A new species of Alsodes (Amphibia, Anura, Leptodactylidae) from Central Chile

A new species of Alsodes (Amphibia, Anura, Leptodactylidae)from Central Chile

César C. Cuevas, J. Ramón Formas

Instituto de Zoología, Casilla 567, Universidad Austral de Chile, Valdivia, Chilee-mail: [email protected] l

Abstract. A new species of frog, Alsodes hugoi, from the temperate Nothofagus forest of Central Chile, isdescribed. Up to now, no sympatric occurrence of other congeneric species is known from the type locality.Karyotypically, this new species is included in the monticola group, whose species have 26 chromosomes .

Resumen. Una nueva especie de rana, Alsodes hugoi, es descrita para el bosque temperado de Nothofagus deChile Central. Hasta hoy no se ha encontrado otra especie congenérica en la localidad típica. Desde el punto devista cariológico, la nueva especie es incluida en el grupo monticola cuyas especies tienen 26 cromosomas.

Introduction

Frogs of the genus Alsodes Bell 1843 (Gallardo, 1970) are distributed in Central andSouthern Chile and along the eastern slopes of the Andes in Argentina (south of MendozaCity). The following species are recognized: A. australis Formas, Ubeda, Cuevas andNúñez, 1997; A. barrioi Veloso, Díaz, Iturra and Penna, 1981; A. gargola Gallardo,1970; A. illotus Barbour, 1922 (Cei, 1980); A. kaweshkari Formas, Cuevas and Núñez,1998; A. laevis Philippi, 1902 (Lynch, 1978); A. montanus Lataste, 1897 (Veloso et al.,1982); A. monticola Bell, 1843; A. nodosus Duméril and Bibron, 1841 (Gallardo, 1970);A. pehuenche Cei, 1976; A. tumultuosus Veloso, Iturra and Galleguillos, 1979; A. vanzoliniiDonoso-Barros, 1974 (Formas, 1981); A. verrucosus Philippi, 1902 (Cei, 1976) andA. vittatus Philippi, 1902 (Formas, 1989). The taxonomic status of A. illotus is not clear(Cei, 1980), and A. laevis has not been collected since its description by Philippi (1902).Alsodes montanus and A. pehuenche have been considered by Díaz (1989) as members ofthe genus Telmalsodes. Wiens (1993) did not agree with this proposition and argued thatboth genera are synonymous. The only character that signi� catively differentiates the twogenera is the presence of plantar webbing in Telmalsodes, but this character also occurs in

© Koninklijke Brill NV, Leiden, 2001 Amphibia-Reptilia 22: 187-198

188 C.C. Cuevas, J.R. Formas

Alsodes gargola (Cei, 1980) and A. tumultuosus (Formas et al., 1997). The most remarkablecharacteristic of the males of the genus Alsodes is the presence of thorny structures onthe � ngers, and round spiny patches on the chest (Formas et al., 1998). Insuetophrynus(I. acarpicus) also shows paired patches of spines on the chest of males, but its pectoralgirdle is pseudo� misternal (Barrio, 1970); in Alsodes species the pectoral girdle is arciferal.

While collecting frogs in the temperate Nothofagus forest in Central Chile, a new speciesof Alsodes was detected. This taxon is described here, including the tadpole and thekaryotype, and compared with other congeneric species.

Materials and methods

The description of the new species is based on four adults (two males and two females)from Central Chile [IZUA (Instituto de Zoología Universidad Austral de Chile) 3118, 3119,3120, 3121], one juvenile and thirteen tadpoles from the same locality (IZUA 3131). Allthe measurements (mm) were taken with digital callipers to the nearest 0.1 mm. Specimenswere measured according to Cei (1962). The following data were recorded: snout-ventlength (SVL); head length; head width; thigh length; tibia length; foot length and nostril-snout distance. Internarial distance was measured according to Cei (1980). Eye diameterwas measured according to Duellman (1970). Tadpoles were staged according to Gosner(1960) and the following measurements were taken according to Formas (1992): totallength; body length; body depth; � n depth; snout-eye distance; snout-nostril distance;eye diameter and mouth width. The karyotype was described from two males (IZUA3191, 3192). The specimens were injected with 0.5% colchicine for four hours, andthen anaesthetised with diethyl ether, cut open ventrally under sterile conditions andthe intestine carefully removed. Metaphase plates were obtained by squashing intestinalepithelium fragments. These were treated hypotonically with distilled water, then � xed inacetic alcohol (1 3), and � nally placed in 45% acetic acid. Small fragments of tissue weresquashed between glass slides and cover slips and dipped in liquid nitrogen. The cover slipwas removed with a razor blade and the chromosomes were allowed to air dry. After oneday the chromosomes were stained for 15 min in Sorensen’s phosphate buffer (pH 6.8),containing 4% Giemsa solution (Cuevas and Formas, 1996). Centromeric positions weredetermined according to Levan et al. (1964). The relative length was determined accordingto Bogart (1970). Microscopic slides and specimens were deposited in the AmphibianCollection of the Institute of Zoology, Universidad Austral de Chile (IZUA).

A new species of Alsodes 189

Results

Alsodes hugoi sp. nov. (� gs. 1, 3)

Holotype. IZUA specimen 3119 is an adult male collected by Lila Brieva, John Val-ladares and César Cuevas on 29 January 1998 at Río Lircay, Altos de Vilches (35°280S,71°110W, 900 m altitude), 66 km E of Talca (by road), Talca Province, Región del Maule,on the western slopes of the Andes (� g. 2).

Paratypes. IZUA 3120 (male), 3118 and 3121 (females). These were collected at thesame site and time as the holotype.

Diagnosis. The following characteristics have been considered to include the new speciesinto the genus Alsodes: males characterized by having thorny structures on the � ngers andround, bilateral spiny patches on the chest; pectoral girdle arciferal. Alsodes hugoi can bedistinguished from its congeners by the following combination of characters: dorsum lightbrown; arms and legs light brown and black-barred; a lateral fold of the skin extendingbetween the posterior border of the arm, and the middle part of the body; iris brilliant

Figure 1. Alsodes hugoi sp. nov. (paratype 3118).


Figure 2. Type locality of Alsodes hugoi sp. nov.

copper in colour; a black triangular zone converging backwards on the head between theeyes; heel reaching the middle of the eye when hind leg is bent forward; tadpole with dorsalaperture of the spiracular tube.

Although both A. hugoi and A. nodosus have the dorsal surface of arms and legsblack-barred, they can be differentiated because the latter lacks the lateral fold of theskin. From the karyological point of view both species can be clearly differentiatedbecause A. nodosus possesses 22 chromosomes (Bogart, 1970) whilst A. hugoi has26. Alsodes hugoi, A. tumultuosus and A. kaweshkari have a distinctive lateral foldof the skin; however, their length and thickness are different. In A. kaweshkari thelateral fold of the skin is thick and folding, reaching the groin; in A. hugoi it is thinand does not reach the groin; in A. tumultuosus it is also thin, but extending to thegroin.

Description of the Holotype. The body is robust and the arms and legs are well developed.The head is shallow, slightly wider than it is long, with a length 28% of the snout-ventlength. The snout is slightly rounded in lateral view and dorsal pro� le (� gs. 3A, B). Theloreal region is slightly rounded in cross section. The nostrils are antero-lateral in position,


Figure 3. Alsodes hugoi sp. nov., morphologica l details of the holotype. (A) Antero-lateral view, (B) antero-dorsalview, (C) plantar view of the left foot. Sexual secondary characters. (D) Ventral view of the chest and palmar viewof the left hand, (E) dorsal view of the left hand.

nearer the tip of the snout than the anterior margin of the eye (� g. 3A). The eye diameteris 1.3 times the internarial distance. A tympanum is absent. A postocular fold is evident,reaching the middle of the body (� g. 3B). The head has minute granules, especially onthe supraocular areas and under the postocular fold. The tongue is rounded, lacking anotch at the tip (paratype IZUA 3120). Choanae are rounded. Dentigerous processes (6-7)


are present on the vomer between the choanae. Forelimbs are robust (� gs. 1 and 3A, B).Fingers are in order of increasing length: I — II — IV and III (� gs. 3D, E). Webbing isabsent on the hand, and the � ngers have rounded tips. The inner palmar tubercle is ovoidand � attened. The outer palmar tubercle is triangular, � attened, and slightly bigger than theinner one. Palmar and subarticular tubercles are present (� g. 3D). The dorsal surface andinternal border of the � rst � nger has strong thorny excrecences, brown in colour (� gs. 3D,E). The internal border of the second � nger has a narrow band of smaller spines (� g. 3E).The chest has two bilateral whitish patches of small spines (� g. 3D). The toes are long,thick and fringed and in order of decreasing length I — II — III — V and IV with roundedtips. Webbing is absent. Between the fourth and � fth toes, there is a reduced membrane.The inner metatarsal tubercle is oval and rounded and larger than the outer one (� g. 3C).A tarsal fold is present, but reduced. The ventral, dorsal and lateral surfaces of the thighshave minute granules. A fold of skin extends between the posterior border of the arm, andthe middle part of the body (� g. 3B). The measurements of the holotype and the paratypesare shown in table 1. Pectoral girdle arciferal and the xiphisternum slightly expanded anddeeply notched (IZUA 3120, 3191).

Colouration. This is based on the type series. In alcohol, the holotype (IZUA 3119) andone paratype (IZUA 3120) are dark grey dorsally with a grey venter. The ventral areasof the anterior parts of the thighs are whitish and the granular regions dark. The throat isdark grey. Two paratypes (IZUA 3118, 3121) are grey dorsally with irregular black spots.A black triangular zone converges backwards on the head between the eyes. The venter,throat and ventral surface of the thighs are whitish. The dorsal surfaces of the legs and armsare black-barred. In life, dorsum, arms and legs are light brown with golden tints. The barson the legs and arms, and the spots on the dorsum are black. The venter is whitish. Theupper part of the iris is golden yellow (� g. 1).

Juvenile. One juvenile (IZUA 3189, collected at the type locality) exhibits the samecharacteristics as the adults; however, some differences were observed. The dorsum, � anks,and dorsal surfaces of the legs are covered with minute granules. The postocular foldreaches the groin. Measurements of this specimen are shown in table 1.

Tadpole description. Thirteen larvae of A. hugoi, stages 25-40 (Gosner, 1960) werecollected at the type locality close to the type series. We assigned these tadpoles to A. hugoibased on distribution, as the only Alsodes species collected here was A. hugoi. In the typelocality, we also captured larvae and adults of Telmatobufo venustus, but their tadpoles areeasily distinguishable. Telmatobufo venustus tadpoles (Díaz et al., 1983) show remarkablemorphological adaptations to mountain streams, such as broad, sucker-like mouths, andwide, strong tails. The measurements of the tadpoles of A. hugoi are shown in table 2.Larvae in stage 25 have ovoid bodies in lateral view, 2.6 times longer than deep. The snoutis round in contour (� g. 4A). Nostrils are small, situated between the eye and snout tip.The eyes are antero-dorsolateral and the interocular distance is similar to the internarial


Table 1. Alsodes hugoi sp. nov.: morphometric data of the type series and a juvenile (all data in mm).

Character Holotype Paratypes Juvenile

IZUA 3119 IZUA 3120 IZUA 3118 IZUA 3121 IZUA 3189

Snout-vent length 69.0 68.4 65.9 70.7 30.4Head length 22.4 22.6 24.1 25.7 10.9Head width 23.8 23.4 24.1 23.0 12.2Thigh length 35.7 34.5 29.1 29.5 14.5Tibia length 33.2 33.4 33.2 32.0 13.9Foot length 51.8 50.6 49.8 49.8 19.3Eye diameter 7.6 7.8 8.0 7.3 4.1Snout-nostril distance 4.9 4.3 4.3 3.5 2.7Internarial distance 5.8 5.9 5.9 5.1 3.3

Table 2. Alsodes hugoi sp. nov.: morphometric data (mean § s; ranges) of the tadpoles (all data in mm).

Character Stage 25 Stage 40n D 11 n D 2

Total length 37:9 § 3:60 62.4-61.3Body length 15:9 § 1:59 26.5-26.2Body depth 6:5 § 0:89 9.9-9.6Fin depth 6:4 § 0:62 9.2-8.3Snout-eye distance 5:4 § 0:34 8.8-8.0Snout-nostril distance 3:3 § 0:40 5.8-5.0Eye diameter 1:4 § 0:16 2.5-2.4Mouth diameter 5:0 § 0:68 8.3-8.3

distance (� g. 4B). Pupils are circular with a golden iris spotted by black reticulations (livespecimens). The oral disc is translucent, in an anteroventral position. The width of the discis 1.4 times the interocular distance. The oral disc is intraangular, with a rostral gap presentand a mental gap absent (� g. 4C). A line of marginal papillae is seen along the periphery ofthe disc, except in the rostral region. Scarce intramarginal papillae are found in the mentalarea. Three to four papillae are found at supra-angular regions. Rostrodonts are widerthan they are high, and are well keratinized. Suprarostrodont s and infrarostrodonts possessserrations. Keratodont formula is normal (1) (1-1)/(1-1) (2) (� g. 4C). The spiracular tubeis sinistral and laterally located, and the aperture is oval with a well-de� ned border, visibleon dorsal view (� g. 4A, B). The proctodeal tube is transparent, wide, and the vent openingdextrally with the distal opening diameter shorter than the internarial distance (� gs. 4B, D).Tail length is 1.5 times the body length, with a straight axis. The dorsal � n begins in a � eshycrest on the posterior third of the body. The ventral � n begins at the end of the proctodealtube. The tail tip is rounded. Fin depth does not exceed the body depth (� g. 4A). Dorsaland ventral � ns are transparent with numerous minute melanophores. The dorsal surfaceappears grey in formalin but dark brown in life. The venter is transparent with dark spots.Two ventral whitish spots are present near the proctodeal tube (� g. 4D).


Figure 4. Tadpole of Alsodes hugoi sp. nov. (stage 25). (A) Lateral view, (B) dorsal view, (C) oral disc, (D) ventralview.

Chromosomes. The examination of 15 metaphase plates from two male specimens(IZUA 3191, 3192), collected in the type locality, showed a karyotype formed by 26 bi-armed chromosomes (� g. 5). The fundamental number (FN) was 52. When chromosomesare arranged in pairs of decreasing length, pairs 1-3 are large (> 100 units), pairs 4-6are intermediate (between 80 to 100 units), and 7-13 small (< 80 units). A remarkablesecondary constriction can be observed on the short arm of one homologue of pair 1. Thechromosome measurements are shown in table 3.


Figure 5. Karyotype of Alsodes hugoi. Arrow shows the secondary constriction.

Table 3. Alsodes hugoi sp. nov.: karyotype data (mean § s) of two male specimens (IZUA 3191, 3192). m Dmetacentric, sm D submetacentric, st D subtelocentric. aChromosomes with secondary constriction. Relativelength was calculated according to Bogart (1970). Arm ratio D long arm/short arm.

Pair Relative length Ratio Type

1a 147:50 § 11:92 1:27 § 0:11 m2 122:10 § 6:33 3:83 § 0:13 st3 103:90 § 4:25 2:00 § 0:77 sm4 98:15 § 6:57 1:32 § 0:09 m5 91:83 § 6:20 5:98 § 0:79 st6 85:31 § 3:84 4:94 § 1:77 st7 64:78 § 4:42 1:84 § 0:15 sm8 61:48 § 3:47 1:28 § 0:20 m9 54:96 § 3:08 1:71 § 0:12 sm

10 50:16 § 2:42 1:56 § 0:10 m11 43:50 § 9:84 1:35 § 0:12 m12 39:61 § 9:41 1:22 § 0:18 m13 31:96 § 9:28 1:53 § 0:44 m

Distribution and ecology. This species is known only from the type locality (Altos deVilches, 900 m altitude, Talca Province) (� g. 2). This woody area (Nothofagus alpina,N. pumilio, N. glauca, Austrocedrus chilensis, Laurelia sempervirens, Drimys winteri andAextoxicon punctatum) is located on the western slopes of the Andes. The type localityis situated in the Mediterranean subhumid region (di Castri, 1968). The annual meantemperature of this region is 14.5 ±C, the relative humidity is 73%, and the mean annualrainfall is 700 mm. The specimens described here were captured on the banks of the


Lircay river (5-7 m wide, 10-12 ±C) during the southern summer (January 1998). Adultsand tadpoles were collected in the water, among the rocks. The juvenile was captured outof the water, under a stone close to the river. The holotype (IZUA 3119) and paratype(IZUA 3120) had nuptial asperities on � ngers and chest and two females (paratypes IZUA3118-3121) presented mature oocites (2.4-2.6 mm diameter) with the animal pole blackand the vegetal pole creamy-white.

Etymology. This new species is named in memory of Professor Dr. Hugo CamposCereceda, for his remarkable contributions to the development of the Natural Sciencesin Chile.

Discussion

Among the leptodactylid frogs, the species of the genus Alsodes are characterized by anarciferal type pectoral girdle, and strong black thorny excrecences on the thumb and chest(males only) (Cei, 1980). Lavilla (1988) diagnosed the genus Alsodes based on larvalcharacteristics. The morphological features of adults and tadpoles of Alsodes hugoi areappropriate reasons to include this species in the genus Alsodes. Cei (1980) considered thattwo morphologically different evolutionary lines exist in the genus Alsodes. This results inone stock having a rather simple horny ornamentation on the thumb pad (e.g. A. nodosusand A. tumultuosus), and another showing a more complex spiny structure on the � rst� ngers (e.g. A. australis, A. kaweshkari) (Formas et al., 1997, 1998). The characteristicsof the nuptial pads of A. hugoi suggest that this species can be included in the stock withcomplex spiny ornamentation. Formas and Vera (1983), based on chromosome number,recognized three groups within the genus: the barrioi group (2n D 34); the monticolagroup (2n D 26) and the nodosus group (2n D 22). The presence of 26 chromosomes inA. hugoi allows its inclusion in the monticola group.

The maximum SVL (70.7 mm) of Alsodes hugoi is larger than that of the othercongeneric species. Among the larger frogs, the external morphology of the foot andchromosome numbers provides useful characteristics to identify each taxon. One exampleof this is A. nodosus, a species characterized by the absence of lateral fringes (presenton the toes in A. barrioi, A. hugoi, A. kaweshkari, and A. tumultuosus) and a karyotypewith 22 chromosomes (Formas and Vera, 1983). While foot morphology of A. hugoiand A. barrioi are similar, these taxa can be distinguished because A. barrioi has34 chromosomes (Formas and Vera, 1983) and A. hugoi has 26. Alsodes hugoi differsfrom A. australis, A. kaweshkari, and A. tumultuosus in the development of the tarsalfold, which is reduced in A. hugoi and well-developed in these other species (A. australis,A. kaweshkari and A. tumultuosus). The dorsal aperture of the spiracular tube of the tadpoleof A. hugoi is characteristic of this species and it is an appropriate feature to distinguishthis larva from the tadpoles of other congeneric species.


Specimens examined

AbbreviationsCU: Carmen Ubeda (personal collection), Argentina.DBCG: Departamento de Biología Celular y Genética, Universidad de Chile, Chile.IBA: Instituto de Biología Animal, Universidad Nacional de Cuyo, Argentina.IZUA: Instituto de Zoología, Universidad Austral de Chile, Chile.MNHN: Museo Nacional de Historia Natural, Chile.MZUC: Museo de Zoología, Universidad de Concepción, Chile.

Alsodes barrioi: IZUA 1629-1630: Cordillera Pelada, Provincia de Valdivia, 1,020 m, Chile.Alsodes gargola: CU 6: Macizo Loncoluan, Provincia de Neuquén, 1,900 m, Argentina.Alsodes montanus: IZUA 824: Estero Cobarrubias; Provincia de Santiago, 2,400 m, Chile.Alsodes monticola: IZUA 1550-1749 : Cordillera Pelada, Provincia de Valdivia, 1,020 m, Chile.Alsodes nodosus: IZUA 756,700: Aguas Claras, Provincia de Petorca, 150 m, Chile.Alsodes pehuenche: IBA 1643: Valle del Pehuenche, Provincia de Mendoza, 2,500 m, Argentina.Alsodes tumultuosus: DBCG 161-162: La Parva, Provincia de Santiago, 2,600 m, Chile.Alsodes vanzolinii: MZUC 12063-12070 : Ramadillas, Provincia de Arauco, 100 m, Chile.Alsodes verrucosus: MNHN 1506: Puerto Edén, Provincia de Ultima Esperanza, 10 m, Chile.

Acknowledgements. The authors would like to give special thanks to Lila M. Brieva and John P. Valladareswho helped us in the � eldwork. One referee, Prof. Dr. U. Sinsch, gave us useful comments on the manuscript.Alberto Veloso, Patricia Iturra, José Navarro (Universidad de Chile), Enrique Pereira (Universidad Nacional deCuyo, Argentina), and Carmen Ubeda (Universidad del Comahue, Argentina) kindly provided specimens forcomparisons . Marcos Navarro expertly prepared the drawings. The photograph of the paratype was taken byRené Navarro. This work was supported by Fondo Nacional de Ciencia y Tecnología (FONDECYT), Project1000426.

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Received: May 15, 2000. Accepted: September 8, 2000.

A new species of Alsodes (Amphibia, Anura, Leptodactylidae) from Central Chile

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