 foot length 5 144head length 5 89 head width 5 101 uppereyelid width 5 23 interorbital distance 5 32eye diameter 5 29 eye-to-nostril distance 525 snout-to-eye distance 5 42 tympanumdiameter 5 12 eye-to-tympanum distance 512 internarial distance 5 21 radio-ulnalength 5 70 hand length 5 94 and Finger-I length 5 58

Variation in external morphologymdashVaria-tion in measurements and proportions is givenin Tables 1 and 2 Other variation is as followsupper eyelid with subconical tubercle (2males) or no tubercle (1 female) tympanicmembrane not evident (1 female) vomerineteeth absent (3 females) or only one toothvisible (1 female 1 male) tongue withoutnotch in posterior border (1 female) dorsumwithout tubercles (1 female) or with some flatwarts (1 male) flanks shagreen without warts(2 males) or with small but pronounced warts(1 male) tubercles on tarsus heel and ulnaenlarged (1 male) supernumerary tubercleson palms and soles less evident (1 female 2males) Usually tubercles on eyelid ulna heeland tarsus are more pronounced in males thanin females

TABLE 1mdashComparison of coloration and morphological characters among species of the Eleutherodactylus orcesi GroupSVL in mm Ventral coloration for specimens preserved in alcohol

SVLmales

SVLfemales

Tympanicannulus

Vocal slitsin males Ventral coloration Source

E huicundo 198ndash213 232ndash287 Visible Present Gray to dark grayor dark brown

This work

E obmutescens 214ndash266 285ndash384 Not visible Absent Cream with reticulationto gray

Lynch 1980

E orcesi 239ndash296 352ndash361 Visible Present Pale brown Lynch 1972 1981E ortizi 181ndash247 243ndash292 Visible Present White with or without

spots andor reticulationThis work

E racemus 252ndash302 299ndash379 Visible Absent White with brown spotsand reticulation

Lynch 1980

E simoteriscus 231ndash251 257ndash314 Visible Absent Pale brown Lynch et al 1996E simoterus 259ndash327 314ndash390 Visible Present Reddish-brown to black Lynch et al 1996E thymelensis 214ndash252 280ndash335 Not visible Present Gray to brown with

black marksLynch 1972 1981


Color variation (in ethanol)mdashBased on fiveadult males and six adult females (paratypes)Labial bars clearly defined (1 male) or absent(1 male) Dorsum from brownish gray tobrownish orange marks on dorsum not out-lined by black and white lines (1 female 1male) dorsum and flanks with pale yellowish-white warts (1 male) Throat and venter fromgrayish cream to dark brownish-gray with orwithout dark gray marks Undersides of limbscream (1 female) or cream with pale yellowishwarts and minute dark gray spots (1 male)Groin and posterior surfaces of thighs brownwith minute cream spots (1 female) or creamwith minute black spots (3 males) Tarsaltubercles not pigmented with white (2 fe-males) Palmar and plantar surfaces cream (1male) Usually females have a darker colora-tion than males

Color in lifemdashColor in life for threeindividuals is as follows

QCAZ 15391 Iris golden brown with fineblack reticulation and dark brown horizontalstreak dorsum greenish brown with darkerbrown markings outlined with narrow blackand pale brown lines white stripe in outeredge of upper eyelid interorbital canthal andsupratympanic stripes dark brown two nar-row white labial bars below eye tympanumpale brown bars in hind and forelimbs weaklydefined groin gray posterior surfaces of thighsdark gray throat and venter cream gray withsmall dark brown spots palmar and planarsurfaces dark gray dorsal and ventral surfacesof Fingers I and II orange yellow and FingersIII and IV brown dorsal and ventral surfacesof Toes IndashIII orange-yellow and Toes IV and Vbrown

QCAZ 15394 (adult male) Same as QCAZ15391 except dorsum uniformly orange-brown flanks light green-gray groin andanterior and posterior surfaces of thighsmustard-yellow throat light yellow-gray ven-ter greenish yellow with light and dark brownmarks

QCAZ 15392 (adult male) Same as QCAZ15391 except middorsal blotch green flanksbrown with small cream spots interorbitalcanthal and supratympanic strip reddishbrown labial bars absent no bars on limbsgroin and posterior surfaces of thighs yellowthroat gray (red when male was calling) ventercream with dark brown warts palmar andplanar surfaces dark orange-brown dorsal andventral surfaces of Fingers I and II orangeand Fingers III and IV orange-brown dorsaland ventral surfaces of Toes IndashIII orange andToes IVndashV orange-brown

Natural historymdashEleutherodactylus hui-cundo occurs in Bosque Siempre VerdeMontano Alto (High Montane EvergreenForest) and Paramo de Frailejones (paramodominated by the plant species Espeletiapycnophylla ssp angelensis) as defined byValencia et al (1999) Individuals were foundat night mostly on terrestrial and epiphyticbromeliads (7 of 11 individuals) branches andleaves 30ndash230 cm (xx 5 1056 6 718 n 5 11)above the ground At the time of collection(August 2000 June 2001) three females(QCAZ 14746 14752ndash53) had mature eggstwo males (QCAZ 15392 15394) were heard

TABLE 2mdashMeasurements of adult males and females ofEleutherodactylus huicundo (mean 6 standard deviation

followed by range in mm)

CharacterMales

(n 5 5)Females(n 5 6)

SVL 206 6 06 257 6 23198ndash213 232ndash287

Tibia length 117 6 06 137 6 06110ndash125 128ndash142

Foot length 116 6 10 135 6 12105ndash128 115ndash146

Head length 76 6 06 94 6 0668ndash82 89ndash104

Head width 83 6 03 98 6 0678ndash87 89ndash107

Upper-eyelid width 21 6 02 24 6 0119ndash23 22ndash26

Interorbital distance 27 6 02 31 6 0225ndash30 29ndash34

Eye length 27 6 02 33 6 0324ndash29 29ndash37

Eye-to-nostril distance 20 6 02 25 6 0117ndash22 22ndash25

Snout-to-eye distance 33 6 03 41 6 0330ndash38 35ndash44

Tympanum diameter 12 6 01 14 6 0211ndash13 12ndash16

Eye-to-tympanum distance 09 6 01 12 6 0207ndash10 09ndash14

Internarial distance 18 6 01 21 6 0216ndash20 20ndash24

Radio-ulna length 59 6 03 68 6 0555ndash62 62ndash74

Hand length 74 6 07 89 6 0667ndash82 80ndash97

Finger-I length 45 6 04 56 6 0440ndash51 49ndash60


calling and three juveniles (QCAZ 14748ndash50)were found

DistributionmdashEleutherodactylus huicundois known only from the montane forest andparamo in the vicinity of El Playon de SanFrancisco (approx 778 379450 W 008 379500 N)at elevations of 3229ndash3700 m ProvinciaSucumbıos Cordillera Oriental of the Andesof northern Ecuador (Fig 11) The proximityof El Playon de San Francisco to theColombian border (5 km) almost assuresthat E huicundo also occurs in Colombia

EtymologymdashIn the quichua language hui-cundo refers to any plant that has the generalform of a bromeliad We use the specific namehuicundo as a noun in apposition that indicatesthe preference of the species to bromeliads

Eleutherodactylus ortizi sp nov

Figs 5ndash10

HolotypemdashQCAZ 16313 an adult femalecollected by Diego Almeida near Nueva

America (778 589540 W 008 159340 N elevation3420 m) Provincia Imbabura Ecuador onJuly 2001

ParatypesmdashSame data as holotype exceptQCAZ 14777 14780 14790 adult femalescollected by Fernando Nogales on 10ndash14 July2000 QCAZ 14792 adult female collected byDiego Almeida on 01 August 2000 QCAZ14793 adult female collected by DiegoAlmeida on 12 July 2000 QCAZ 14778 adultmale collected by Fernando Nogales on 14July 2000 QCAZ 14782ndash4 adult male col-lected by Fernando Nogales on 10ndash14 July2000 QCAZ 14786 adult male collected byDiego Almeida on 19 August 2000 QCAZ14787 adult male collected by FernandoNogales on 2 August 2000 QCAZ 14788 adultmale collected by Diego Almeida on 3 August2000 QCAZ 14789 adult male collected byDiego Almeida on 11 July 2000 QCAZ 14805adult male collected by Fernando Nogales on1 August 2000 QCAZ 16310ndash2 adult malescollected by Fernando Nogales on July 2001

FIG 5mdashEleutherodactylus ortizi (A) QCAZ 14777 SVL 5 243 mm female (B) Non-collected individual (C) QCAZ14789 SVL 5 198 mm male (D) QCAZ 14783 SVL 5 209 mm male


QCAZ 14763 adult male collected by DiegoAlmeida near El Chamizo (778 469030 W 008299350 N elevation 3264 m) Provincia deCarchi Ecuador on 20 July 2001 QCAZ14765ndash6 14769 adult males collected byDiego Almeida and Benjamin McCormicknear El Chamizo on 17ndash19 July 2000 andQCAZ 14770ndash72 adult males collected byDiego Almeida Benjamin McCormick andFernando Nogales near El Chamizo on 6ndash7August 2000

DiagnosismdashEleutherodactylus ortizi has ananteriorly exposed frontoparietal fontanelleAdditionally E ortizi is diagnosed by having(1) skin of dorsum shagreen flanks in femaleswith numerous flat warts in males areolate

venter areolate dorsolateral folds absent (2)tympanic annulus evident round 231ndash464of eye length (xx 5 366 6 54 n 5 26)tympanic membrane poorly-defined (3) eye-to-nostril distance 594ndash815 eye length (xx 5696 6 55 n 5 26) snout rounded in dorsaland lateral views without papilla at tip (Fig 5)(4) upper eyelid narrower than interorbitalspace with or without one small tuberclecranial crests absent (5) vomerine teethabsent or reduced and difficult to see (6)males with vocal slits and median subgularvocal sac nuptial pads absent (7) Finger Ishorter than Finger II fingers with expandeddiscs (Fig 7) (8) fingers with fleshy lateralfringes (Fig 7) (9) ulnar tubercles usuallyabsent if present small (10) heel usually withone small tubercle tarsus without tubercles onouter edge (11) two prominent metatarsaltubercles inner oval two to three times size ofsubconical outer tubercles supernumeraryplantar tubercles usually low and difficult tosee (Fig 7) (12) toes with fleshy lateral fringes(Fig 7) (13) in ethanol dorsum usuallyuniform gray to brown interorbital andcanthal stripes gray to brown white stripe inouter edge of eyelid and canthus rostralislabial bars and bars on limbs usually absentthroat whitish cream venter uniform whitishcream to cream with dark gray reticularpattern groin and posterior surfaces of thighs

FIG 6mdashDorsal (A) and ventral (B) views of holotype ofEleutherodactylus ortizi QCAZ 16313 SVL 5 292 mmfemale

FIG 7mdashVentral view of right hand (A) and foot (B) ofholotype of Eleutherodactylus ortizi QCAZ 16313 SVL 5292 mm female


usually whitish cream palmar and plantarsurfaces whitish cream (Figs 6 7) (14) adultssmall males 181ndash247 mm SVL (xx 5 216 617 n 5 19) females 243ndash292 mm SVL (xx 5267 6 18 n 5 7)

Comparison with similar speciesmdashSpeciesof the Eleutherodactylus orcesi Group differfrom most other Eleutherodactylus by havingan anteriorly exposed frontoparietal fontanelle(Lynch et al 1996) Characters that distin-guish species of the E orcesi Group arepresented in Table 1 Additionally Eleuther-odactylus ortizi differs from E obmutescens byhaving a gray to brown dorsum (dark brown inE obmutescens) from E orcesi by havinga shagreen dorsum (areolate in E orcesi) andfrom E racemus by having a canthal stripe(absent in E racemus) Eleutherodactylusortizi differs from E simoteriscus by havingvocal slits (absent in E simoteriscus Lynch etal 1996) and a mostly cream venter (venterpale brown in E simoteriscus Lynch et al1996) from E simoterus by having relativelylarge discs on fingers (discs relatively small inE simoterus Lynch 1980) and by lackingwarts on dorsum (present in E simoterusLynch 1980) from E thymelensis by lackingparavertebral ridges (present in E thymelen-sis) and from E huicundo sp nov by usuallylacking tarsal tubercles (present in E hui-cundo) Males of E ortizi have areolateflanks (flanks shagreen to warty in E huicundo)and small or non-evident tubercles on eyelidulna and heel (tubercles pronounced in malesof E huicundo)

Description of holotypemdashAdult female(QCAZ 16313) with head narrower than bodyhead wider than long (head length 94 headwidth) snout round in dorsal and lateral viewsrelatively short (snout-to-eye distance 161SVL 20 SVL) without papilla at tipcanthus rostralis distinct loreal region con-cave nostrils slightly protuberant directedlaterally interorbital area flat broader thanupper eyelid (upper eyelid width 861 in-terorbital distance) cranial crests absentupper eyelid bearing one small tubercletympanic membrane not visible anteroventralhalf of tympanum distinct supratympanic foldobscures dorsal and posterodorsal borders oftympanum tympanum diameter 410 eyelength one postrictal tubercle enlarged non-conical Choanae small nearly elliptical not

concealed by palatal shelf of maxillary dentig-erous process of vomer oblique posteromedialto choanae bearing 1ndash3 teeth each shortestdistance between dentigerous processes 363distance between choanae tongue longer thanwide granular posterior border of tongue withsmall notch

Skin of head and dorsum of body shagreendorsolateral folds absent flanks with numer-ous large flat warts venter areolate with wartypelvic patch discoidal fold not well definedanal sheath absent Forearm slender radio-ulna length 260 SVL antebrachial tubercleand tubercle on elbow small ulnar tuberclessmall and barely distinguishable ulnar foldabsent hand longer than radio-ulna length(hand length 364 SVL) fingers with fleshylateral fringes joining at base to form basalweb fingers lengths I II IV III palmartubercle bifid thenar tubercle oval (Fig 7)

FIG 8mdashLateral view of head of males of Eleuther-odactylus huicundo (A) QCAZ 14754 SVL 5 213 mmand of E ortizi (B) QCAZ 16310 SVL 5 237 mm


subarticular tubercles round prominent su-pernumerary palmar tubercles present (Fig7) disc cover on Finger I not well developedthose of Fingers IIndashIV expanded (Fig 7) outerdisc covers of fingers larger than those of toesall disc covers with elliptical ventral disc padsdefined by grooves (Fig 7)

Hind limbs slender tibia length 521 SVLfoot slightly shorter than tibia (foot length961 tibia length) tarsal tubercles absentheel with small tubercle toes with fleshylateral fringes joining at base to form basalweb (Fig 7) subarticular tubercles roundprominent inner metatarsal tubercle ovalabout twice the size of subconical outertubercle supernumerary plantar tuber-cles poorly defined (Fig 7) all disc covers ex-

panded toes with defined pads disc padsnearly elliptical Toe lengths I II III V IV (Fig 7) tip of Toe V reaching proximalborder of distal subarticular tubercle of Toe IVToe III reaching distal border of medial sub-articular tubercle of Toe IV

Color in ethanol of holotype (Fig 6)mdashDorsum uniform gray flanks paler thandorsum Interorbital and canthal stripes faintsupratympanic stripe absent two labial barsbelow eye poorly defined White stripe onouter edge of upper eyelid continuous alongcanthus rostralis to tip of snout Area limitedby interorbital and canthal stripes paler thanrest of dorsum Limbs uniform gray withoutbars Groin anterior and posterior surfaces ofthighs throat venter undersides of limbs and

FIG 9mdashColor variation in adult males of Eleutherodactylus ortizi Dorsal view of (A) QCAZ 14763 SVL 5 236 mm(B) QCAZ 14778 SVL 5 231 mm (C) QCAZ 14782 SVL 5 231 mm and (D) QCAZ 14787 SVL 5 212 mm


palmar and plantar surfaces whitish creamventral surface of discs of Fingers I and II andToes IV and V dark gray (Fig 6)

Measurements of holotype (mm)mdashSVL 5292 tibia length 5 152 foot length 5 146head length 5 110 head width 5 117 uppereyelid width 5 31 interorbital distance 5 36eye diameter 5 39 eye-to-nostril distance 526 snout-to-eye distance 5 47 tympanumdiameter 5 16 eye-to-tympanum distance 509 internarial distance 5 26 radio-ulnalength 5 76 hand length 5 106 andFinger-I length 5 67

Variation in external morphologymdashVaria-tion in morphological measurements andproportions is given in Tables 2 and 3 Based

on 19 males and 7 females other variation is asfollows flanks areolate (all males) or with flatwarts (1 female) small papilla at tip of snout (1male) no tubercle on upper eyelid (2 females10 males) one or two vomerine teeth visible (4males) pronounced notch on posterior borderof tongue (1 female 2 males) or without notch(1 male) no ulnar tubercles (1 female 6males) no antebrachial tubercle and tubercleon elbow (2 females 6 males) no tubercle onheel (1 female 9 males) or three smalltubercles on heel (1 female) small tarsaltubercles (3 males)

Color variation (in ethanol)mdashEleuthero-dactylus ortizi is remarkably variable Someof color patterns that differ from the holotype

FIG 10mdashColor variation in adult males of Eleutherodactylus ortizi Ventral view of (A) QCAZ 14763 SVL 5 236 mm(B) QCAZ 14778 SVL 5 231 mm (C) QCAZ 14782 SVL 5 231 mm and (D) QCAZ 14787 SVL 5 212 mm


(Fig 6) are as follow (1) dorsum flanksventer and groin whitish cream with dark grayreticulation two dark gray labial bars limbsand posterior surfaces of thighs whitish creamwith dark gray bars (Figs 9A 10A) (2) dorsumgray with dark gray marks flanks gray withwhite spots outlined by black limbs gray withsmall dark gray spots groin cream posteriorsurfaces of thighs cream with elongated darkgray marks venter cream with black reticula-tion and white spots outlined with black (Figs9B 10B) (3) dorsum whitish cream flankswhitish cream with black marks in the anteriorhalf limbs cream without bars groin andposterior surfaces of thighs whitish creamventer whitish cream with some dark grayflecks and black spots next to arm insertiondisc pads of Finger III and IV cream (Figs 9C10C) and (4) dorsum pale gray with somesmall black spots on posterior half flanks palegray with few white spots outlined with blacklimbs pale gray without bars groin andposterior surfaces of thighs cream ventercream with numerous dark gray flecks andsome white spots outlined with black (Figs9D 10D)

Detailed variation is noted below Dorsumcream to brown (5 males) with small dark graymarks or spots (4 females 7 males) or darkgray oblique stripes (1 male) Labial barspresent (1 male) Supratympanic stripe brown(1 male 1 female) Flanks cream to grayish

brown with dark gray flecks (1 male) graywarts (1 male 1 female) white spots outlinedwith black (1 female 6 males) dark grayreticular pattern (2 males) or with dark grayoblique bars (2 males) Limbs with faint bars (1female) or clearly defined bars (2 males)Forelimbs and shanks brown with small darkgray spots (1 female 1 male) Concealedsurfaces of limbs and groin cream with smalldark gray marks (3 males) gray with minutecream spots (1 female 1 male) brown withdarker marks (1 male) or dark gray with whitespots (1 male) Posterior surfaces of thighsdark brown with minute cream spots (1female) Undersides of limbs pale brown (1male) Throat grayish cream (1 female) orcream with dark gray reticular pattern (1female) Venter whitish cream to creamy gray(1 male) with midventral pale brown stripe (1female) small gray spots (2 females 4 males)

TABLE 3mdashProportions (in percentages) of adult males andfemales of Eleutherodactylus huicundo

CharactersMales


Head widthSVL 386ndash414 348ndash416Head lengthSVL 343ndash396 345ndash397Head widthHead length 975ndash1206 967ndash1135Eye-to-nostril distanceEye

length 708ndash760 647ndash862Eyelid widthInterorbital

distance 733ndash885 706ndash839Tympanum diameterEye

length 414ndash480 364ndash448Radio-ulna lengthSVL 266ndash297 254ndash288Hand lengthSVL 332ndash387 310ndash387Tibia lengthSVL 556ndash587 481ndash584Hand lengthRadio-ulna

length 1117ndash1367 1203ndash1406Foot lengthTibia length 948ndash1041 878ndash1028Foot lengthSVL 530ndash601 467ndash593Finger-I lengthHand

length 573ndash623 613ndash663

TABLE 4mdashMeasurements of adult males and females ofEleutherodactylus ortizi (mean 6 standard deviation


CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292















Hand width 45 6 04 59 6 0735ndash50 50ndash68


faint gray reticular pattern (1 female 2 males)to defined reticular pattern (3 males) or withwhite spots outlined with black (1 female 5males)

Color in lifemdashColor in life for four individ-uals is as follows

QCAZ 14765 Iris brownish copper dorsumpale brown with gold specks hind andforelimbs brown with pale green bars groindark brown with bright green patches throatgreenish brown venter white with dark brownspecks (field notes of D Almeida on 17 July2000)

QCAZ 14777 Dorsum uniform brownsupraorbital and canthal stripes grayish whiteflanks brown with white spots outlined withblack (from color slide Fig 5A)

Uncollected individual Dorsum uniformbrown flanks dark gray with greenish yellowspots (from color slide Fig 5B)

QCAZ 14789 Dorsum uniformly greenishyellow (from color slide Fig 5C)

QCAZ 14783 Dorsum yellow with darkbrown marks flanks yellow (from color slideFig 5D)

EcologymdashEleutherodactylus ortizi occursin Evergreen High Montane Forest (BosqueSiempre Verde Montano Alto) EspeletiaParamo (Paramo de Frailejones Paramo domi-nated by the plant species Espeletia pycno-phylla ssp angelensis) and HerbaceousParamo (Paramo Herbaceo) as defined byValencia et al (1999) Individuals were ob-

served at night in secondary montane forest(32 of 71 individuals) paramo (24 of 71 indi-viduals) primary montane forest (8 of 71individuals) and agricultural lands (7 of 71individuals Schultz et al 2000) The frogswere in terrestrial bromeliads (39 of 61individuals) other plants (27 of 61 individuals)or on the ground (5 of 61 individuals)

DistributionmdashEleutherodactylus ortizi isknown only from the montane forest and par-amo near El Chamizo (778 469030W 008 299350N 3264 m Provincia Carchi Ecuador) andNueva America (778 589540 W 008 159340 N3420 m Provincia Imbabura Ecuador) Bothlocalities are in the Cordillera Oriental of theAndes of northern Ecuador (Fig 11)

EtymologymdashThe specific name is a noun inthe genitive case and is a patronym forFernando Ortiz-Crespo one of the most notedEcuadorian ornithologists Fernando Ortiz-Crespo was well known for his research inthe Galapagos Islands and the Andes Hedrowned while conducting fieldwork in theLaguna de la Mica in the high Andes ofEcuador on 13 September 2001

TABLE 5mdashProportions (in percentages) of adult males andfemales of Eleutherodactylus ortizi

CharactersMales






length 1161ndash1429 1232ndash1395Foot lengthTibia length 797ndash963 894ndash980Foot lengthSVL 439ndash511 465ndash519Hand widthHand length 577ndash676 588ndash800

FIG 11mdashDistribution of Eleutherodactylus huicundo(triangle) and E ortizi (circles) in Ecuador


KEY TO SPECIES OF THE

ELEUTHERODACTYLUS ORCESI GROUP

1 Tympanic annulus not externally vis-ible under skin - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 2Tympanic annulus externally visibleunder skin - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 3

2 Dorsum usually having paravertebralfolds adult males with vocal slits - - - -

- - - - - - - - - - - - - - - - - Eleutherodactylus thymelensisDorsum lacking paravertebral foldsadult males lacking vocal slits - - - - - - - - - -

- - - - - - - - - - - - - - - - Eleutherodactylus obmutescens3 Skin of dorsum mostly areolate or

with flat warts - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 4Skin of dorsum mostly shagreen - - - - - - - - - - - - 6

4 Skin of dorsum mostly areolate - - - - - - -

- - - - - - - - - - - - - - - - - - - - - - - - - - Eleutherodactylus orcesiSkin of dorsum mostly with flat warts - - - - 5

5 In adult females SVL 314ndash390 mmin adult males SVL 259ndash327 mmadult males with vocal slits - - - - - - - - - - - - - -

- - - - - - - - - - - - - - - - - - - - Eleutherodactylus simoterusIn adult females SVL 257ndash314 mmin adult males SVL 231ndash251 mmadult males without vocal slits - - - - - - - - -

- - - - - - - - - - - - - - - - Eleutherodactylus simoteriscus6 In adult females SVL 299ndash379 mm

in adult males SVL 252ndash302 mmadult males lacking vocal slits sinuousparavertebral folds sometimes pre-sent - - - - - - - - - - - - - - - Eleutherodactylus racemusIn adult females SVL 295 mm inadult males SVL 250 mm adultmales with vocal slits paravertebralfolds absent - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - - 7

7 Ulnar and tarsal tubercles presentbackground of venter gray to darkgray - - - - - - - - - - - - - - Eleutherodactylus huicundoUlnar and tarsal tubercles absent orminute background of venterwhite - - - - - - - - - - - - - - - - - - Eleutherodactylus ortizi

RESUMEN Describimos dos especies nue-vas de Eleutherodactylus e hipotetizamos quepertenen al grupo de especies reconocidocomo E orcesi Ambas especies habitan losAndes del norte del Ecuador a altitudes sobrelos 3000 m La presencia de una fontanelaexpuesta entre los frontoparietales distinguea las nuevas especies de la mayorıa de especiesdel genero Eleutherodactylus en donde lafontanela frontoparietal esta cubierta por los

huesos frontoparietales Adicionalmente lasnuevas especies se caracterizan por poseerdientes vomerinos reducidos o ausentes yconspicuos pliegues laterales en los dedos delas manos y pies Las nuevas especies puedenser diferenciadas entre ellas por la presenciaausencia de determinados tuberculos y por suspatrones de coloracion

AcknowledgmentsmdashWe thank L Trueb W E Duell-man L Fitzgerald H Alamillo E Bonaccorso C SheilO Torres-Carvajal E O Wiley and an anonymousreviewer for critically reviewing this manuscript L Trueband J E Simmons for facilitating access to the KUspecimens L A Coloma for granting loans from QCAZand J D Lynch for comments on the manuscript andchecking the unidentified specimens described in thiswork Research was supported by The University ofKansas the Fundacion para la Conservacion de Ecosiste-mas Amenazados (Numashir) and a fellowship from theFundacion para la Ciencia y Tecnologıa del Ecuador(FUNDACYT) under the sponsorship of the Departa-mento de Ciencias Biologicas of the Pontificia UniversidadCatolica del Ecuador Fieldwork was founded by theOxford University at UK through the Andinoherps project

LITERATURE CITED

GUAYASAMIN J M 2004 A new species of Eleutherodac-tylus (Anura Leptodactylidae) from the northwesternlowlands of Ecuador Herpetologica 60103ndash116

LYNCH J D 1971 Evolutionary relationships osteologyand zoogeography of leptodactylid frogs MiscellaneousPublications Natural History Museum The Universityof Kansas 531ndash238

mdashmdashmdash 1972 Two new species of frogs (Eleutherodacty-lus Leptodactylidae) from the paramos of northernEcuador Herpetologica 28141ndash147

mdashmdashmdash 1980 New species of Eleutherodactylus ofColombia (Amphibia Leptodactylidae) I five newspecies from the paramos of the Cordillera CentralCaldasia 13165ndash188

mdashmdashmdash 1981 Leptodactylid frogs of the genus Eleuther-odactylus in the Andes of northern Ecuador andadjacent Colombia Miscellaneous Publications of theMuseum of Natural History University of Kansas 721ndash46

LYNCH J D AND W E DUELLMAN 1997 Frogs of thegenus Eleutherodactylus (Anura Leptodactylidae) inwestern Ecuador systematics ecology and biogeogra-phy The University of Kansas Natural History MuseumSpecial Publication 231ndash236

LYNCH J D P M RUIZ-CARRANZA AND M C ARDILA-ROBAYO 1996 Three new species of Eleutherodactylus(Amphibia Leptodactylidae) from high elevations of theCordillera Central of Colombia Caldasia 18329ndash342

mdashmdashmdash 1997 Biogeographic patterns of Colombian frogsand toads Revista de la Academia Colombiana deCiencias Exactas Fısicas y Naturales 21237ndash248

SAVAGE J M 1987 Systematics and distribution of theMexican and Central American rainfrogs of theEleutherodactylus gollmeri group (Amphibia Leptodac-tylidae) Fieldiana Zoology New Series 331ndash57


mdashmdashmdash 1997 A new species of rainfrog of the Eleuther-odactylus diastema group from the Alta Talamancaregion of Costa Rica Amphibia-Reptilia 18241ndash247

SIMMONS J E 2002 Herpetological collecting andcollections management Society for the Study ofAmphibians and Reptiles Herpetological Circular161ndash70

SCHULTZ N H FORSEY B MCCORMICK J MATHER-HILLON AND L FROLICH 2000 A Herpetological Surveyof the High Andes of Northern Ecuador Report of theOxford University Expedition

VALENCIA R C CERON W PALACIOS AND R SIERRA1999 Las formaciones naturales de la Sierra delEcuador Pp 79ndash108 In R Sierra (Ed) Propuesta

Preliminar de Clasificacion de Vegetacion para elEcuador Continental Proyecto INEFANGEF-BIRFand EcoCiencia Quito Ecuador

APPENDIX I

Specimens Examined

Eleutherodactylus obmutescens (KU 144086ndash89144091ndash106) Eleutherodactylus orcesi (KU 130314ndash16177815ndash16 140001 218021ndash23) Eleutherodactylus race-mus (KU 168941ndash68) Eleutherodactylus thymelensis (KU117719ndash21 117724ndash70)


Herpetological Monographs 18 2004 127ndash141 2004 by The Herpetologistsrsquo League Inc

TWO NEW SPECIES OF FROGS (LEPTODACTYLIDAEELEUTHERODACTYLUS) FROM THE HIGH ANDES OF

NORTHERN ECUADOR

JUAN M GUAYASAMIN14 DIEGO ALMEIDA-REINOSO

2 AND FERNANDO NOGALES-SORNOSA3

1Natural History Museum amp Biodiversity Research Center Department of Ecology and Evolutionary BiologyThe University of Kansas Lawrence KS 66045-7561 USA

2Universidad Central del Ecuador Facultad de Filosofıa Letras y Ciencias de la EducacionEscuela de Biologıa y Quımica Av America Quito Ecuador

3Fundacion Ecologica Arcoiris Segundo Cueva Celi 03-15 y Clodoveo CarrionCiudadela Zamora Loja Ecuador

ABSTRACT We describe two new species of Eleutherodactylus that are hypothesized to belong to the Eorcesi Group Both species are found in Andean habitats of northern Ecuador at elevations above 3000 m Thepresence of an anteriorly exposed frontoparietal fontanelle distinguishes the two new species from most othermembers of the genus in which the frontoparietal fontanelle is covered by frontoparietal bones Additionallyboth species have fingers and toes with fleshy lateral fringes and vomerine teeth reduced or absent The newspecies differ from one another mostly by tuberculation and coloration patterns

Key words Andes Ecuador Eleutherodactylus orcesi Leptodactylidae New species

THE ELEUTHERODACTYLUS ORCESI GROUP wasrecognized by Lynch (1981) for an assemblageof species distributed in the high Andes(gt3000 m) of Colombia and Ecuador Theonly diagnostic feature cited for the group wasthe presence of an anteriorly exposed fronto-parietal fontanelle (Lynch et al 1996) Cur-rently the Eleutherodactylus orcesi Groupcontains six species E obmutescens Lynch1980 E orcesi Lynch 1972 E racemus Lynch1980 E simoteriscus Lynch et al 1996 Esimoterus Lynch 1980 and E thymelensisLynch 1972 All of these species live inparamos at elevations of 3000ndash4150 m andreplace one another along the geographic axisof the Andes of Colombia and Ecuador(Lynch 1981 Lynch et al 1997) If the groupis monophyletic its pattern of distribution iscongruent with an allopatric mode of specia-tion (Lynch et al 1997)

It is not surprising that fieldwork in poorlycollected Andean localities results in thediscovery of undescribed species Herpetolog-ical expeditions by Oxford University of UK incooperation with the Pontificia UniversidadCatolica del Ecuador and the UniversidadCentral del Ecuador to localities in montaneforests and paramo in the Andes of northernEcuador have revealed two more species of the

Eleutherodactylus orcesi Group Herein wedescribe these newly discovered species

MATERIALS AND METHODS

Specimens were euthanized in the field byimmersion in chloretone fixed in 10 forma-lin and preserved in 70 ethanol as sug-gested by Simmons (2002) We examinedalcohol-preserved specimens of frogs fromthe herpetological collections at Museo deZoologıa of the Universidad Catolica delEcuador (QCAZ) Natural History Museumof The University of Kansas (KU) andInstituto de Ciencias Naturales of the Uni-versidad Nacional de Colombia (ICN) Inaddition to the type series of the two newspecies specimens examined are listed inAppendix I Morphological measurementswere taken as described in Guayasamin(2004) and are (1) snoutndashvent length (SVL)(2) tibia length (3) foot length (4) head length(5) head width (6) interorbital distance (7)upper eyelid width (8) internarial distance (9)eye-to-nostril distance (10) snout-to-eye dis-tance (11) eye diameter (12) tympanumdiameter (13) eye-to-tympanum distance(14) radioulna length (15) hand length (16)Finger-I length Sexual maturity was deter-mined by the presence of eggs or convolutedoviducts in females and by the presence ofvocal slits in males For ease of comparison4 CORRESPONDENCE e-mail juanmkuedu

127


SYSTEMATICS


Figs 1ndash4 8


























SVLmales

SVLfemales

Tympanicannulus



This work


Lynch 1980




Lynch 1980












CharacterMales


SVL 206 6 06 257 6 23198ndash213 232ndash287





















Figs 5ndash10




































CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined




SYSTEMATICS


Figs 1ndash4 8


























SVLmales

SVLfemales

Tympanicannulus



This work


Lynch 1980




Lynch 1980












CharacterMales


SVL 206 6 06 257 6 23198ndash213 232ndash287





















Figs 5ndash10




































CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined






















SVLmales

SVLfemales

Tympanicannulus



This work


Lynch 1980




Lynch 1980












CharacterMales


SVL 206 6 06 257 6 23198ndash213 232ndash287





















Figs 5ndash10




































CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined

















SVLmales

SVLfemales

Tympanicannulus



This work


Lynch 1980




Lynch 1980












CharacterMales


SVL 206 6 06 257 6 23198ndash213 232ndash287





















Figs 5ndash10




































CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined










SVLmales

SVLfemales

Tympanicannulus



This work


Lynch 1980




Lynch 1980












CharacterMales


SVL 206 6 06 257 6 23198ndash213 232ndash287





















Figs 5ndash10




































CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined











CharacterMales


SVL 206 6 06 257 6 23198ndash213 232ndash287





















Figs 5ndash10




































CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined







Figs 5ndash10




































CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined

































CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined



























CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined




















CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined














CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined







CharactersMales










CharacterMales


SVL 216 6 17 267 6 18181ndash247 243ndash292





























CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined















CharactersMales


























LITERATURE CITED
















APPENDIX I

Specimens Examined




















LITERATURE CITED
















APPENDIX I

Specimens Examined








APPENDIX I

Specimens Examined



Documents

NORTHERN ECUADOR URL: ...multimedia20stg.blob.core.windows.net/...alcohol-preserved specimens of frogs from the herpetological collections at Museo de Zoologı´a of the Universidad