TWO NEW SPECIES OF ATELOPUS (ANURA: BUFONIDAE)...(from Laguna La Cocha, Departamento Narin˜o,...

229

Herpetologica, 58(2), 2002, 229–252q 2002 by The Herpetologists’ League, Inc.

TWO NEW SPECIES OF ATELOPUS (ANURA: BUFONIDAE)FROM ECUADOR

LUIS A. COLOMAMuseo de Zoologı́a, Centro de Biodiversidad y Ambiente, Departamento de Ciencias Biológicas, Pontificia

Universidad Católica del Ecuador, Av. 12 de Octubre y Roca, Aptdo. 17-01-2184, Quito, Ecuador

ABSTRACT: Two new species of Atelopus from Ecuador are described; both are from the high-lands of the Cordillera Occidental of the Andes, one from the Chimbo Basin and the other fromthe Páramos del Cajas. I include osteological data of both species and the description of the gas-tromyzophorous tadpoles of one of the new species. Patterns of diversity and endemism of Atelopusjustify the necessity of further research and conservation efforts directed toward Andean habitats.

Key words: Anura; Bufonidae; Atelopus; A. guanujo new species; Atelopus nanay new species;Taxonomy; Osteology; Tadpole; Andes; Ecuador

THE speciose neotropical bufonid genusAtelopus contains about 70 recognizedspecies (Frost, 2000). Individuals of Ate-lopus are easily collected because they arediurnal, slow-moving, sometimes abun-dant, and often conspicuously colored.Therefore, many specimens have been de-posited at scientific collections since the19th century. Nevertheless, ongoing explo-rations in the neotropics and more de-tailed analyses of museum specimens con-tinue to reveal previously unknown spe-cies.

Most species of Atelopus occur in thenorthern Andes (Lötters, 1996), and sevennominal species have been recorded fromthe Cordillera Occidental of Ecuador: Ate-lopus arthuri Peters, 1973; A. coynei Mi-yata, 1980; A. exiguus (Boettger, 1892); A.ignescens (Cornalia, 1849); A. longirostrisCope, 1868; A. lynchi Cannatella, 1981;and A. mindoensis Peters, 1973. All ofthese species are endemic to the Cordil-lera Occidental except A. ignescens, whichalso occurs in the Cordillera Oriental (Col-oma et al., 2000).

Herein, I describe two new species fromthe Cordillera Occidental of the Ecuador-ian Andes; one is from Chimbo Valley andthe other is from Páramos del Cajas.

MATERIALS AND METHODS

Specimens examined, other than typespecimens, are listed in Appendix I. Mu-seum acronyms follow Leviton et al.(1985), Frost (1985), and Duellman

(1993). Sexual maturity was determined bythe presence of eggs or convoluted ovi-ducts in females, and by the presence ofnuptial excrescences covering the dorsumof Finger I in males. Throughout the text,I use the term ‘‘spiculae’’ to refer to pus-tular warts and ‘‘coni’’ to refer to spiculaewith pointed projections (Duellman andTrueb, 1986). Measurements were takenfollowing Gray and Cannatella (1985),with the addition of sacrum width (SW 5the length from the outer margin of thesacral diapophysis to the outer margin ofthe contralateral side). Measurements aregiven only for adults. Abbreviations ofmeasurements follow Coloma et al. (2000);they are SVL (snout–vent length), TIBL(tibia length), FOOT (foot length), HLSQ(head length from the squamosal tosnout), IOD (interorbital distance),HDWD (head width), EYDM (eye diam-eter), EYNO (eye to nostril distance),ITNA (internarial distance), RDUL(length of flexed forearm), HAND (handlength), THBL (thumb length), and SW(sacrum width). Measurements of frogsand tadpoles were made to the nearest 0.1mm with MAX-CALt digital calipers.Measurements of tadpoles were made un-der a dissecting microscope. When com-paring SVL of Atelopus, raw data were logtransformed and a Student’s unpaired t-test was used to assess the significance ofdifferences between means. Measure-ments (mean, sample size, and standarddeviation) for A. sorianoi (females and

230 [Vol. 58, No. 2HERPETOLOGICA

males) and A. chrysocorallus (females)were taken from La Marca (1996, 1983),respectively. Data on skeletons were takenfrom cleared and alizarin-red–alcian-bluestained specimens, and they were pre-pared following a modified version of thetechnique of Dingerkus and Uhler (1977).Osteological nomenclature follows Duell-man and Trueb (1986), Trueb and Cloutier(1991), and Trueb (1993), but nomencla-ture for the skeletal elements of the handfollows Fabrezi and Alberch (1996).

Webbing formulae are indicated in themanner described by Savage and Heyer(1969), with modifications suggested byMyers and Duellman (1982) and Savageand Heyer (1997). The webbing formulais a subjective approximation, because insome cases the distinction between web-bing and lateral fringes is ambiguous; insuch cases, a range is indicated when a sin-gle specimen is described. In the diagno-ses, a range also indicates intraspecific var-iation. Staging of larvae follows Gosner(1960). Drawings were made using a cam-era lucida attached to a Wildt M3B mi-croscope. Ecological distributions of eachspecies follow the Natural Formations ofEcuadorian Sierra defined by Valencia etal. (1999). Climatological data for the nat-ural formations were taken from Cañadas-Cruz (1983).

SYSTEMATICS

Atelopus guanujo sp. nov.

Holotype.—QCAZ 11103, adult female,from Guanujo (018 339 200 S, 798 009 240W; 2900 m), Provincia Bolı́var, Ecuador,obtained on 13 April 1983 between 0945h and 1030 h, by Marinus S. Hoogmoedand Luis A. Coloma.

Paratypes.—MHNG 2271.11–38 fromGuanujo, 2600 m, obtained in December1985 by Luis A. Coloma; QCAZ 1884,1885 (cleared-and-stained preparation),1905, obtained at the same locality in De-cember 1986 by Luis A. Coloma; QCAZ1886–92, 11205 (cleared-and-stainedpreparation), 11250–1 (cleared-and-stained preparation), 11253 (cleared-and-stained preparation), KU 289426 (cleared-and-stained preparation), obtained at the

same locality in 10 April 1988 by MarioTapia and Giovanni Onore; KU 193267from near Guaranda, Provincia Bolı́var,Ecuador, obtained by unknown collectorin June 1980; KU 178388–89 from Guanu-jo, 2923 m, obtained in 15 January 1978by John D. Lynch; MHNG 1151.21 fromGallo Rumi, Provincia Bolı́var, Ecuador,obtained in April 1964 by Karl Von Snei-dern.

Diagnosis.—(1) A moderate-sized spe-cies (SVL in females 35.0–46.0, x̄ 5 39.6,n 5 12; males 29.9–37.1, x̄ 5 33.9, n 529); (2) hindlimbs short (tibia length/SVL5 0.322–0.409, n 5 41); (3) phalangealformula of hand 2-2-3-3, basal webbingabsent; (4) foot webbing formula I(0–1)—(0–2)II(½–2)—(1–3)III(1–3)—(2–4)IV(2–4)—(1–3)V; (5) snout acuminate, protrud-ing beyond lower jaw; (6) tympanic mem-brane, tympanic annulus and stapes ab-sent; (7) dorsal surfaces of body usuallysmooth, eight (out of 38) individuals beara few scattered small warts and spiculae,(8) white spiculae (cream in preservative)present on forearm, flanks and dorsal sur-faces of thighs; (9) vertebral neural pro-cesses inconspicuous; (10) dorsum orangeto dark brown (cream to dark brown inpreservative); (11) venter uniform orange(cream to pale yellow in preservative); (12)gular region without warts, spiculae orconi.

In having a plain orange-red dorsum,Atelopus guanujo is most similar to A.bomolochos (populations from Cutchil andChilpe, Provincia Azuay, Ecuador), A. sp.(from Laguna La Cocha, DepartamentoNariño, Colombia; see Coloma et. al.,2000, for a discussion of the taxonomic sta-tus of this population therein included un-der the name A. ignescens), A. carboner-ensis, A. chrysocorallus, and A. sorianoifrom the Andes of Venezuela. It differsfrom A. sp. and A. bomolochos by havingwhite spiculae (in life) on forearm, flanks,and dorsal surfaces of thighs (black in A.sp., yellow in A. bomolochos). Atelopus gu-anujo is significantly smaller than A. car-bonerensis, A. chrysocorallus, A. sorianoi,A. sp. and A. bomolochos (Table 2). Fur-ther, A. guanujo differs from A. sp. bylacking a patch of spiculae and coni on the

June 2002] 231HERPETOLOGICA

FIG. 1.—Holotype of Atelopus guanujo (QCAZ11103): (A) dorsal and (B) ventral views. Female SVL5 37.5 mm.

gular and pectoral regions, from A. bom-olochos in lacking green or greenish or-ange dorsal coloration, from A. carboner-ensis and A. sorianoi in having a less pro-tuberant snout and lacking postocularcrest. Furthermore, it differs from A. car-bonerensis and A. chrysocorallus by havingvocal slits in males and lacking a row ofwarts on the dorsolateral surfaces of thebody in males.

Description of holotype.—(Figs. 1, 2A).Head about as long as wide, HLSQ andHDWD less than one third SVL (HLSQ/SVL 5 0.293, HDWD/SVL 5 0.283);snout acuminate in dorsal view; in lateralview, profile of tip of snout to the anteriormargin of jaw curved and slightly protu-berant; swollen gland on tip of snout ab-sent; nostrils slightly protuberant, directedlaterally, situated at level of apex of lowerjaw; canthus rostralis distinct, weakly con-cave from eye to nostril; loreal region con-cave; lips not flared; interorbital regionand occiput flat, smooth; eyelid flaredwithout distinct tubercles; postorbital crestslightly raised, glandular; pretympanic andtympanic areas covered with warts; tym-panic membrane and tympanic annulusabsent; temporal area with spiculae; cho-anae small, rounded, widely separated;tongue twice as long as wide, broadest an-teriorly, free for half its length posteriorly;ostia pharyngea absent.

Forearm relatively short (RDUL/SVL 50.290); palmar tubercle round; supernu-merary palmar tubercles distinct; thenarand subarticular tubercles distinct; digitaltips with round pads; thumb relatively long(THBL/HAND 5 0.621), apparently hav-ing two phalanges; webbing on hands ab-sent, fingers lacking lateral fringes; relativelength of fingers III . IV . II . I. Tibiarelatively short (TIBL/SVL 5 0.376); foldon distal half of inner edge of tarsus ab-sent; inner metatarsal tubercle oval; outermetatarsal tubercle round, raised, abouttwo thirds length of inner metatarsal tu-bercle; supernumerary plantar and subar-ticular tubercles conspicuous; digital padsdistinct; webbing formula of foot I1—1II1—(1–2)III(1–2)—2IV(2–3)—(1–1½)V; relative length of toes IV . V 5 III. II . I.


FIG. 2.—Lateral view of head of holotype of (A) Atelopus guanujo and (B) A. nanay (QCAZ 1922). Scale5 5 mm.

Dorsal surfaces smooth with scatteredspiculae, increasing in number toward theflanks; flanks and anterior and proximalupper surfaces of forelimbs bearing nu-merous, evenly distributed spiculae;throat, chest, belly, undersides of hin-dlimbs rugose, but free of warts; cloacaopens as an inconspicuous tube at midlev-el of thighs, directed posteriorly; wartspresent lateral to cloacal opening, low.

Coloration in preservative (70% etha-nol): dorsal surface of head and bodybrown, becoming paler brown posteriorlyon body, flanks, dorsal surfaces of limbs;minute gray stippling regularly distributedon dorsum of body (viewed at 83 magni-fication); spiculae cream; dorsal surfaces ofFingers I and II cream; tips of digits ofhand cream dorsally, brown adjacent artic-ular region of phalanges; throat, chest, bel-ly, and ventral surfaces of limbs yellowishcream; outer metatarsal tubercle cream,brown surrounding plantar areas; proximalend of tongue lacking black pigmentation.

Measurements (in mm): SVL 37.5,TIBL 14.1, FOOT 16.9, HLSQ 11.0,HDWD 10.6, ITNR 3.4, EYDM 3.0,EYNO 2.7, RDUL 10.9, HAND 10.3,THBL 6.4, SW 11.0.

Variation.—Meristic variation is de-scribed in Table 1. The paratypes resemblethe holotype with the following notewor-

thy exceptions. The proximal undersides ofthighs and pelvic area have a patch of are-olate skin in all but holotype. Sexual di-morphism is evident in that females arelarger than males (Table 1). Males have vo-cal slits and keratinized nuptial pads onthe dorsal and inner surfaces of the thumband inner side of the proximal phalanx ofFinger II. Forelimbs are long and slenderin females, but short and robust in males(Table 1). Morphological features of handand foot of a female paratype (KU 178389)are depicted in Fig. 3. MHNG 2271.28and 2271.30 have abnormal right and lefthindlimbs, respectively; they lack shanksand distal elements.

All paratypes possess a brown, narrowmiddorsal line; in most specimens, it isconspicuous, but in 7 (QCAZ 1886–87,1905, MHNG 2271.13, 2271.16, 2271. 25,2271.28) of 38 it is indistinct. Dorsal col-oration varies in preservative from uniformyellowish cream to dark brown. Some in-dividuals have one to three small darkbrown marks on dorsum. One specimen(MHNG 2271.16) possesses a creamy yel-low dorsum with ill-defined brown marks.Two specimens (QCAZ 1884, 1886) havedark brown areas on the flanks. The ventervaries from uniform pale cream to paleyellow in preservative. A young female(KU 193267) possesses a dark brown patch


TABLE 1.—Measurements (in mm) of adults of Atelopus guanujo and A. nanay; n 5 sample size. Mean andone standard deviation are listed above range.

A. guanujo

Males(n 5 29)

Females(n 5 12)

A. nanay

Males(n 5 17)

Females(n 5 2)

SVL 33.9 6 1.6(29.9–37.1)

39.6 6 3.0(35.0–46.0)

25.2 6 1.0(23.6–27.3)

35.7(31.7–39.6)

TIBL 12.8 6 0.9(10.1–14.4)

14.2 6 0.3(13.8–14.8)

9.2 6 0.6(7.9–10.1)

11.6(9.9–13.3)

FOOT 14.9 6 1.0(12.5–16.8)

(n 5 28)

16.5 6 0.9(15.3–18.0)

9.4 6 0.7(8.5–10.8)

13.1(10.7–15.4)

HLSQ 10 6 0.4(9.0–11.3)

11.3 6 0.5(10.5–12.1)

7.8 6 0.5(7.0–8.9)(n 5 16)

10.2(8.9–11.4)

HDWD 10.1 6 0.5(8.8–11.1)

10.7 6 0.5(9.7–11.3)

7.8 6 0.3(7.5–8.6)

9.7(8.6–10.7)

EYDM 3.4 6 0.3(2.7–4.1)

3.5 6 0.3(3.0–4.1)(n 5 11)

2.9 6 0.2(2.6–3.2)(n 5 16)

3.3(3.1–3.4)

EYNO 2.3 6 0.2(1.9–2.7)

2.7 6 0.2(2.5–3.0)(n 5 11)

2.0–0.2(1.7–2.2)(n 5 16)

2.4(2.1–2.6)

ITNA 3.6 6 0.2(3.2–3.9)(n 5 28)

3.8 6 0.6(3.3–4.2)

2.9 6 0.3(2.6–3.3)(n 5 16)

3.6(3.3–3.9)

RDUL 9.6 6 0.6(8.7–10.8)(n 5 16)

11.1 6 0.7(9.9–11.7)

7.8 6 0.5(7.1–8.5)(n 5 16)

9.5(8.6–10.4)

HAND 8.5 6 0.5(7.6–9.4)(n 5 27)

10.5 6 0.6(9.6–11.3)

6.3 6 0.3(5.8–7.0)(n 5 16)

8.5(6.9–10.1)

THBL 5.7 6 0.4(4.6–6.4)

6.7 6 0.5(6.1–7.5)

4.0 6 0.2(3.6–4.2)(n 5 16)

5.5(4.4–6.6)

SW 9.4 6 0.6(7.7–10.2)(n 5 28)

11.7 6 1.4(9.1–14.0)

6.4 6 0.4(5.8–7.1)(n 5 16)

9.4(8.2–10.5)

on the undersides of the thighs and pelvicregion.

Color in life (Fig. 4): Color transparen-cies of specimens (no museum numbersassociated) from Guanujo reveal that thedorsum, dorsal surfaces of limbs (exceptfingers and toes) and flanks are nearly uni-form reddish orange to uniform orange orbrownish orange. Dorsal surfaces of fin-gers and toes are yellowish orange withbrown areas on the articular regions ofphalanges. Creamy white spiculae are pre-sent on the flanks and forelimbs. The irisis black and lacks a conspicuous pupilaryring.

Tadpoles.—Tadpoles of this species areunknown.

Distribution and natural history.—Ate-lopus guanujo is known only from the typelocality and its vicinity (Guaranda, GalloRumi) in the Chimbo Basin of the Cor-dillera Occidental of Ecuador, ProvinciaBolı́var (Fig. 5). These localities are be-tween 2600 and 2923 m in humid cloudforest in the upper Rı́o Chimbo Valley. An-nual mean rainfall is 1000–2000 mm, andthe annual mean temperature is 12–18 C.Frogs were collected at disturbed CloudMontane Forest areas near the Rı́o Salinasand Rı́o Illangama. Formerly, this species


FIG. 3.—Ventral view of (A) right hand and (B) right foot of paratype (KU 178389, female) of Atelopusguanujo. Scale 5 2 mm.

FIG. 4.—Atelopus guanujo (no number associated)from Guanujo.

was abundant (personal observation); how-ever, despite repeated efforts to locate thisfrog, the last living individuals were seenin 10 April 1988 by Mario Tapia. Some of

these individuals were in amplexus (MarioTapia, personal communication).

Etymology.—The specific name is anoun in apposition and is dedicated to theGuanujo, a Quechua people inhabitingpart of the region where this species for-merly was abundant. The specific namealso refers to the type locality, near thetown that bears the same name as the in-digenous group. This species is known lo-cally as ‘‘puca sapo,’’ Quechua and Spanishwords, respectively; meaning ‘‘red toad’’.

Atelopus nanay sp. nov.

Holotype.—QCAZ 1922, adult male,from Las Tres Cruces (4 km W of LagunaToreadora; 28 469 S, 798 149 W; Parque Na-cional del Cajas; 4000 m), Provincia Azuay,Ecuador, obtained on 28 July 1989 by


FIG. 5.—Map of Ecuador showing localities of Ate-lopus guanujo (triangles) and A. nanay (circle).

Stella de la Torre, Felipe Campos-Yánez,and Luis A. Coloma.

Paratypes.—QCAZ 1916–21, 1923(cleared-and-stained skeletal preparation),1924–27, KU 217474–78, same data as ho-lotype; CM 117018 from 35 km WNW ofCuenca, Provincia Azuay, Ecuador, col-lected 9 November 1987 by an unknowncollector; QCAZ 13938 from road Cuenca-Molleturo, Provincia Azuay, Ecuador, col-lected 31 December 1981 by FernandoOrtiz-Crespo.

Diagnosis.—(1) A small Atelopus (SVLin females 31.7–39.6, x̄ 5 35.7, n 5 2;males 23.6–27.3, x̄ 5 25.2, n 5 17); (2)hindlimbs short (tibia length/SVL in fe-males 0.312–0.336, n 5 2; males 5 0.331–0.402, n 5 17); (3) phalangeal formula ofhand 1-2-3-3, webbing absent; (4) footwebbing formula I(0–½)—(½–1)II(½–1)—(1–2)III(1–2)—(2–3)IV(2–3)—1V; (5)snout acuminate, protruding beyond lowerjaw; (6) tympanic membrane, tympanic an-nulus and stapes absent; (7) dorsal surfacesusually smooth, in some cases bearing fewscattered, flat warts; (8) scattered whitespiculae (cream in preservative) in flanks,limbs, and dorsum; (9) vertebral neuralprocesses absent; (10) dorsum black(cream to dark brown in preservative); (11)venter cream with green and brown marks

in life; plantar and palmar tuberclescream; (12) gular region without warts,spiculae or coni.

In having a black dorsum, Atelopus na-nay resembles A. ignescens (as redefinedby Coloma et al., 2000) from the northernEcuadorian Andes, and has a size similarto that of A. exiguus from the southernEcuadorian Andes in Provincia del Azuay.It differs from these taxa by having a 1-2-3-3 phalangeal formula (as observed inQCAZ 1923; skeletal preparation) insteadof 2-2-3-3 (A. ignescens, n 5 4; A. exiguus,n 5 4; data taken from Coloma, 1997, andColoma et al., 2000, respectively). Atelo-pus nanay is significantly smaller than A.ignescens (Table 2). Also, A. nanay lacksconi on flanks, and lacks a triangular markwith black spiculae and coni in the gular-pectoral region (present in A. ignescens).Atelopus exiguus has a green dorsum, yel-low spiculae, and a yellow venter in life,whereas A. nanay has a black dorsum,white spiculae, and a venter black tocream with green and brown marks. Fur-thermore, A. nanay differs from A. exiguusby lacking pigmentation of proximal end oftongue and by having minute dark spotsregularly distributed on dorsum of (viewedat 83 magnification).

Description of holotype.—(Figs. 2B, 6,7). Head about as long as wide, HLSQ andHDWD slightly less than one third SVL(HLSQ/SVL 5 0.328, HDWD/SVL 50.316); snout truncate in dorsal view; inlateral view, profile of tip of snout to an-terior margin of jaw curved and slightlyprotruding beyond lower jaw; swollengland on tip of snout absent; nostrils slight-ly protuberant, directed laterally, situatedat level of apex of lower jaw; canthus ros-tralis distinct, weakly concave from eye tonostril; loreal region concave; lips notflared; interorbital region and occiput flat,smooth; eyelid and canthus rostralis flaredwithout distinct tubercles; postorbital crestflared; tympanic region without warts;tympanic membrane and tympanic annu-lus absent; warts in temporal area present,barely raised; choanae small, rounded,widely separated; tongue twice as long aswide, broadest anteriorly, free for half its


TABLE 2.—Results of Student’s unpaired t-test showing significant differences between SVL of Atelopus guan-ugo and each of five species of Atelopus (above) and SVL of A. nanay and A. ignescens (below). Range, mean,

standard deviation (SD), and samples size (n) are indicated above results.

Species Females Males

A. bomolochos 38.3–50.4x̄ 5 44.7, SD 6 3.2, n 5 13t 5 4.078, df 5 23, P , 0.001

32.1–42.1x̄ 5 36.9, SD 6 3.8, n 5 5t 5 2.960, df 5 32, P 5 0.0057

A. carbonerensis 45.6–54.1x̄ 5 49.4, SD 6 4.3, n 5 4t 5 4.981, df 5 14, P , 0.001

—

A. chrysocorallus 47.1–50.4x̄ 5 48.9, SD 6 1.2, n 5 8t 5 6.062, df 5 18, P , 0.001

—

A. sorianoi 42.2–49.1x̄ 5 44.9, SD 6 2.6, n 5 5t 5 6.415, df 5 15, P , 0.001

38.2–39.8x̄ 5 38.9, SD 6 0.8, n 5 5t 5 6.779, df 5 32, P , 0.001

A. sp. 40.3–52.2x̄ 5 45.2, SD 6 3.6, n 5 10t 5 4.090, df 5 20, P , 0.001

36.7–41.0x̄ 5 38.1, SD 6 1.6, n 5 6t 5 5.664, df 5 33, P , 0.001

A. ignescens 35.6–48.2x̄ 5 42.5, SD 6 3.3, n 5 15t 5 3.552, df 5 15, P , 0.0029

34.2–41.4x̄ 5 37.8, SD 6 1.9, n 5 15t 5 24.911, df 5 30, P , 0.001

length posteriorly; vocal slits present; ostiapharyngea absent.

Forearm length nearly one third SVL(RDUL/SVL 5 0.328); fingers notwebbed, lacking lateral fringes; thenar,palmar, and subarticular tubercles distinct;palmar tubercle round; supernumerarypalmar tubercles absent; digit tips withround pads; keratinized nuptial pad cov-ering entire dorsal and lateral surfaces ofthumb and inner side of Finger II; fore-arm swollen; thumb relatively short(THBL/HAND 5 0.641), apparently withone phalanx; relative length of fingers III. IV . II . I. Tibia relatively short(TIBL/SVL 5 0.402); no fold on distal halfof inner edge of tarsus; inner metatarsaltubercle oval; outer metatarsal tubercleround, noticeably raised, slightly smallerthan inner metatarsal tubercle; supernu-merary plantar tubercles absent; subarti-cular tubercles at base of digits barelyraised; digital pads distinct, round; web-bing covering all digits except two or threephalanges of Toe IV; foot webbing formulaI0—½II½—1III1—(2–3)IV2—1V; rela-tive length of toes IV . V 5 III . II . I.

Dorsal surfaces smooth with few wrin-kles on head and limbs; flanks with few,scattered, barely-raised spiculae, most nu-

merous ventrolaterally; throat, chest, belly,and undersides of forearm rugose, but freeof warts; cloacal patch with few wrinkles,but not differentiated from adjacent skinon undersides of thighs; cloacal opening atmidlevel of thighs with surrounding skinheavily wrinkled.

Color in life: dorsum black; spiculae onflanks white; palmar and plantar tuberclescream; venter cream with large greenmarks; gular region with brown marks(Field Notes of L. A. Coloma, 28 July1989).

Coloration in preservative (70% etha-nol): dorsum, flanks and dorsal surfaces oflimbs black; minute gray stippling regular-ly distributed on dorsum of body (viewedat 83 magnification); spiculae cream; up-per lip cream suffused with brown; well-defined black stripe bordering the lowerlabium at inner and outer margins; under-sides of limbs black with cream flecks;throat cream with brown marks; chest andabdomen cream; palmar, thenar, metatar-sal, and subarticular tubercles on palmarand plantar surfaces cream contrastingwith black on surrounding surfaces; ablack, elliptical cloacal patch on under-sides of thighs; proximal end of tonguelacking black pigmentation. The iris is


FIG. 6.—Holotype of Atelopus nanay: (A) dorsaland (B) ventral views. Male SVL 5 24.4 mm.

black and lacks a conspicuous pupilaryring.

Measurements (in mm): SVL 24.4,TIBL 9.8, FOOT 8.0, HLSQ 8.0, HDWD7.7, ITNR 2.7, EYDM 2.6, EYNO 2.1,RDUL 8.0, HAND 6.4, THBL 4.1, SW6.9.

Variation.—Meristic variation is de-scribed in Table 1. The paratypes resemblethe holotype, with the following notewor-thy exceptions: females are larger thanmales; the snout of females is less protu-berant than in males; the forearm of fe-males is not swollen; and nuptial pads areabsent in females. One female (CM117018) differs in having more numerous

spiculae on the flanks, anterodorsal surfaceof the arm and dorsal surfaces of thethighs, and a few scattered spiculae on thedorsum of body (not on head); another fe-male (QCAZ 13938) lacks spiculae.

In males, the snout is acuminate in dor-sal view; the top of the snout is sloped an-teroventrally in lateral view. A few spiculaeare present on the anterior surface of thearm in three males (QCAZ 1917–18, 1926)(out of 12), and spiculae are virtually ab-sent on the flanks in four males (QCAZ1916, 1920, 1924–25). Malformations areapparent on QCAZ 1921 and 1927: theformer lacks Toe III on the left foot, andthe latter has two toes on the right foot.

In preservative, ventral color in malesvaries from uniform cream (e.g., QCAZ1918) to cream with dark brown marks(e.g., QCAZ 1927), to nearly uniform darkbrown (e.g., QCAZ 1925). In one speci-men (QCAZ 1921), the entire palmar sur-face is cream; thus, there is no markedcontrast between tubercles and adjacentsurfaces. In one female (CM 117018), theventer is cream with black marks, whereasin another (QCAZ 13938) the venter andventral surfaces of limbs are entirely black,except the border of the mandible, whichis brownish. In life, color variation mainlyis on the venter. QCAZ 1923 had a creamventer washed with diffuse green andlarge, brown marks; QCAZ 1924 had anentirely green venter and QCAZ 1925 hada brown venter with diffuse white marks.

Tadpoles.—Measurements (mm) of de-velopmental stages of a series of 14 larvae(QCAZ 3672) with the same data as theholotype are given in Table 3. The follow-ing description is based on an individual inStage 28 (QCAZ 3672) (Fig. 8B). Type-IVtadpole of Orton (1953). Tadpoles belongto the gastromyzophorous ecomorpholog-ical guild defined by Altig and Johnston(1989).

Body length (tip of snout–base of venttube) 5.9 mm, total length 16.2 mm. Bodyelongately ovoid, dorsoventrally com-pressed, about two thirds as high as wide,greatest width at anterior half of body.Snout bluntly rounded in dorsal view andin profile; body constricted at level of eyesand at spiracle; nostrils small, about one-


FIG. 7.—Ventral view of (A) right hand and (B) right foot of holotype of Atelopus nanay. Scale 5 1 mm.

TABLE 3.—Measurements (in mm) of developmentalstages sensu Gosner (1960) of 14 tadpoles of Atelopusnanay (QCAZ 3672); n 5 sample size, SD 5 standard

deviation.

Stage n

Body lengthmean 6 SD

(range)

Total lengthmean 6 SD

(range)

25 2 4.4 6 0.1(4.3–4.5)

11.2 6 0.4(10.0–11.4)

26 3 4.9 6 0.4(4.5–5.2)

13.0 6 0.7(12.4–13.8)

27 7 5.2 6 0.4(4.8–5.9)

14.2 6 1.1(12.7–15.5)

28 2 5.7 6 0.3(5.5–5.9)

15.0 6 1.7(13.8–16.2)

third the distance from eyes to tip ofsnout; eyes dorsal, directed dorsolaterally,diameter 0.7 mm; interocular distance 1.3mm (taken from the medial edges of thecorneas). Spiracle sinistral, half free, elon-

gate, ventral to horizontal body axis, di-rected posterodorsally, originating at mid-body; diameter of opening about equal tolength of free tube; vent tube short, me-dial. Caudal musculature robust anteriorly,narrowing abruptly posterior to midlengthof tail, terminating in a narrow strip justanterior to end of tail; dorsal fin highestposterior to midlength of tail; tail length36.4% of total length; dorsal and ventralfin heights 0.7 mm at midlength of tail,dorsal fin beginning well posterior to body,ventral fin beginning posterior to venttube, tip of tail rounded.

Mouth ventral, surrounded by labiaforming complete oral disc 3.8 mm wide;one row of marginal papillae on upper la-bium, on vertex between upper and lowerlabia, and on one-fourth of lower labium;papillae on rest of lower labium absent;one submarginal papilla anterior to vertex


FIG. 8.—Tadpole of A. nanay (QCAZ 3672): (A) lateral, (B) dorsal, and (C) ventral views. Scale 5 2 mm.

on upper labium; one submarginal papillajust posterior to vertex on lower labium.Labial tooth row formula 2/3; rows com-plete, about equal in length; jaw sheathsabout equal in length, upper jaw sheathnarrow, but wider medially; lower jawsheath V-shaped. Large ventral suctorialdisc covering area from posterior labiumto just anterior to end of body (at base ofvent tube), forming a complete, round discwith raised edges; suctorial disc broadenedat juncture with labium, few wrinkles pre-sent just posterior to juncture; roof of suc-torial disc bearing an elliptical fold poste-riorly; skin on edge and roof of discsmooth, skin on posterior margin of rooffinely rugose.

In preservative, dorsum, sides of body,and belly uniform brown; spiracle brown;tail musculature uniform brown for twothirds of its length, whereas distal one-

third white with scattered brown pigment.Dorsal fin brown at base, with numerousflecks posteriorly forming a reticulate pat-tern; scattered flecks on anterior half ofventral fin and reticulate pattern posteri-orly; oral disc brown; suctorial disc trans-lucent, except for brown flecks on roof. Inlife (based on color transparencies), bodyentirely black and fins translucent withblack flecks.

Comparisons: The tadpole of Atelopusnanay is similar to that of A. ignescens(Duellman and Lynch, 1969) and A. car-rikeri (senior synonym of A. leopereziiRuı́z-Carranza, et al., 1994 fide Coloma,1997), which are the only larvae of Atelo-pus (among 30 species for which infor-mation is available; Coloma et al., 2000)having an entirely black body. Tadpoles ofA. nanay are slightly smaller (total lengthof tadpoles in Stage 25 5 10.0–11.4, x̄ 5


11.2, n 5 2) than larvae of A. ignescens(total length of tadpoles in Stage 25 511.5–14.7, x̄ 5 13.2, n 5 10; data fromDuellman and Lynch, 1969), and muchsmaller than those of A. carrikeri [Ruı́z-Carranza et al. (1994) stated that thelength of tadpoles of A. carrikeri is twicethe length of previously known Atelopuslarvae]. Furthermore, tadpoles of A. nanaypossess a white blotch on the distal thirdof the tail musculature, whereas the tailmusculature of A. ignescens is entirelyblack. Tadpoles of A. nanay resemblethose of A. exiguus and A. peruensis inhaving a shallow tail. However, the lattertwo have pale (in life, pale golden and blu-ish white, respectively) marks on dorsumof body and caudal musculature (Colomaet al., 2000).

Distribution and natural history.—Thisspecies is known only from paramo habi-tats at the type locality, which is in Her-baceous Paramo in the Cordillera Occi-dental in Ecuador (Fig. 5). Annual meanprecipitation is 1000–2000 mm and annualmean temperature is 3–6 C. By day, on 28July 1989, some specimens were activeclose to streams and springs in an area ofmany interconnected pools; others wereon land under rocks. A female, collectedon 9 November 1987, has convoluted ovi-ducts with large ovarian eggs. Atelopus na-nay was found syntopically with Eleuther-odactylus cryophilius, Telmatobius niger,and Gastrotheca pseustes. Tadpoles werefound in a stream close to a spring whereadults of this species also were present.Some of the specimens described hereinare the last seen alive at that locality (in1989), despite subsequent efforts (28 No-vember 1992, 19 February 1993, 19 De-cember 1998) to find them.

Etymology.—The specific name nanayis a Quechua word, which means ‘‘sad-ness.’’ The name is used in reference tothe sadness that Ecuadorian indigenouspeoples often express when referring tothe disappearance of highland Atelopus.This name is used to call attention to cat-astrophic extinction of frogs of the genusAtelopus in the Ecuadorian Andes.

Osteology

The following description applies toadults of Atelopus guanujo and A. nanay,except when indicated otherwise. Intra-specific variation was not assessed in A. na-nay. Descriptions are based on clearedand double-stained skeletal preparations[QCAZ 1885 (male), 11205 (female),11250–51 (males), 11253 (male) of A. gu-anujo and QCAZ 1923 (male of A. na-nay)], except as indicated otherwise [forsome features, I include specimen KU289426 (male of A. guanujo)].

Skull.—(Figs. 9, 10). In dorsal view, theskull is triangular. A cartilaginous medialseptum nasi, paired tectum nasi, andpaired oblique cartilages are visible dor-sally as a continuous, thin strip at the an-terior end of the sphenethmoid and at theanterolateral border of nasals. The solumand septum nasi are synchondrotically (orsynostotically) fused to the sphenethmoid.The septum nasi contributes to the medial,anterodorsal configuration of the snoutthat is protruding relative to the anteriormargin of maxilla. Between the tectumnasi and the premaxilla are the alary andprenasal cartilages.

The septomaxilla is a U-shaped bonewith medial and lateral rami; the anteriorend is closed. The lateral ramus is broadand bears a nasal process. The latter variesin shape from an elongate, cylindrical andwell-differentiated process in Atelopus na-nay to a pyramidal, poorly differentiatedprocess in A. guanujo. Atelopus guanujoand A. nanay have a foramen in the lateralramus.

The entire surfaces of the single sphe-nethmoid are sculptured with a rough ap-pearance, except its posterior fourth,which is cartilaginous and borders the an-terolateral margins of the optic foramina.The posterior limits of ossification of sphe-nethmoid are at about one-third the lengthof the margin of the orbit in Atelopus na-nay, and at varying levels from one-thirdto slightly posterior to midway of orbitalregion in A. guanujo.

Paired prootics are synostotically unitedventromedially; dorsally, they are fused tothe posterolateral margins of frontoparie-


FIG. 9.—Skull of Atelopus guanujo (QCAZ 1885, male) in (A) dorsal, (B) ventral, and (C) lateral views.Scale 5 2 mm. Abbreviations: exoc 5 exoccipital; fpar 5 frontoparietal; max 5 maxilla; neopl 5 neopalatine;op 5 operculum; pmax 5 premaxilla; pro 5 prootic; prsph 5 parasphenoid; pter 5 pterygoid; spmax 5septomaxilla; qj 5 quadratojugal; spheth 5 sphenethmoid; sq 5 squamosal.

tals. Prootic surfaces are slightly rugose.The lateral portion (crista parotica) lacks aposterodorsal projection beyond the endof the posterior margin of the cranium.The anterior margin of prootic forms theposterior edge of the optic foramen. Theoculomotor and prootic foramina are lo-cated at the anteroventral portion of theprootic. The posteroventral margin of theprootic surrounds the ventral margins ofthe postotic foramen.

Both paired exoccipitals are synostoti-cally united dorso- and ventromediallycompletely encircling the foramen mag-num in bone. The posterior margins of ex-occipitals form the posteromedial wall ofthe otic capsule, the occipital condyle, and

the margin of the foramen magnum. Apostotic foramen is located just lateral tothe occipital condyle. The anteroventralmargins of exoccipitals are in contact withthe medial posteroventral margin of theparasphenoid.

A tympanic membrane, tympanic an-nulus, and stapes are absent. A well-de-veloped cartilaginous operculum lies in thefenestra ovalis. The operculum is conicaland oval in lateral view. The operculum isentirely cartilaginous in Atelopus nanay,whereas it is mineralized in A. guanujo(except in QCAZ 1885, in which it is most-ly cartilaginous).

The nasals are broadly triangular andseparated at midline; each possesses an


FIG. 10.—Skull of Atelopus nanay (QCAZ 1923, male) in (A) dorsal, (B) ventral, and (C) lateral views.Scale 5 2 mm.

elongate maxillary process that extendsventrolaterally toward the maxilla. Themaxillary process is pointed; the process isnarrowly separated from the maxilla.

The frontoparietals are paired, rectan-gular bones that roof the dorsal and dor-solateral surfaces of the braincase, fromthe level of the anterior margin of the orbitto close to the margin of the foramen mag-num. Medial articulation of the frontopar-ietals is absent in Atelopus nanay andvaries intraspecifically in A. guanujo; inthree specimens, more than half thelength of the bones is articulated whereasin three others a medial articulation is ab-sent. A suture line is visible in the speci-

mens with an articulated condition. Thelateral border of the frontoparietal, at levelof posterior half of orbit, lacks a flange,except in two specimens (out of five) of A.guanujo that possesses a small flange. Oc-cipital grooves are present; the grooves inA. guanujo are partially roofed, the pos-terior half of the groove is covered; the leftgroove in A. nanay is completely roofedwhereas the right groove is covered alongthe posterior three-fourths of its length.Posteriorly the frontoparietals are synos-totically fused posterolaterally to the pro-otics, and posteromedially to the medialanterodorsal margins of the exoccipitals.

The paired medially-separated vomers


are crescent-shaped, triradiate, and eden-tate. The anterior ramus of the vomer isdirected parallel to the midline in Atelopusnanay and anterolaterally and ventrally to-ward the anterior tip of the maxilla in A.guanujo; the terminus of the anterior ra-mus is pointed in A. nanay whereas is ei-ther pointed (QCAZ 11250, 11253) ortruncate in A. guanujo. In A. guanujo, theanterior ramus varies from long to short;it is long and nearly reaches the maxillaein QCAZ 11251; it is shorter or as long asthe prechoanal ramus in QCAZ 1885. InA. nanay, the anterior ramus is about aslong as prechoanal ramus. Pre- and po-stchoanal rami surround the medial, an-tero-, and posteromedial margins of thechoana. The prechoanal ramus is directedtowards the maxillae, whereas the po-stchoanal ramus is either parallel to themidline or directed posterolaterally. Therami are pointed in A. nanay, but in A.guanujo their distal ends may be roundedor truncate. The vomers do not contact theoverlying sphenethmoid.

The neopalatine is an elongate, trian-gular, or subarcuate bone that underliesthe planum antorbitale and sphenethmoidorbital margin; it extends from the sphe-nethmoid toward the maxilla. A gap existsbetween the lateral margin of neopalatineand maxilla. The maxillary half of neopa-latine is widest and its lateral border isusually pointed; the medial end is pointedand the amount of overlap with spheneth-moid is variable. The anterior border ofthe neopalatine is slightly rounded in Ate-lopus nanay and two specimens (QCAZ1885, 11251) of A. guanujo, whereas it hasan apex in the other three specimens(QCAZ 11205, 11250, 11253) of A. guanu-jo.

The parasphenoid has an inverted T-shape and invests the ventral surface of theneurocranium; it extends from the levelposterior to neopalatines (from the ante-rior half of the orbit) toward, but notreaching the margin of the foramen mag-num; its surfaces are smooth. The cultri-form process abuts [in Atelopus nanay,and one specimen (QCAZ 1885) A. gu-anujo] or underlies the posterior part ofthe sphenethmoid. The terminal end of

the cultriform process is rounded in A. na-nay and in three individuals of A. guanujo;it is acutely pointed in other three of A.guanujo. The parasphenoid alae are ori-ented perpendicular or slightly posterolat-eral to the cultriform process. The poste-rior margin of parasphenoid is nearlystraight, bearing a subquadrangular (A. gu-anujo) or rounded (A. nanay) posteriorprocess medially.

Each premaxilla consists of a dorsal ala-ry process, a biradiate pars palatina, andan edentate, pars dentalis in the anteriormargin. The alary process projects antero-laterally. In Atelopus guanujo, the lowerhalf of the alary process is wider than theupper half, whereas in A. nanay, it is onlyslightly wider. The pars palatina has twowell-differentiated processes, a medial(palatine) and a lateral process; the lateralprocess is almost as twice the length of thepalatine process. The medial process ofthe pars palatina is a well-developed, tri-angular, pointed ramus in A. nanay. In A.guanujo, there is intraspecific variation;the medial process varies from well-devel-oped in QCAZ 1885 to not well-differen-tiated in QCAZ 11253. A foramen is pre-sent along the length of the medial processof the pars palatina, and a ridge is evidenton its ventral surface. The inner margin ofthe pars palatina, where the palatine andlateral processes come together, forms aconcave border. The posterior end of thepars dentalis is pointed in A. guanujo (ex-cept in QCAZ 11205, in which it is trun-cate) and A. nanay and articulates with themaxillae. In one specimen of A. guanujo(QCAZ 1885), a small, sesamoid oval bonelies at the contact zone between the parspalatina of premaxillae and maxillae.

The maxilla is an elongate, edentatebone located between the premaxilla andthe quadratojugal. A horizontal pars pala-tina of the maxilla extends along the lin-gual margin of the maxilla; posteriorly thisshelf contacts the anterior ramus of thepterygoid. The maxilla is deepest anteri-orly and bears the pars facialis which issubtriangular and directed medially; itsdorsal margin is nearly even in Atelopusnanay and bears small bumps that nearlycontact the neopalatine or the maxillary


process of the nasal in A. guanujo. The an-terior end of the maxilla is truncate,whereas the posterior end is pointed. Theanterior end slightly overlaps the posteriorend of premaxilla. The posterior end of themaxilla is separated from the quadratoju-gal; in A. nanay, it is separated by a lengthsimilar to that of the quadratojugal, where-as in A. guanujo, the distance of separationvaries from one third to the full length ofquadratojugal.

The quadratojugal is a small, L-shapedbone underlying the margin of the ventralarm of the squamosal; the posterior ramusarticulates with the pars articularis of thepalatoquadrate and the anterior ramus isseparated from the maxilla.

The squamosal bears one or two dorsalrami, and a ventral ramus. The otic ramus(posterior, dorsal ramus) is moderately ex-panded towards the dorsolateral surface ofthe prootic; the otic ramus varies in thedegree to which it overlaps the prootic;sculpturing on this element is nearly ab-sent. The otic ramus overlaps nearly theentire length of the crista parotica (QCAZ11253) or leaves only about one fourth ofthe posterior end free. Atelopus nanay andone specimen of A. guanujo (QCAZ 1885)lack a clearly differentiated zygomatic ra-mus (anterior, dorsal ramus) whereas fourspecimens of A. guanujo bear a small tri-angular zygomatic ramus. In one specimen(QCAZ 11250), there is a small foramenlocated in the zygomatic ramus. The anglebetween the dorsal part of the squamosaland the anterior margin of the dorsal por-tion of the ventral ramus varies from acuteto nearly perpendicular in A. guanujo andis nearly perpendicular in A. nanay. Thelower portion of the ventral ramus is flatand blade-like and invests the palatoquad-rate laterally. The squamosal has a keelthat extends from the outer side of dorsalarm toward the outer, upper half of theventral arm.

The triradiate pterygoid bears anterior,medial, and posterior rami; the anterior ra-mus articulates with the maxilla anteriorly;the medial ramus invests the pseudobasalprocess of the prootic, and a short, poste-rior ramus invests the medial surface ofthe palatoquadrate. The articulation of the

medial pterygoid ramus with the prooticinvolves two poorly differentiated headsthat articulate with the ventrolateral sur-face of the pseudobasal process of the oticcapsule.

The palatoquadrate is primarily cartilag-inous and is invested by the ventral arm ofthe squamosal laterally and the posteriorramus of pterygoid medially and postero-medially. The ventral pars articularis as-sociated with the quadratojugal is cartilag-inous with a slight mineralization in twospecimens of Atelopus guanujo (QCAZ11205, 11250).

The lower jaw is composed of fourpaired elements. Anteriorly, the mento-meckelian bone bears a cartilaginous me-dial margin that forms a symphysis withthe opposite medial margin. Posteriorlythe dorsal portion of the mentomeckelianbone is fused to the dentary; the latter isthin when viewed laterally. The edentatedentary overlaps the angulosplenial dor-sally at the anterior portion and extendslaterally toward the posterior region. Theangulosplenial is located at the lingual sur-face of the mandible from a point behindone length of mentomeckelian bone andto the posterior end of the mandible. Be-tween the dentary and angulosplenial aMeckel’s cartilage extends from a pointjust behind the mentomeckelian bone tothe posterior end of the mandible. Meck-el’s cartilage is invested by the dentary an-terodorsally and anterolaterally, and by theangulosplenial posteroventrally and poster-omedially. The dentary terminates poste-riorly at a level just anterior to the begin-ning of the curve of angulosplenial.

Hyobranchium.—(Fig. 11). The hyob-ranchium consists of a cartilaginous plate,two hyalia, a pair of basally narrow alary(anterolateral) processes, usually a pair ofposterolateral processes, and two ossifiedposteromedial processes. There is varia-tion in shape of the plate (5 corpus), aswell as in the presence of the posterolat-eral processes. The plate is longer thanwide in Atelopus guanujo (ratio of hyoidplate length/hyoid plate minimum width2.17–3.91, n 5 6) and A. nanay (3.54, n 51). Endochondral mineralization in corpusof hyoid plate are present in two speci-


FIG. 11.—Ventral view of hyobranchium of (A)Atelopus guanujo (QCAZ 1885, male) and (B) A. na-nay (QCAZ 1923, male). Dotted pattern indicatescartilage. Drawings of posteromedial processes maybe distorted because they are not in a horizontalplane. Scale 5 2 mm.

FIG. 12.—Ventral views of pectoral girdles of (A)Atelopus guanujo (QCAZ 1885, male) and (B) A. na-nay (QCAZ 1923, male). Dotted pattern indicatescartilage. Calcified areas are indicated by irregularstippling on the overlay. Scale 5 5 mm.

mens of A. guanujo: QCAZ 11250 pos-sesses mineralization, which covers the en-tire corpus, whereas QCAZ 11251 pos-sesses mineralization at lateral and poste-rior margins of corpus. Each hyale bears ashort anterior process, which may bestraight or deflected laterally. The lengthof the anterior process varies intraspecifi-cally in A. guanujo, but it does not rejointhe hyale. Two specimens of A. guanujo(QCAZ 11251, 11253) also bear a smallprocess on the hyale lateral to the anteriorprocess; in A. nanay, this process is absent.As described by the configuration of thehyalia, the shape of the hyoglossal sinusvaries; the lateral margins of the hyoglossalsinus diverge anteriorly except in onespecimen of A. guanujo (QCAZ 11250) inwhich they are almost parallel. The an-terolateral processes always are present,and vary in shape. The narrow base inboth species supports an anterior flangeand another smaller posterior flange. Pos-terolateral processes vary in length inthree specimens of A. guanujo and are ab-sent in three, whereas in A. nanay they areabsent. The long, bony posteromedial pro-cesses bear cartilaginous ends. In fourspecimens (QCAZ 1885, 11205, 11251,KU 289426) (out of six) of A. guanujo,each posteromedial process bears a smalllateral expansion at about half its length.

Pectoral girdle.—(Fig. 12). Each half of

the firmisternal pectoral girdle is com-posed of the suprascapula, scapula andzonal area. The cartilaginous suprascapulais fan shaped and is mineralized. Variationexists in degree of mineralization fromscattered to heavy in Atelopus guanujo,and with little mineralization in A. nanay.A bony, bicapitate cleithrum is present onthe anterior margin of the suprascapula.The cleithrum is L-shaped, with the elon-gate arm located along the anterior marginand the short arm along the scapular bor-der of suprascapula. The scapula is shortand stocky, and fused to clavicle and cor-acoid at its medial end. The pars acromi-alis and pars glenoidalis of scapula are notwell defined; but their margins are evidentby virtue of a separation zone that forms


a small oval fenestra located at the innermargin of glenoid cavity. A prezonal ele-ment (5 omosternum) is absent. Zonalcomponents are clavicles, procoracoids,coracoids and epicoracoids. The clavicle isoriented transversely or slightly antero-medially; it is fused at its distal end withthe pars acromialis of the scapula and dis-tal end of coracoid.

At midline, the epicoracoid cartilagesare synchondrotically fused throughouttheir lengths and lack any trace of a suture.Anteriorly, at midregion, the epicoracoidsare fused to procoracoid cartilages andform an epicoracoid bridge. Posteriorly,the mineralized epicoracoid cartilages arenot fused to the sternum. A suture line isvisible between the platelike epicoracoidsand the sternum. The shape of this sutureline possess a triangular projection at mid-length that extends anteriorly, except intwo specimens (QCAZ 11250–51) of Ate-lopus guanujo on which the suture line isnearly straight. A calcified procoracoid car-tilage extends from midlength to medialend of clavicles; the clavicle borders it dor-sally and ventrally. Epicoracoids and cor-acoid margins form the medial margin ofthe pectoral fenestra.

Other differences in the pectoral girdleinclude the degree of development of pro-cesses along the anterior margin of theclavicle and scapula, and the length andshape of the sternum. There are two tri-angular processes; the proximal, a clavic-ular acromium, is larger than the distal.The former lies at the joint between theclavicle and scapula; its orientation variesfrom nearly parallel in relation to the lon-gitudinal axis in two (out of five) Atelopusguanujo to forming an angle in A. guanujoand A. nanay. The distal process is welldifferentiated in A. guanujo but absent inA. nanay.

The sternum varies in length. The ratiosof sternum length to total length of girdleat midline are 0.45–0. 54 (n 5 6) in Ate-lopus guanujo, and 0.58 for A. nanay.There is no clear differentiation betweena mesosternun and a xiphisternum. Thesternum is entirely calcified, except for adistal cartilaginous terminus. One speci-men of A. guanujo (QCAZ 11250) has a

distal mineralized terminus. In a specimenof A. guanujo (QCAZ 1885), cartilaginousregions are present at the anterior andposterior ends. The sterna are expandeddistally in A. guanujo and about uniformin width throughout its length in A. nanay;the ratios of sternum maximum width tosternum minimum width are 1.25–2.05 (n5 6) in A. guanujo, and 1.34 for A. nanay.The terminal end of the sternum is trun-cate and unicapitate in both species.

Pelvic girdle.—(Fig. 13). The pelvic gir-dle is V-shaped in dorsal aspect in bothspecies; it is composed of paired ilia, is-chia, and pubes. The ilial shafts are cylin-drical and bear cartilaginous anterior tips.Each ilium extends anteriorly beneath thesacral diapophysis; the distal end of the il-ium lies at the anterior border of the di-apophysis in one specimen (QCAZ 1885),whereas in others it does not reach theborder. Posteriorly, the ilia are united in amedial symphysis with the ischia and pu-bes; the pubis is ossified. The iliosacral ar-ticulation is Type IIA (sensu Emerson,1979), thus the ilium is attached to the di-apophysis by a broad, transverse ligament.There is a small, sesamoid bone anteriorto the distal end of the ilium in one spec-imen of Atelopus guanujo.

Vertebral column.—(Fig. 13). The ver-tebral column contains nine elements. Theatlas complex is formed by the completefusion of Presacrals I and II. There are sixindependent trunk vertebra (PresacralsIII–VIII), the sacrum, and the coccyx. Ver-tebral centra are procoelus, and the sa-crum bears a bicondylar articulation withthe coccyx. The holochordal centra are cy-lindrical and solid. The neural arches ofPresacrals IV–VIII are not imbricate. Ver-tebral shields and free ribs are absent.

In the atlas complex, one pair of trans-verse processes is present and they are ori-ented anterolaterally in Atelopus guanujo,whereas in A. nanay, the transverse pro-cesses are oriented nearly perpendicular tothe midline. In Presacrals III–VIII, the an-terior and posterior margins of the trans-verse processes are even in both species.The proximal end of the transverse pro-cesses is slightly wider than the terminalend in Atelopus guanujo (except in one


FIG. 13.—Dorsal views of vertebral columns and pelvic girdles of (A) Atelopus nanay (QCAZ 1923, male),(B) A. guanujo (QCAZ 1885, male), and (C) A. guanujo (QCAZ 11205, female). Scale 5 5 mm.

specimen, QCAZ 1885) and A. nanay ow-ing to the presence of a flange, which di-minishes in size distally. The transverseprocesses are slightly shorter than the sa-cral diapophyses and their ends form aslightly concave line. The transverse pro-cesses of Presacrals III and VIII are thelargest. Presacral IV bears the widesttransverse processes in A. guanujo; how-ever, in A. nanay, Presacral III bears widertransverse processes than Presacral IV.Orientation of the transverse processesvaries markedly. The processes of Presa-cral III project posteroventrally, whereasthose of Presacral IV have less pronounced

posterior orientation; the processes of Pre-sacrals V–VIII become oriented more an-teriorly. In one specimen of A. guanujo(QCAZ 11250), all the transverse process-es are oriented nearly perpendicular to themidline. In A. guanujo, the spacing amongtransverse processes between vertebravaries intraspecifically because of sexualdimorphism, with males having smallerspaces than females (Fig. 13B,C). Sculp-turing on the dorsal surfaces of neuralarches is indistinct in both species. A cen-tral pit in most vertebrae is absent; thereare scattered small pits in one individual(QCAZ 1885) of A. guanujo.


The shape of the sacrum varies and isrelated to the sacral diapophyses, whichare expanded. One specimen of Atelopusguanujo (QCAZ 11250) bears a nearly hor-izontal anterior and posterior border,whereas in A. nanay and others of A. gu-anujo, the anterior border is more angularand directed anterolaterally, and the pos-terior border extremely angular. In A. na-nay, the sacrococcygeal articulation is bi-condylar; in A. guanujo, the sacrum andcoccyx are fused dorsally and ventrally intwo specimens, and unfused in four. Thecoccyx is an elongate, subcylindrical bonethat has an inverted triangular anteriorend, which is formed by small vestigial lat-eral processes. Lateral processes are ab-sent in one specimen (QCAZ 11253) of A.guanujo, whereas in other specimens(QCAZ 1885, 11205) are present only atone side. Lateral flanges along nearly en-tire length of coccyx are absent.

Forelimb.—A humerus, radioulna, car-pal elements, four digits (II–V), and a pre-pollex make up the forelimb. Intercalaryelements are absent. Torsion is absent inDigit II (first finger). The following carpalelements were observed: a postaxial cen-trale (5 radiale), an ulnare, the ElementY, distal Carpals 3, 4, and 5 fused in a sin-gle bone, and distal Carpal 2. Additionally,two other elements were identified. Be-cause their origin and homologies to carpalelements in other taxa are uncertain, I pro-visionally consider these two bones to besesamoid bones (Sesamoids I and II). Ses-amoid I is located under distal Carpal 3 14 1 5; it is round. Sesamoid II is a smaller,round bone located at the distal end of theradius at the posterior upper side of thepostaxial centrale, and is larger in Atelopusguanujo than in A. nanay. Terminal pha-langes on the digits are T-shaped. Thephalangeal formula is 2-2-3-3 in Atelopusguanujo, whereas A. nanay has a reducedDigit II (5 Finger I) with a formula of 1-2-3-3. The prepollex is calcified. It pos-sesses two elements in Atelopus nanay andone to three elements in A. guanujo. InQCAZ 1885, the prepollex consists of twoelements on the left hand and three on theright hand. In one specimen (QCAZ11250), the prepollex is a single bone with

the proximal end nearly completely fusedto the adjacent proximal end of the firstphalanxe of Finger I. In QCAZ 11251, theprepollex of the right hand is similar to thelatter, but the left hand has a prepollexwith two elements.

Hind limb.—The hind limb comprises afemur, tibiofibula, calcified cartilago plan-taris, tarsal elements, five metatarsals, fivedigits, and prehallux. The tibiofibula issmaller in Atelopus nanay than in A. gu-anujo. The ratios of female tibiofibulalengths to snout–vent lengths are 0.32–0.39 (n 5 13) in A. guanujo, and 0.31 forA. nanay. The tarsal elements include thetibiale (5 calcaneum), fibulare (5 astrag-alus), Tarsal I, Tarsals 2 1 3 (fused), cen-trale, and os sesamoides tarsale. Atelopusnanay has one os sesamoides tarsale thatis elliptical to round in shape and is locat-ed ventral to the distal head of the fibu-lare. The same bone also is present in A.guanujo, and a smaller one is located ven-tral to Tarsal I. In some specimens of A.guanujo (e.g., QCAZ 11205, 11251,11253), two to four additional os sesamo-ides tarsale are located close to the twolarger ones. The phalangeal formula of thepes is 2-2-3-4-3. The prehallux is ossifiedwith the distal end cartilaginous in A. na-nay, whereas is either completely ossifiedor bears a cartilaginous end in A. guanujo.In A. nanay, the number of elementsvaries bilaterally, and there is one elementon one side and two on the other side. InA. guanujo, there are one to three ele-ments, and intraindividual bilateral varia-tion occurs.

DISCUSSIONKnowledge of the alpha-level systemat-

ics of Atelopus continues to increase withspecies descriptions (Ardila-Robayo andRuiz-Carranza, 1998; Ibañez et al., 1995;Lötters and Henzl, 2000; Rivero and Mo-rales, 1995). However, hypotheses of re-lationships among Atelopus include fewspecies and/or are poorly supported (LaMarca, 1983; Lescure, 1973, 1981; Lynch,1993; McDiarmid, 1971, 1973; Peters,1973; Rivero, 1963a,b, 1984; Rivero andSerna, 1993; Rivero and Morales, 1995).Placement of the new species described


herein in any of the species groups pro-posed by several authors (e.g., Lynch,1993; Peters, 1973) is ambiguous. The A.flavescens group of Lynch (1993) wouldaccommodate A. nanay based on its pha-langeal formulae; however, the historicalreality of this group has been questioned(Coloma, 1997; Vélez-Rodrı́guez andRuiz-Carranza, 1997) because of intra-in-dividual and intraspecific variation notedin the lack of a terminal phalanx on thumb,which defines it. The phylogenetic alloca-tion of A. guanujo within the A. ignescensgroups—either sensu Peters, 1973 or sen-su Lynch, 1993—is doubtful because oflack of evidence of their monophyly, aproblem which was already recognized byboth authors. The A. ignescens group sen-su Peters (1973) is diagnosed based on thelength of limbs relative to body length,whereas the ignescens group sensu Lynch(1993) lacks diagnostic characters. Al-though I could allocate the new species tophenetic groupings, I prefer not to do so,because I think that such assignmentmight introduce bias to future attempts toinfer phylogenetic relationships.

The description of two new species inthis paper brings the total number of Ate-lopus from Ecuador to 18. Among these,15 species occur in the Andes at elevations.1000 m (Coloma and Quiguango, 2000).The endemism and diversity of Atelopuscorroborates a pattern of biodiversitywhich resides primarily in montane com-ponents when contrasted to lowland eco-systems (Lötters, 1996; see also Petersonet al., 1993 for similar biodiversity patternsin Mexico). In lowland ecosystems, al-though alpha diversity (number of speciesat a single site) may be high, beta diversity(in two or more sites in a given region) ofsome groups of plants and animals is lessthan in the Andes: for example, in Ecu-adorian bamboos (Clark, 1997), Ecuador-ian Compositae (Funk, 1997), SouthAmerican amphibians (Duellman, 1999),Colombian frogs (Lynch et al., 1997), andEcuadorian frogs (Coloma and Quiguango,2000). Alpha and beta diversity are higherin the Andes than in lowland ecosystemsin Ecuadorian frogs such as Centrolenidae(Duellman and Burrowes, 1989; Lynch

and Duellman, 1973), Colostethus (Colo-ma, 1995), and Eleutherodactylus (Lynchand Duellman, 1980, 1997), among others.These patterns suggest the need for fur-ther biological research and conservationefforts directed to Andean habitats, wheredisappearance of frog populations are amatter of concern.

RESUMENSe reconocen dos especies de Atelopus

de Ecuador nuevas para la ciencia; la unade la Hoya de Chimbo, la otra de los Pá-ramos del Cajas; ambas provienen de re-giones altas en la Cordillera Occidental delos Andes. Se proveen descripciones osteo-lógicas de ambas especies y del renacuajode torrente de una de las especies nuevas.Los patrones de diversidad y endemismode Atelopus justifican la necesidad de ma-yores investigaciones biológicas y esfuerzosde conservación dirigidos a habitats andi-nos.

Acknowledgments.—For the loan of specimens, Ithank M. C. Ardila-R. (ICN), E. J. Censky (CM), W.E. Duellman (KU), W. R. Heyer (USNM), E. LaMarca (ULAGB), B. Schätti and J. Mariaux(MHNG), and J. Vindum (CAS). I am also indebtedto my field companions: F. Campos-Yánez, M. S.Hoogmoed, and S. de la Torre. G. Onore and M.Tapia kindly helped me obtain additional specimensof Atelopus guanujo.

Parts of this study were included in my Ph.D. dis-sertation presented to the Department of Systematicsand Ecology of The University of Kansas. W. E.Duellman, R. Timm, and L. Trueb provided valuablesuggestions and encouragement. S. Lötters, S. Ron,and J. J. Wiens reviewed preliminary versions of thismanuscript. S. Lötters, and J. E. Simmons helped as-semble bibliographic references. A. Quiguango drewFigs. 2A and 12. G. Onore and A. Quiguango helpedin numerous ways, and I am greatly indebted tothem.

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Accepted: 12 June 2001Associate Editor: Joseph Mendelson III

APPENDIX I

Specimens Examined

Atelopus bomolochos.—ECUADOR: ProvinciaAzuay: Vic. E of Cuenca, 2535 m, (CAS 85l39–41);Sevilla de Oro, 2800 m, 82 km E of Cuenca, (CAS85341, 93912; paratopotypes); 3 km E of Sigsig, 2460m, (QCAZ 1893–94); 10 km S Cutchil, 2900 m,(QCAZ 1861–69, 1896–99); 7.6 km S Cutchil, 2880m, (QCAZ 1900); 6.2 km S Cutchil, 2800 m, (QCAZ1901); Rı́o Chilpe, 10–15 km E Luz–Marı́a, aprox.2500 m, (QCAZ 1860); 13 km NE Nieves, (CM117039); Provincia Cañar: Between Juncal and Gen-eral Morales, approx. 2500 m, (CAS 93906, paratype);


Provincia Loja: Loja (locality questioned by Peters[1973:37]), (USNM 236942–44); Provincia MoronaSantiago: Sucúa, (USNM 283538–41).

Atelopus carbonerensis.—VENEZUELA: EstadoMérida: 32 km NW Mérida, road to Azulita, 2010 m,(KU 132857, KU 132860, KU 132864, KU 132867–68, KU 132871).

Atelopus ignescens.—ECUADOR: ProvinciasNapo and Pichincha: Páramo de Guamanı́, La Virgen,3800–4200 m, (MHNG 2410.1–6, 2409.95–100,2273.48, 2273.61, 2273.81–82, 2273.84–97).

Atelopus sorianoi.—VENEZUELA: Estado Méri-da: Paramito de San Francisco, 2450 m [QCAZ 4910,4911 (C&S)].

Atelopus sp.—COLOMBIA: Departamento Nari-ño: 8 Km E Pasto, 3050 m, (KU 169267); 12 Km EPasto, 3050 m, (KU 169284, 169292); Vereda de SanFernando, 7.5 Km E Pasto, 2800 m, (KU 169267);Hacienda San Gerardo, 2900–3000 m, [ICN 450,459, 469, 471, 472 (C&S), 473, 481 (C&S)]; El En-cano, 2800 m, [ICN 3633 (C&S), 3636–38]; VolcánGaleras, 3200 m, [ICN 3406].

TWO NEW SPECIES OF ATELOPUS (ANURA: BUFONIDAE)...(from Laguna La Cocha, Departamento Narin˜o,...

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