A NEW SPECIES OF TELMATOBIUS (ANURA: LEPTODACTYLIDAE) … · 253 Herpetologica, 59(2), 2003, 253–270 q 2003 by The Herpetologists’ League, Inc. A NEW SPECIES OF TELMATOBIUS (ANURA:

253

Herpetologica, 59(2), 2003, 253–270q 2003 by The Herpetologists’ League, Inc.

A NEW SPECIES OF TELMATOBIUS(ANURA: LEPTODACTYLIDAE) FROM RIO VILAMA,

NORTHERN CHILE, AND THE REDESCRIPTIONOF T. HALLI NOBLE

J. RAMON FORMAS1,4, EDGAR BENAVIDES2,3, AND CESAR CUEVAS1

1Instituto de Zoologıa, Universidad Austral de Chile, Casilla 567, Valdivia, Chile2Casilla 160-C, Departamento de Zoologıa, Facultad de Ciencias Naturales y Oceanograficas,

Universidad de Concepcion, Concepcion, Chile

ABSTRACT: We describe a new species, Telmatobius vilamensis, from Rıo Vilama, near San Pedrode Atacama, Chile. Previously confused with Telmatobius halli Noble, 1938, this new species isdiagnosed by distinct characteristics of the adult and larval morphology and chromosomes. The newspecies is the southernmost representative of the genus Telmatobius in Chile. Telmatobius halliNoble, 1938 is redescribed, and both taxa are compared with eight Chilean congeners that occurin the Andes.

Key words: Anura; Karyotype; Leptodactylidae; Osteology; San Pedro de Atacama; Tadpoles;Telmatobius halli redescription; Telmatobius vilamensis new species

SPECIES of the genus Telmatobius rep-resent one of the most diverse componentsof the Andean batrachofauna above 2000m. The genus is comprised of around 50species distributed from southern Ecuadorto different latitudes on the eastern (228519 S, 688 119 W) and western (298 399 S,688 369 W) slopes of the Andes of northernChile and Argentina. New species have re-cently been described primarily from Bo-livia and Peru (de la Riva, 1994a,b; Lavillaand Ergueta, 1995a,b, 1999; Salas andSinsch, 1996). Nevertheless, recent her-petological exploration of the southwest-ern limits of the range of the genus hasresulted in the unexpected discovery ofthree new Chilean species of Telmatobius:T. dankoi (Formas et al., 1999), T. philippii(Cuevas and Formas, 2002), and T. fron-teriensis (Benavides et al., 2002a). Thesethree are in addition to five other speciesof Telmatobius previously reported fromnorthern Chile (T. peruvianus Wiegmann,1835; T. marmoratus [Dumeril and Bi-bron, 1841]; T. halli Noble, 1938; T. pefau-ri Veloso and Trueb, 1976; and T. zapa-huirensis Veloso et al., 1982; Fig. 1). An-

3 PRESENT ADDRESS: Department of Zoology andM. L. Bean Life Science Museum, Brigham YoungUniversity, Provo UT 84602, USA.

4 CORRESPONDENCE: e-mail, [email protected]

other new species described herein is thesouthernmost representative of the genusin Chile. Diagnosis of the new species isbased on external morphology of adultsand tadpoles, osteology, karyotype, chro-mosomal C-banded patterns, and Ag-NOR(nucleolar organizer region) position. Be-cause this new species has been confusedwith T. halli (e.g., Cei, 1962; Veloso et al.,1982), a redescription, diagnosis, and illus-trations of the holotype of T. halli are pre-sented. Finally, we provide morphologicaland karyological comparisons among Chi-lean species as well as a key to their iden-tification.

MATERIALS AND METHODS

This description is based on 25 adultfrogs, a juvenile, and 5 tadpoles collectedat Rıo Vilama near San Pedro de Atacama,Chile (Fig. 1). Geographic coordinateswere obtained using a Trimble Global Po-sitioning System (GPS). With the help ofnets, specimens were collected belowdense vegetation in flowing water. Photo-graphs, notes on the habitat, and color inlife were taken in the field at the time ofcapture. Vouchers and additional speci-mens used in this study (Appendix I) aredeposited in the following collections: Co-leccion Boliviana de Fauna (CBF); Insti-tuto de Zoologıa, Universidad Austral de

254 [Vol. 59, No. 2HERPETOLOGICA

FIG. 1.—Distribution of the species of Telmatobiusin Chile. The inset shows the type locality of Telma-tobius vilamensis.

Chile (IZUA); Museo de Zoologıa, Univ-ersidad de Concepcion (MZUC); Depar-tamento de Biologıa Celular y GeneticaUniversidad de Chile (DBCUCH); Depar-tamento Biomedico, Universidad de An-tofagasta (DBMUA); and Museo Nacionalde Historia Natural, Chile (MNHN). Theredescription of T. halli Noble is based onthe holotype and two paratypes depositedin the American Museum of Natural His-tory (AMNH).

The following variables were measuredusing dial calipers (to the nearest 0.1 mm):snout–vent length (SVL), tibia length(knee to heel), foot length (proximal edgeinner metatarsal tubercle), head length(posterior corner of jaw to tip of thesnout), head width (from posterior cornerof jaw), interorbital distance, eye–nostrildistance (from anterior corner of eye), in-ternarial distance, eye diameter (betweenanterior and posterior corners of eye),hand length (proximal edge of outer meta-carpal tubercle to tip of third finger), andradioulnar length (elbow to distal edge ofouter metacarpal tubercle). Sexual matu-rity was assessed by presence of eggs infemales and the degree of development ofnuptial excrescences in males. Formulasfor toe webbing are those of Savage andHeyer (1967), as modified by Myers and

Duellman (1982). Osteological observa-tions were made on three adult specimens(two males IZUA 3146 [SVL 44.16, 46.81mm] and a female IZUA 3224 [SVL 47.51mm]); cleared-and-stained skeletons wereprepared using the methods of Hollister(1934) and Song and Parenti (1995). Car-pal elements were identified according toFabrezi and Alberch (1996).

Tadpoles (IZUA 3132) were staged ac-cording to Gosner (1960). The followingmeasurements were taken: total length(distance from the tip of the snout to thetip of the tail), body length (distance fromthe tip of the snout to the origin of thehind limb), body depth (maximum dis-tance between the dorsal and ventral sur-faces of the body), fin depth (maximumdistance between margin of dorsal andventral fins), snout–nostril distance (dis-tance from the tip of the snout and theanterior border of the nostril), eye diam-eter (maximum distance between the an-terior and posterior margins of the eye),and mouth width (maximum distance be-tween the lateral margins of the oral disc).

Chromosomal characteristics were re-corded from two male specimens (IZUA3225–26), which were injected with 0.1%colchicine. After 2 h, frogs were anaesthe-tized with diethyl ether. A ventral incisionwas made under sterile conditions, and in-testines were removed. Metaphase plateswere obtained by macerating intestinal ep-ithelium fragments. These were hypoton-ically treated with distilled water, fixed inacetic acid-alcohol (1:3), and finally placedin 45% acetic acid. Small fragments of tis-sue were placed between a glass slide andcover slip and dipped in liquid nitrogen.Thereafter, the cover slip was removedwith a razor blade and chromosomes wereallowed to air dry. After 3 d, chromosomeswere stained for 15 min in Sorensen’sphosphate buffer (pH 6.8), containing 4%Giemsa solution (Formas, 1991). Chro-mosomes were stained to reveal C-bandpatterns and Ag-NOR position, followingthe methodologies of Sumner (1972) andRufas et al. (1982). Centromeric positionswere determined according to Levan et al.(1964).

June 2003] 255HERPETOLOGICA

ACCOUNTS OF SPECIES

Telmatobius vilamensis sp. nov.

Holotype.—IZUA 3080, adult male col-lected by C. Cuevas, J. R. Formas, and C.Jara at the Rıo Vilama (228 519 430 S, 688239 250 W; 3110 m), 6.5 km NE (by road)of San Pedro de Atacama, Provincia ElLoa, Region de Antofagasta, Chile, on 17April 1997 (Fig. 1).

Paratypes.—IZUA 3081, adult male;IZUA 3224 (cleared-and-stained adult fe-male); same data as holotype; CBF 3760–62 (two females and a juvenile); MZUC24104–06 (three males); MZUC 25725–38(nine females and five males); collected byE. Benavides, H. Dıaz, M. Hengst, and J.C. Ortiz at the type locality on 17 Decem-ber 1998.

Diagnosis.—A moderate-sized Telmato-bius (38.36–50.81 mm) having the follow-ing combination of characters: (1) premax-illary and maxillary teeth absent, (2) vo-mers rudimentary or absent, (3) neopala-tines reduced, (4) tongue nearly ovoid,elongate, almost adhered to the floor ofmouth, posterior border free, (5) choanaelarge, circular, (6) tympanum, tympanicannulus, and columella absent, (7) snoutdepressed in lateral view, (8) postfemoralfolds present, (9) dorsum olive green (inlife), (10) cranium poorly ossified, (11) 26bi-armed chromosomes, (12) toes webbed,(13) outer border of Toe V moderatelyfringed.

Telmatobius vilamensis differs from allother known Telmatobius by having aprominently pointed snout (Fig. 2A), aswell as a lean, hydrodynamic body shape.Among Chilean species, T. dankoi mostclosely resembles T. vilamensis in overallmorphology. Both taxa share the followingcharacters: postfemoral folds (thinner andsmaller in T. vilamensis), premaxillary andmaxillary teeth absent, oral disc transan-gular, tympanum and tympanic annulusabsent, and size moderate. Telmatobius vi-lamensis is distinguished by its smoothskin (spiny in both sexes of T. dankoi),tongue nearly ovoid (rounded in T. dan-koi), snout strongly depressed (not de-pressed in T. dankoi), and cranium poorlyossified (well ossified in T. dankoi).

Despite obvious external differences,both T. vilamensis and T. dankoi have beenconfused with T. halli (e.g., Cei, 1962,1986; Northland et al., 1990; Veloso et al.,1982). These misidentifications probablyresulted from the incomplete nature of theoriginal description of T. halli (Noble,1938) and the lack of an illustration of thetype specimen. In addition to overall ex-ternal morphology, T. halli also is distin-guished by the absence of a postfemoralfold (present in T. dankoi and T. vilamen-sis); small maxillary teeth (absent in T.dankoi and T. vilamensis); round, thicktongue (nearly ovoid in T. vilamensis); andsmooth skin (spiny in T. dankoi).

Body size distinguishes T. vilamensis(SVL 38.36–50.81 mm) from the larger T.philippii (SVL 41.49–53.70 mm) and thesmaller T. fronteriensis (SVL 31.62–43.20mm). Telmatobius vilamensis further dif-fers from these taxa by having a postfe-moral fold (absent in T. philippii and T.fronteriensis) and lacking vomerine (pre-sent but reduced in T. philippii and T.fronteriensis), premaxillary, and maxillaryteeth (present in T. philippii and T. fron-teriensis). Telmatobius vilamensis differsfrom T. marmoratus by having a smoothdorsum and head (granular in T. marmor-atus) and in the absence of premaxillaryand maxillary teeth (present in T. marmor-atus). Telmatobius peruvianus, T. pefauri,and T. zapahuirensis have nuptial spineson the chest, throat, and ventral surfacesof the arms; this character is absent in T.vilamensis. Telmatobius vilamensis differsfrom the northernmost species (T. peru-vianus, T. marmoratus, T. pefauri, T. za-pahuirensis; Fig. 2) by lacking premaxillaryand maxillary teeth (present in the north-ernmost species). Telmatobius huayra(Lavilla and Ergueta, 1995a), a Bolivianspecies only 150 km distant from T. vila-mensis, differs from T. vilamensis by hav-ing a well developed postocular fold andsubcylindrical body; T. vilamensis addition-ally has a postfemoral fold (absent in T.huayra).

Description of holotype.—Adult male(SVL 5 46.95 mm); head large, depressed,narrower than body; head length 30.5% ofSVL; head broader than long (head width/


FIG. 2.—Morphological details of the male holotype (IZUA 3080) of Telmatobius vilamensis. Lateral (A),dorsal (B), and ventral (C) views of head; nuptial asperities (D); palmar (E) and plantar surfaces (F); andpostfemoral folds (G).


SVL 5 0.33; head length/head width 50.97). Snout subovoid in dorsal view (Fig.2B); margin of upper jaw flared; snoutstrongly depressed in lateral view (Fig.2A); loreal region flat; nostrils slightlyprominent, oriented anterodorsolaterally;internarial distance 24.3% head width; in-ternarial region flat; nostrils located ap-proximately midway between margin ofeye and anterior terminus of snout; can-thus rostralis indistinct, straight in dorsalview, elevated in lateral view. Eye moder-ately large (30.9% head length), orientedanterodorsolaterally; tympanum, tympanicannulus, and columella absent; supratym-panic fold moderately developed, extend-ing posteroventrally from posterior cornerof eyelid, terminating dorsal to forelimb;lips thin; maxillary and premaxillary teethabsent; dentigerous processes of vomersabsent; choanae large (50.2% internarialdistance) and subcircular; tongue ovoid,elongate, with posterior border free, un-notched, attached through approximately80% of its length anteriorly.

Forelimbs thin, with dermal antebran-chial fold; dorsal surfaces of wrist and sec-ond fingers granular (Fig. 2D); ventral sur-faces of arm and forearm with minutewhite spines; Finger I slightly longer thanFinger II; relative lengths of digits on fore-limb: III . IV . I . II; palmar webbingabsent; tips of fingers rounded; lateralfringes present on Fingers II and III; in-ner palmar tubercle large, depressed andfusiform; diameter of outer metacarpal tu-bercle about 66.6% inner tubercle, ovoid,not depressed; subarticular tubercles sim-ple, rounded, scarcely protruding; twosubarticular tubercles on digits III and IV,one on digits I and II; numerous super-numerary tubercles present (Fig. 2E);nuptial pad present on inner surface ofpollex, consisting of a weakly cornifiedplate with numerous, small, conical, darkspines (Fig. 2D).

Hind limbs long (approximately 158.2%SVL) and slender (Fig. 2G); toes long,thin; length of toes: IV . V 2 III . II .I; webbing formula: I (1½— 2½) II (1½—2) III (1—3½) IV (3½—1½) V; webbingdiminishing distally to form wide fringesalong lateral margins of toes (Fig. 2F); out-

er border of Toe V moderately fringed; in-terdigital plantar webbing with smoothmargins; tips of toes rounded, slightlysmaller than tips of fingers; inner metatar-sal tubercle small, subelliptical and prom-inent; outer metatarsal tubercle small,rounded, and prominent, about 20% sizeof inner tubercle; subarticular tuberclesrounded, small, raised; subarticular tuber-cle formula: I (1), II (1), III (2), IV (3), V(2); supernumerary plantar tubercles few,small; tarsal fold distinct, extending ap-proximately 66.6% length of tarsus, contin-uous distally with fringe along inner mar-gin of Toe I (Fig. 2F); postfemoral foldwell developed (Fig. 2G).

Skin smooth dorsally; skin of ventergranular with numerous, transparent, mi-nute spines; throat smooth (Fig. 2C); clo-acal opening directed posteriorly at dorsallevel of thighs; opening round and orna-mented with folds and papillae; ventralsurface of postfemoral fold with smallgranules and minute transparent spines;ventral skin of thigh loose, folded. Mea-surements of holotype and paratypes arepresented in Table 1.

Color in preservative.—Dorsal surfacesof body, upper arms, and legs dark gray;scattered, distinct, dark brown spots ondorsum; minute spots on dorsal surfaces ofarms and legs; venter and throat cream;ventral surfaces of legs with small, irregu-lar white spots.

Color in life.—Dorsum dark green withdark brown spots; venter and throat white.

Osteology.—Prootic ossification limitedto anteromedial part of otic capsule and apair of small dorsal ossifications associatedwith the lateral margins of the epiotic em-inence. Dorsum of braincase with pairedfontanelles (Fig. 3A), an anterior oval fron-tal fontanelle separated from single, small,round parietal fontanelle by taenia tectitransversalis. Rostral cartilages nearly en-tirely exposed with a pair of small, slendernasals located laterally and adjacent tomargin of tectum nasi and anterior to pla-num antorbitale on each side; vomers en-tirely absent; neopalatines present as short,slender bones underlying middle part pla-num antorbitale on each side of cranium.Minute septomaxillae visible anteriorly.


TA

BL

E1.

—M

orph

omet

ric

data

(in

mm

)fo

rth

ety

pese

ries

ofTe

lmat

obiu

svi

lam

ensi

san

dT.

halli

,pr

ovid

ing

mea

n6

1SD

and

rang

e.

Var

iabl

es

Telm

atob

ius

halli

Hol

otyp

eA

MN

HA

-447

53fe

mal

e

Par

atyp

eA

MN

HA

-447

54fe

mal

e

Par

atyp

eA

MN

HA

-447

58ju

veni

le

Telm

atob

ius

vila

men

sis

Hol

otyp

eIZ

UA

3080

mal

e

Par

atyp

es

Mal

e(n

59)

Fem

ale

(n5

11)

Juve

nile

SVL

Hea

dle

ngth

Hea

dw

idth

57.0

6

16.1

5

18.7

5

48.0

4

14.2

7

16.5

8

33.0

2

9.86

12.3

4

46.9

5

15.5

0

15.5

1

48.0

06

1.72

(44.

73–5

0.81

)12

.96

60.

63(1

2.05

–13.

89)

16.2

96

0.66

(15.

50–1

7.54

)

44.5

06

3.20

(38.

36–4

8.21

)12

.27

60.

86(1

1.05

–13.

36)

15.0

86

1.02

(12.

96–1

6.36

)

31.1

9

10.5

11.3

1

Inte

rorb

ital

dist

ance

Inte

rnar

ial

dist

ance

Eye

–nos

tril

dist

ance

6.04

3.65

3.65

4.91

3.03

3.31

3.16

2.43

1.91

4.79

3.24

3.20

4.87

60.

27(4

.47–

5.32

)3.

406

0.33

(2.7

9–3.

81)

3.40

60.

19(3

.16–

3.73

)

4.68

60.

32(4

.27–

5.26

)3.

226

0.22

(2.9

2–3.

53)

3.25

60.

16(3

.05–

3.54

)

3.72

2.32

2.55

Eye

diam

eter

Han

dle

ngth

Rad

ioul

nar

leng

th

4.17

12.2

0

12.1

1

4.67

11.9

8

10.1

4

3.25

7.78

6.46

4.65

8.57

10.7

9

4.19

60.

34(3

.74–

4.85

)10

.34

60.

97(8

.18–

11.4

1)10

.33

60.

59(9

.32–

11.2

0)

3.97

60.

19(3

.72–

4.34

)10

.03

60.

78(8

.56–

11.2

2)9.

126

1.04

(7.1

9–10

.36)

3.40

7.69

8.58

Tib

iale

ngth

Foo

tle

ngth

24.0

3

40.2

1

20.2

6

32.2

7

24.0

3

23.2

1

19.4

9

33.1

3

21.6

26

1.48

(19.

01–2

3.54

)34

.31

62.

10(3

0.46

–36.

62)

20.0

66

1.34

(17.

48–2

1.77

)32

.51

62.

57(2

8.71

–36.

83)

14.1

1

22.0

2


FIG. 3.—Skull of Telmatobius vilamensis, IZUA 3224, in dorsal (A), ventral (B) and lateral (C) views.Pectoral girdle (D) and hyoid plate (E) of T. vilamensis.

Frontoparietals well developed with dis-tinct laminae perpendicularis; acuminateanteriorly and overlying dorsolateral brain-case from anterior part of orbit posteriorlyto angle of epiotic eminence; frontoparie-tals with broad medial separation except atposterior margin where the pair is fused;frontoparietals investing in medial marginsof prootics and anteromedial margins of

exoccipitals. Parasphenoid robust, nearlytriangular; alae short, broad, and associat-ed with prootic anteriorly and exoccipitalsposteriorly; posteromedial process trun-cate; cultriform process underlying most oforbital region of braincase and formingventral margin of prootic foramen (Fig.3B). Maxillary arcade composed of robustpremaxillae with massive partes dentalis


anteriorly, slender maxillae with low partesfacialis, and lacking preorbital processes,quadratojugals short, not articulating withmaxillae; maxillae edentate. Pterygoidsslender, lacking flanges; anterior ramusterminating medial to pterygoid cartilagein anterior part of orbit; posterior ramusinvesting medial surface of massive pala-toquadrate cartilage; medial ramus short,terminating on anteroventral margin ofotic capsule and not in contact with pro-otic or parasphenoid bones. Squamosals T-shaped in lateral (Fig. 3C) aspect withslender otic and zygomatic rami aboutequal in length; otic ramus located alongdorsolateral margin of cartilaginous cristaparotica; ventral ramus distinctly curved(posteriorly concave), investing lateral sur-face of palatoquadrate cartilage and dis-tinctly separated from quadratojugal. Jawsmoderately short with jaw articulation ly-ing anterior to level of fenestra ovalis.Operculum absent. Stapes and tympanicannulus absent.

Hyoid corpus broad, about 50% broaderthan length at midline; hyoglossal sinusbroad, shallow; hyalia simple, uniform inshape, and lacking processes; anterolateralprocesses of hyoid short, slender, arcuate,directed anterolaterally; posterolateralprocesses short, slender, arcuate, directedposterolaterally; posteromedial processesossified, robust, broadly separated fromone another anteriorly, terminating in car-tilage posteriorly (Fig. 3E).

Pectoral girdle arciferal and robust (Fig.3D). Clavicles stocky, well ossified, appar-ently incorporating ossification of procor-acoid cartilages laterally; transverse axis ofclavicular region straight, anterior marginof bone arcuate. Coracoids short, stocky;sternal end expanded, about twice widthof glenoid end; width of glenoid headabout 50% greater than midshaft width ofcoracoid. Pectoral fenestra shallow, morethan twice as broad as long, medial half ofanterior margin and medial margin carti-laginous, posterior and anterolateral mar-gins formed by coracoid and clavicle, re-spectively. Epicoracoid cartilages well de-veloped; anterior termini anterior to me-dial end of clavicles and broadly separatingthese bones. Omosternum well developed

with cartilaginous manubrium and termi-nal ossification. Sternum short and broad,mostly ossified with cartilaginous margins.Scapula short, robust, only slightly longerthan coracoid; distinctly bicapitate, partesacromialis and glenoidalis approximatelyequal in size; pars acromialis much smallerthan lateral head of clavicle. Suprascapulanot broadly expanded; anterior third ofblade ossified as cleithrum; posterior mar-gin ossified.

Vertebral column composing eight, pro-coelus, nonimbricate, independent presa-cral vertebrae (Fig. 4A,B); vertebrae shortand broad, overall width of neural archabout three times width of centrum, cen-trum width approximately equal to cen-trum length. Presacral I (atlas) moderatelywidely separated, shallow cervical cotyles;presacrals bearing neural spines; relativelengths of transverse processes and sa-crum: II , sacrum , V 5 VI 5 VII 5VIII , III 5 IV; transverse process of Pre-sacral II distally expanded, oriented an-terolaterally; transverse processes of Pre-sacral III arcuate, oriented laterally; trans-verse processes of Presacral IV slightly ex-panded distally, with slight posterolateralorientation; orientation of transverse pro-cesses of Presacrals V–VIII approximatelyuniform with those of V oriented postero-laterally, those of Presacrals VI–VII ori-ented laterally. Sacral diapophyses round,not dilated, oriented posterolaterally; sa-crum with bicondylar articulation withurostyle. Urostyle robust, bearing dorsalcrest that is best developed anteriorly,flanked by lateral flanges of bone that di-minish in size posteriorly.

Overall length of pelvic girdle approxi-mately equal to length of sacrum plus pre-sacral vertebral column. Ilial shaft robust,bearing dorsolateral crest, with low, later-ally oriented dorsal prominence; interilialprofile a narrow U-shape, width of U atbase about half the distance between an-terior ends of ilia; ilium forming anteriormargin of round acetabulum; preacetabu-lum forming approximately a 908 angle toilial shaft; ilia articulating with one anothermedially, forming posterior margin of ac-etabulum; ventral margin of acetabulum


FIG. 4.—Vertebral column of Telmatobius vilamensis in dorsal (A) and ventral (B) views. Lateral view ofpelvic girdle (C), carpus (D), and tarsus (E) of T. vilamensis. Abbreviations: u 5 ulnare; r 5 radiale; Y 5Element-Y; pp 5 prepollex; 2 5 Distal Carpal 2; 5–3 5 Distal Carpal 5–3.


formed by slightly ossified cartilaginouspubis (Fig. 4C).

Humerus longer, more robust than ra-dioulna; humeral crests well developed,especially crista medialis; phalangeal for-mula of manus: 2-2-3-3; terminal phalan-ges pointed; seven well ossified carpal el-ements present: prepollex (two elements),Distal Carpal 2 independent, Distal Car-pals 3–5 fused, Element-Y, radiale, ulnareindependent (Fig. 4D); femur and tibiasimilar in length; tibiale and fibulare fusedproximally and distally, half length of fe-mur; phalangeal formula of the foot: 2-2-3-4-3; prehallux with two elements; twoadditional metatarsal bones of unresolvedhomology (Fig. 4E).

Description of tadpoles.—Larvae (Stag-es 35–40) large (total length 77.71–78.15mm; body length 31.79–34.06 mm) androbust; tail relatively short (1.5 3 bodylength) and thick; body ovoid in lateralview (Fig. 5A); lateral profile of tip ofsnout gently rounded; nostrils ovoid, notprotuberant; apertures situated anterodor-solaterally, with cutaneous fringe (Fig. 5B);internarial distance 150% interocular dis-tance; nostrils closer to anterior border ofeye than tip of snout; eyes circular, locatedin shallow depression; situated anterodor-solaterally; in life, iris with melanophoresand golden tints; width of oral disc 120%interocular distance; oral disc anteroven-tral, transangular; rostral gap present andmental gap absent (Fig. 5C); single row ofmarginal papillae present along peripheryof disc, marginal papillae absent in rostralregion; one row of intramarginal papillae(n 5 13–16) in mental area; papillae pre-sent in infra-angular and supra-angular re-gions 14–17, respectively; rostrodonts wid-er than tall well keratinized; suprarostro-donts and infrarostrodonts with serrationsand dark brown pigmentation; keratodontsformula [(1) (1-1)/(1-1) (2)]; spiraculartube sinistral, short, situated laterally; ap-erture oval, diameter 110% eye diameter;proctodeal tube triangular, opaque (in 10%formalin), wide; vent opening dextral, notvisible in ventral position; distal end ovoid(Fig. 5D); dorsal fin not extending ontobody; ventral fin begins at end of procto-deal tube; tail tip rounded; maximum

width of dorsal and ventral fins slightlyposterior of midlength of tail; fin depth ex-ceeding body depth; myomeres and finswith irregular, dark brown spots; body darkbrown (formalin 10%); internal organs notvisible; hind limbs with minute melano-phores.

The tadpole of T. vilamensis is general-ized (Orton, 1953), having characters as-sociated with anuran larvae inhabiting loticand benthic habitats (e.g., depressed body,anteroventral oral disc, robust caudal mus-culature, dorsolateral eyes, and low fins).Measurements of tadpoles are given in Ta-ble 2.

Chromosomes.—Examination of 19metaphases from two males revealed adiploid number of 2N 5 26. All chromo-somes are bi-armed and the fundamentalnumber (NF) is 52. Pairs 1, 4, 8–13 aremetacentric; Pairs 2, 6, and 7 are sub-metacentric; and Pairs 3 and 5 are subtel-ocentric (Fig. 6A). Pair 6 (sm) has a sec-ondary constriction in the smaller arm.Pairs 1–4 are large (.100 units); Pairs 5and 6 are intermediate (between 80 and100 units), and Pairs 7–13 are small (,80units). A summary of the relative lengths,arm ratios, and types of chromosomes ispresented in Table 3.

The C-band karyotype, based on fiveplates, shows constitutive heterochromatinin the pericentromeric region of all chro-mosomes, except in Pair 4, which showsextensive positive staining in the telomericends of the long arms (Fig. 6B). Thin het-erochromatic bands can be discerned atsome telomeres, especially on the shortarms of Pairs 1, 2, and 7. A thin interstitialband was observed on the long arm of Pair5. The Ag-NOR were revealed in thesmaller arm of Pair 6 (Fig. 6C).

Distribution and ecology.—This speciesis only known from the type locality, theintersection of the Rıo Vilama and theroad between San Pedro de Atacama andEl Tatio (6.5 km by road from San Pedrode Atacama; Fig. 1). The type locality is asemidesert area with scarce vegetation(Ephedra andina and Atriplex atacamen-sis), located in the tropical marginal region(Di Castri, 1968). In San Pedro de Ata-cama, the annual mean temperature is


FIG. 5.—Tadpole of Telmatobius vilamensis (Stage 35). Lateral (A), dorsal (B), and ventral (D) views; oraldisc (C).

13.3 C and rainfall is very scarce (Hajekand Di Castri, 1975). Adults and tadpoles(Stages 34–40) were collected with a netbelow aquatic plants on the banks of theRıo Vilama (5–7 m wide, 10–12 C) duringthe austral autumn (11 April 1997). Malespecimens had nuptial asperities, and onetadpole was near the end of metamorpho-sis (Stage 46, SVL 5 32.0 mm). Bufo spi-

nulosus and lizards of the genus Liolaemuswere also present.

Etymology.—The name vilamensis is aLatin adjective and refers to the type lo-cality, Rıo Vilama.

Telmatobius halli NobleTelmatobius halli Noble, 1938:1 [Holo-

type: AMNH A-44753 from a warm spring


TABLE 2.—Measurements (mm) of the tadpoles ofTelmatobius vilamensis, providing mean 6 1 SD and

range.

CharacterStage 35(n 5 3)

Stage 40(n 5 1)

Total length

Body length

Body depth

80.69 6 2.98(77.71–83.67)32.49 6 0.70(31.79–33.19)16.76 6 0.99(15.77–17.75)

78.15

34.06

16.58

Fin depth

Snout–nostril distance

8.89 6 0.90(7.50–9.89)6.64 6 0.53(6.11–7.17)

9.73

6.29

Eye diameter

Mouth width

2.80 6 0.16(2.64–2.96)7.31 6 0.26(7.05–7.57)

2.64

7.82

near Ollague, 10,000 ft. altitude, ProvinciaEl Loa, Chile].

Diagnosis.—A large Telmatobius (46.95–57.06 mm) having the following combina-tion of characters: (1) premaxillary teeth ab-sent, maxillary teeth rudimentary; (4)tongue round, thick; with posterior borderfree, unnotched; slightly long than wider;attached through more than 75% of itslength; (5) choanae large, subrectangular;(6) tympanum and tympanic annulus ab-sent; (7) snout truncate in dorsal view; (8)postfemoral folds absent; (9) dorsum yellowcream (in preservative); (12) toes extensive-ly webbed; (13) outer border of Toe Vwidely fringed.

Redescription of holotype.—Adult fe-male (SVL 5 57.06 mm); head narrowerthan body; head length 28.6% of SVL;head wider than long (head width/SVL 50.33; head length/head width 5 0.86).Snout truncate in dorsal view (Fig. 7B);snout moderately short in lateral view (Fig.7A); loreal region nearly horizontal; snoutslightly truncate in lateral view; nostrils notprotuberant, oriented anterodorsolaterally;internarial distance 19.6% head width; in-ternarial region convex; nostrils closer totip of snout than eye; canthus rostralis in-distinct in dorsal profile. Eye moderatelylarge (25.5% head length), oriented anter-odorsolaterally; tympanum and tympanicannulus absent; supratympanic fold barelyvisible; lips thin; rudimentary maxillary

teeth present, completely embedded with-in labial mucosa; dentigerous process ofvomers absent; choanae large (60.3% in-ternarial distance) and subrectangular;tongue thick, round, with posterior borderfree, unnotched, slightly longer than wide,attached through more than 75% of itslength anteriorly.

Forelimbs thin, with dermal forearmfold; dermal wrist fold absent; Finger Ilonger than Finger II; relative lengths ofdigits of forelimb: III . IV . I . II; pal-mar webbing absent (Fig. 7C); tips of fin-gers round; rudimentary lateral fringes be-tween Fingers I and II, and II and III;inner palmar tubercle oval, distinct; di-ameter of outer metacarpal tubercle about66.6% inner tubercle, pear-shaped, slightlyraised, distinct; one round distinct subar-ticular tubercle distally on Finger IV; in-distinct distal subarticular tubercle on Fin-gers II and III; distinct subarticular tuber-cles proximally on each finger; one distinctsupernumerary tubercle present at base ofFingers II and III. Three or more indis-tinct supernumerary tubercles present atbase of each finger; supernumerary palmartubercles also present along outer edge ofinner metacarpal tubercle (Fig. 7C).

Hind limb, long (approximately 150.4%SVL) and robust; length of toes: IV . III. V . II . I (Fig. 7D); webbing formulaI (1⅓—2) II (1⅔—3) III (2—3½) IV(3⅔—1½) V; webbing diminishes distallyto form wide fringes along lateral marginsof toes; outer border of Toe V widelyfringed from base to tip; tips of toes spher-ical, about equal in size to finger tips; innermetatarsal tubercle elongate, raised; outermetatarsal tubercle round, small, raised,about half size of inner tubercle; subarti-cular tubercles round, some indistinct, dis-tributed on toes as follows: I (1), II (1), III(6), IV (8), V (4); single, proximal tubercledistinct on Toes II and V; two proximal in-distinct supernumerary tubercles on ToesIV and III; indistinct supernumerary tu-bercles around outer metatarsal tubercle(Fig. 7D); tarsal fold distinct, extendingapproximately 66.6% length of tarsus, con-tinuous distally with fringe along innermargin of Toe I; postfemoral fold absent.

Skin smooth dorsally and ventrally, with-


FIG. 6.—Chromosomes of Telmatobius vilamensis. Standard (A) and C-banded (B) karyotypes. Ag-NOR(C).

out spines or warts; cloacal opening at up-per level of thigh; opening round orna-mented with folds; transverse fold of skinpresent dorsal to the cloacal opening; ven-tral skin of thigh loose, folded, pustular.Measurements of the holotype and para-types are presented in Table 1.

Color in preservative.—Dorsum of bodyuniform brown to tan; paler minute speck-ling on flanks; ventral surfaces of forearmwith white speckles; venter of body uni-

form pale cream; tips of fingers and toesyellow; hand and foot tubercles lightcream. Color in life unknown.

Distribution and ecology.—Telmatobiushalli is only known from the type locality(near Ollague; 218 139 S, 688 169 W; 3700m; El Loa Province, Region de Antofagas-ta, Chile). The type series (five adult fe-males and six tadpoles) was collected froma warm spring in June (Noble, 1938). Dur-ing two consecutive periods (March and


TABLE 3.—Relative length, arm ratio (mean and stan-dard deviations), and type of chromosomes of Tel-

matobius vilamensis.

Pair Relative length* r† Type‡

12345

178.07 6 14.52144.38 6 10.67124.55 6 8.43115.70 6 4.0597.18 6 4.94

1.22 6 0.132.17 6 0.844.08 6 0.521.46 6 0.243.58 6 0.61

msmstmst

6§789

60.22 6 4.6851.85 6 1.3447.57 6 1.7743.50 6 3.19

2.12 6 0.471.72 6 0.351.61 6 0.341.64 6 0.28

smsmmm

10111213

41.42 6 2.0238.50 6 2.5933.92 6 2.6227.50 6 2.47

1.15 6 0.081.32 6 0.061.43 6 0.321.38 6 0.18

mmmm

* Calculated according to Bogart (1970).† Ratio of short arm divided into long arm.‡ Chromosomic types (m 5 metacentric, r 5 1.0–1.7; sm 5 submeta-

centric, r 5 1.71–3.0; st 5 subtelocentric, r 5 3.01–4.5) according toLevan et al. (1964).

§ Pair with secondary constriction.

FIG. 7.—Morphological details of the female ho-lotype (AMNH A-44753) of Telmatobius halli. Lateral(A) and dorsal (B) views of the head. Palmar (C) andplantar (D) surfaces.

December 1998), we searched for addi-tional specimens at the type locality. How-ever, no T. halli were collected. Ollague issituated in the Andean Region (Di Castri,1968); the annual mean temperature ofthis region is 6.8 C and the rainfall rangesfrom 0.0 mm (October) to 31.8 mm (Jan-uary).

DISCUSSION

Species of Telmatobius are character-ized by a significant amount of inter- andintraspecific morphological variation (Cei,1986; Trueb, 1979; Wiens, 1993) that pre-cludes taxonomic arrangement based onunique character states in each species.

Discordant patterns of variation of thefew informative external and osteologicalcharacters have resulted in the progressiverejection of each synapomorphy proposedfor the genus. Thus, the monophyly of Tel-matobius is suspect (see the extended dis-cussion in Formas et al., 1999). As sug-gested by Wiens (1993), generic allocationis based on two character states (fronto-parietals fused posteriorly, and nuptial ex-crescences on Finger I only; both presentin T. vilamensis) that, although not univer-sal, are present in most Telmatobius as wellas in the type for the genus (T. peruvianusWiegmann, 1835).

At this point, the generic assignment of

T. vilamensis would seem questionable;nevertheless, it should be noted that zoo-geographical evidence, as well as allozymeand morphometric data, support the affin-ity of T. vilamensis to other species cur-rently referred to Telmatobius species ofthe Bolivian Altiplano and the AntofagastaRegion (Benavides et al., 2002b).

Telmatobius vilamensis, like other Tel-matobius, has 26 bi-armed chromosomes(Kuramoto, 1990). This 26-chromosomekaryotype (NF 5 52) was considered to beancestral for telmatobiine frogs by Formasand Cuevas (2000) and Reig (1972), butBrum-Zorrilla and Saez (1968) reported 22chromosomes in an unidentified specimenof the Telmatobius marmoratus groupfrom La Paz (Bolivia). This exceptionalkaryotype should be verified, because allother Telmatobius species have a symple-siomorphic arrangement (2N 5 26; seeBarbieri, 1954; Cordova et al., 1987; Cue-vas and Formas, 2002; Dıaz and Veloso,


1979; Formas et al., 1999; Morescalchi,1973; Veloso and Trueb, 1976). It is alsopossible that the 2N 5 22 karyotype is asynapomorphy for a particular clade of Tel-matobius.

On a comparative basis, the standardkaryotype of Telmatobius species is uni-form, but the C-band karyotypes revealsome specific differences. For instance, T.vilamensis has an interstitial heterochro-matic band in Pair 5, whereas, in T. are-quipensis, the short arm of Pair 13 is to-tally heterochromatic (Cordova et al.,1987). Telmatobius dankoi exhibits pro-nounced pericentromeric bands in Pairs 1and 9 (Formas et al., 1999), and T. philip-pii lacks pericentromeric heterochromatinin Pair 1 (Cuevas and Formas, 2002). Inall cases analyzed, the secondary (nucleo-lar) constriction is always located in theshort arm of Pair 6. The number and dis-tribution of heterochromatic bands mayoffer additional phylogenetically informa-tive characters for future studies of thegroup.

Finally, it is worth noting that T. vila-mensis, T. dankoi, T. halli, T. philippii, andT. fronteriensis are known only from theirrespective type localities. In the case of T.halli, no additional specimens have beenreported since the species was first col-lected in 1935 (Noble, 1938). The restrict-ed distributions of these species can be ex-plained by the aridity of the western slopesof the Andes, which effectively separatesbodies of water as reduced independentdrainages in an area of less than 250 km2

total. Geographic isolation of small popu-lations could have produced the surprisingspecies diversity and unusual degree ofdifferentiation shown by this group of Tel-matobius. However, the evolutionary con-sequence of small populations is loss ofheterozygosity (Hartl and Clark, 1997).Benavides et al. (2002b) screened levels ofallozyme diversity in T. vilamensis and T.fronteriensis and found values of H0 50.0058 and H0 5 0.00, respectively. Re-cently, Montgomery et al. (2000) experi-mentally confirmed a positive correlationbetween allozyme diversity and populationsize; thus, it is plausible that these speciesare endangered.

KEY TO ADULT TELMATOBIUS OF CHILE

1. Snout rounded or pointed in dorsal view. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Snout truncated in dorsal view;tongue rounded, unnotched; attachedanteriorly along 75% of its length; pre-maxillary teeth absent, maxillary teethreduced; dorsum uniform brown totan (in preservative); belly pale cream;tips of toes and fingers yellow . . T. halli

2. Tibio tarsal join does not reach the pos-terior border of the eye . . . . . . . . . . . 3Tibio tarsal join reaches or exceedsthe posterior border of eye . . . . . . . . 4

3. Skin on head strongly granular; nuptialspines extensive on dorsolateral sur-face of thumb; irregularly distributedspines on chest, inner metacarsal tu-bercle, lips, and ventral surface ofarm; tongue attached anteriorly about66% of its length; vomerine teeth re-duced; skin on dorsum, belly, flanksand dorsal surfaces of legs stronglygranular; dorsum gray (in preserva-tive) with irregular dark spots (mar-morations) . . . . . . . . . . . . T. marmoratusSkin on head without granules; nuptialspines reduced, not reaching innermetacarpal tubercle; tongue attachedanteriorly for about 80% of its length;skin on flanks strongly granular; dor-sum dark gray (in preservative) withdark brown irregular spots; belly giltcopper (in life) . . . . . . . . . . . T. philippii

4. Xiphisternum notched . . . . . . . . . . . . . . 5Xiphisternum unnotched; tympanicring absent; nuptial spines extensiveon chest, throat, and dorsolateral andventral surfaces of thumb, reachingthe inner metacarpal tubercle; tongueattached 66% of its length; vomerineteeth absent; dorsum brown to gray(in preservative) . . . . . . . . T. peruvianus

5. Flanks without queratinous spines . . . . 6Flanks with minute, brown, queratin-ous spines (males and females); nup-tial pad extending along dorsolateralsurface of thumb; minute dark spinesirregularly distributed on throat, anddorsal and ventral surfaces of hand;postfemoral folds present; craniumwell ossified; dorsum gray to brown(in preservative) . . . . . . . . . . . T. dankoi

6. Snout not depressed . . . . . . . . . . . . . . . 7Snout strongly depressed (wedge-shaped); without vomerine, premaxil-lary, and maxillary teeth; nuptial


spines small, restricted on dorsal andlateral surfaces of thumb; postfemoralfolds present (in preservative and life);cranium poorly ossified; dorsum gray(in preservative), olive-green (in life). . . . . . . . . . . . . . . . . . . . . . T. vilamensis

7. Subarticular tubercles present on all toes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

Subarticular tubercles present on ToesI, II, and III; nuptial spines extensiveon chest and dorsolateral surfaces ofthumb; tongue attached anteriorly85% of its length; dorsum light brown(in preservative) . . . . . . T. zapahuirensis

8. Adult size .54 mm; nuptial spines ex-tensive on chest, throat, anterior bor-der of arm, and dorsoventral surfacesof thumb, reaching the inner metacar-pal tubercle; tongue attached 66% ofits length; dorsum light brown (in pre-servative) with minute gray irregularlydistributed spots . . . . . . . . . . . T. pefauriAdult size ,45 mm; nuptial spineslarge, black, extensive on dorsolateralsurface of thumb; nuptial pad contin-uous with inner metacarpal tubercle;throat and ventral surface of thighsand arms with leaden (in preservative)irregular spots . . . . . . . . . T. fronteriensis

RESUMEN

Describimos una nueva especie; Telma-tobius vilamensis del Rıo Vilama, cerca deSan Pedro de Atacama, Chile. Previamen-te confundida con T. halli Noble, 1938,esta nueva especie es descrita en base acaracteres de la morfologıa externa de laslarvas y adultos, osteologıa y cromosomas.La nueva especie es el representante massureno del genero Telmatobius en Chile.Adicionalmente, se redescribe T. halli y secomparan ambos taxa con las restantes es-pecies del genero reportadas para Chile.

Acknowledgments.—We thank L. Trueb, J. Sites,and three anonymous referees for suggestions thatgreatly improved the manuscript; C. Cole (AMNH),J. Aparicio (CBF), and J. C. Ortiz (MZUC) for loanof specimens; and N. Bernal, C. Mehling, and R.Bain (AMNH) for pictures of the T. halli holotype.We also thank C. Jara for field assistance, M. Navarrofor art work, and L. Brieva for drafting versions ofthe manuscript. This study was supported by FondoNacional de Ciencia y Tecnologıa (ProyectoS960021).

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Accepted: 5 June 2002Associate Editor: Joseph Mendelson III

APPENDIX ISpecimens Examined

Telmatobius dankoi.—CHILE: Provincia El Loa:Las Cascadas, 2260 m, MNHN 3006, IZUA 2108–10,2112 (two cleared-and-stained adults), 2113 (ninespecimens), 2106 (two tadpoles), 2107 (five tadpoles),DBMUA 45, 46, 53, 57.

Telmatobius halli.—CHILE: Provincia El Loa:Ollague, 3050 m, AMNH A-44753–54, A-44758.

Telmatobius marmoratus.—CHILE: Provincia Pa-rinacota: Lago Chungara, 4270 m, MZUC 24520,24524, 24526. BOLIVIA: Provincia Murillo: Valle deLa Paz, CBF 03244–56, CBF 1009–11, CBF 468–70.

Telmatobius pefauri.—CHILE: Provincia Parina-cota: Quebrada de Zapahuira, 3270 m, DBCUCH629, 632–634, 641, 652.

Telmatobius peruvianus.—CHILE: Provincia Pa-rinacota: Putre, 3200 m, MZUC 24642–43.

Telmatobius vilamensis.—CHILE: Provincia ElLoa: Rio Vilama, 3110 m, IZUA 3080–81, IZUA 3224(cleared-and-stained adult), IZUA 3146 (two cleared-and-stained adults), IZUA 3225–26, IZUA 3132 (fivetadpoles); CBF 3760–62; MZUC 24104–06, MZUC25725–38.

Telmatobius zapahuirensis.—CHILE: Provincia


Parinacota: Quebrada de Zapahuira, 3270 m, DBCG630, 639.

Telmatobius philippii—CHILE: Provincia El Loa:Quebrada de Amincha, 3700 m, IZUA 3093, 3087;Provincia El Loa: Quebrada del Inca, 3700 m, IZUA3088–92, 3193–95 (three cleared-and stained speci-

mens, chromosomes), 3094 (seven tadpoles), 3196–97 (chromosomes).

Telmatobius fronteriensis.—CHILE: Provincia ElLoa: Puquios, 4150 m, MZUC 25094, 25095–103,25261–78, 25095, and 25103 (cleared-and stainedadults).

Documents

A NEW SPECIES OF TELMATOBIUS (ANURA: LEPTODACTYLIDAE) … · 253 Herpetologica, 59(2), 2003, 253–270 q 2003 by The Herpetologists’ League, Inc. A NEW SPECIES OF TELMATOBIUS (ANURA: