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

August 23, 2017 | Autor: Cesar Garcia Cuevas | Categoria: Zoology, Taxonomy, Biodiversity, Chromosomes
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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, Chile e-mail: [email protected] l Abstract. A new species of frog, Alsodes hugoi, from the temperate Nothofagus forest of Central Chile, is described. 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 de Chile Central. Hasta hoy no se ha encontrado otra especie congenérica en la localidad típica. Desde el punto de vista 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 and Southern Chile and along the eastern slopes of the Andes in Argentina (south of Mendoza City). The following species are recognized: A. australis Formas, Ubeda, Cuevas and Núñez, 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. vanzolinii Donoso-Barros, 1974 (Formas, 1981); A. verrucosus Philippi, 1902 (Cei, 1976) and A. 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 of the genus Telmalsodes. Wiens (1993) did not agree with this proposition and argued that both genera are synonymous. The only character that signiŽ catively differentiates the two genera is the presence of plantar webbing in Telmalsodes, but this character also occurs in © Koninklijke Brill NV, Leiden, 2001

Amphibia-Reptilia 22: 187-19 8

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Alsodes gargola (Cei, 1980) and A. tumultuosus (Formas et al., 1997). The most remarkable characteristic of the males of the genus Alsodes is the presence of thorny structures on the Ž 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 pectoral girdle is pseudoŽ misternal (Barrio, 1970); in Alsodes species the pectoral girdle is arciferal. While collecting frogs in the temperate Nothofagus forest in Central Chile, a new species of Alsodes was detected. This taxon is described here, including the tadpole and the karyotype, 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). All the measurements (mm) were taken with digital callipers to the nearest 0.1 mm. Specimens were measured according to Cei (1962). The following data were recorded: snout-vent length (SVL); head length; head width; thigh length; tibia length; foot length and nostrilsnout distance. Internarial distance was measured according to Cei (1980). Eye diameter was measured according to Duellman (1970). Tadpoles were staged according to Gosner (1960) and the following measurements were taken according to Formas (1992): total length; body length; body depth; Ž n depth; snout-eye distance; snout-nostril distance; eye diameter and mouth width. The karyotype was described from two males (IZUA 3191, 3192). The specimens were injected with 0.5% colchicine for four hours, and then anaesthetised with diethyl ether, cut open ventrally under sterile conditions and the intestine carefully removed. Metaphase plates were obtained by squashing intestinal epithelium fragments. These were treated hypotonically with distilled water, then Ž xed in acetic alcohol (1 3), and Ž nally placed in 45% acetic acid. Small fragments of tissue were squashed between glass slides and cover slips and dipped in liquid nitrogen. The cover slip was removed with a razor blade and the chromosomes were allowed to air dry. After one day 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 were determined according to Levan et al. (1964). The relative length was determined according to Bogart (1970). Microscopic slides and specimens were deposited in the Amphibian Collection of the Institute of Zoology, Universidad Austral de Chile (IZUA).

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Results Alsodes hugoi sp. nov. (Ž gs. 1, 3) Holotype. IZUA specimen 3119 is an adult male collected by Lila Brieva, John Valladares 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 (Ž g. 2). Paratypes. IZUA 3120 (male), 3118 and 3121 (females). These were collected at the same site and time as the holotype. Diagnosis. The following characteristics have been considered to include the new species into the genus Alsodes: males characterized by having thorny structures on the Ž ngers and round, bilateral spiny patches on the chest; pectoral girdle arciferal. Alsodes hugoi can be distinguished from its congeners by the following combination of characters: dorsum light brown; arms and legs light brown and black-barred; a lateral fold of the skin extending between the posterior border of the arm, and the middle part of the body; iris brilliant

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

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

copper in colour; a black triangular zone converging backwards on the head between the eyes; heel reaching the middle of the eye when hind leg is bent forward; tadpole with dorsal aperture of the spiracular tube. Although both A. hugoi and A. nodosus have the dorsal surface of arms and legs black-barred, they can be differentiated because the latter lacks the lateral fold of the skin. From the karyological point of view both species can be clearly differentiated because A. nodosus possesses 22 chromosomes (Bogart, 1970) whilst A. hugoi has 26. Alsodes hugoi, A. tumultuosus and A. kaweshkari have a distinctive lateral fold of the skin; however, their length and thickness are different. In A. kaweshkari the lateral fold of the skin is thick and folding, reaching the groin; in A. hugoi it is thin and does not reach the groin; in A. tumultuosus it is also thin, but extending to the groin. 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-vent length. The snout is slightly rounded in lateral view and dorsal proŽ le (Ž gs. 3A, B). The loreal region is slightly rounded in cross section. The nostrils are antero-lateral in position,

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

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

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are present on the vomer between the choanae. Forelimbs are robust (Ž gs. 1 and 3A, B). Fingers are in order of increasing length: I — II — IV and III (Ž gs. 3D, E). Webbing is absent on the hand, and the Ž ngers have rounded tips. The inner palmar tubercle is ovoid and  attened. The outer palmar tubercle is triangular,  attened, and slightly bigger than the inner one. Palmar and subarticular tubercles are present (Ž g. 3D). The dorsal surface and internal border of the Ž rst Ž nger has strong thorny excrecences, brown in colour (Ž gs. 3D, E). The internal border of the second Ž nger has a narrow band of smaller spines (Ž g. 3E). The chest has two bilateral whitish patches of small spines (Ž g. 3D). The toes are long, thick and fringed and in order of decreasing length I — II — III — V and IV with rounded tips. Webbing is absent. Between the fourth and Ž fth toes, there is a reduced membrane. The inner metatarsal tubercle is oval and rounded and larger than the outer one (Ž g. 3C). A tarsal fold is present, but reduced. The ventral, dorsal and lateral surfaces of the thighs have minute granules. A fold of skin extends between the posterior border of the arm, and the middle part of the body (Ž g. 3B). The measurements of the holotype and the paratypes are shown in table 1. Pectoral girdle arciferal and the xiphisternum slightly expanded and deeply notched (IZUA 3120, 3191). Colouration. This is based on the type series. In alcohol, the holotype (IZUA 3119) and one paratype (IZUA 3120) are dark grey dorsally with a grey venter. The ventral areas of the anterior parts of the thighs are whitish and the granular regions dark. The throat is dark 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 arms are black-barred. In life, dorsum, arms and legs are light brown with golden tints. The bars on the legs and arms, and the spots on the dorsum are black. The venter is whitish. The upper part of the iris is golden yellow (Ž g. 1). Juvenile. One juvenile (IZUA 3189, collected at the type locality) exhibits the same characteristics as the adults; however, some differences were observed. The dorsum,  anks, and dorsal surfaces of the legs are covered with minute granules. The postocular fold reaches the groin. Measurements of this specimen are shown in table 1. Tadpole description. Thirteen larvae of A. hugoi, stages 25-40 (Gosner, 1960) were collected at the type locality close to the type series. We assigned these tadpoles to A. hugoi based on distribution, as the only Alsodes species collected here was A. hugoi. In the type locality, we also captured larvae and adults of Telmatobufo venustus, but their tadpoles are easily distinguishable. Telmatobufo venustus tadpoles (Díaz et al., 1983) show remarkable morphological adaptations to mountain streams, such as broad, sucker-like mouths, and wide, 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 snout is round in contour (Ž 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

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

Snout-vent length Head length Head width Thigh length Tibia length Foot length Eye diameter Snout-nostril distance Internarial distance

Holotype

Paratypes

Juvenile

IZUA 3119

IZUA 3120

IZUA 3118

IZUA 3121

IZUA 3189

69.0 22.4 23.8 35.7 33.2 51.8 7.6 4.9 5.8

68.4 22.6 23.4 34.5 33.4 50.6 7.8 4.3 5.9

65.9 24.1 24.1 29.1 33.2 49.8 8.0 4.3 5.9

70.7 25.7 23.0 29.5 32.0 49.8 7.3 3.5 5.1

30.4 10.9 12.2 14.5 13.9 19.3 4.1 2.7 3.3

Table 2. Alsodes hugoi sp. nov.: morphometric data (mean § s; ranges) of the tadpoles (all data in mm). Character Total length Body length Body depth Fin depth Snout-eye distance Snout-nostril distance Eye diameter Mouth diameter

Stage 25 n D 11

Stage 40 nD2

37:9 § 3:60 15:9 § 1:59 6:5 § 0:89 6:4 § 0:62 5:4 § 0:34 3:3 § 0:40 1:4 § 0:16 5:0 § 0:68

62.4-61.3 26.5-26.2 9.9-9.6 9.2-8.3 8.8-8.0 5.8-5.0 2.5-2.4 8.3-8.3

distance (Ž g. 4B). Pupils are circular with a golden iris spotted by black reticulations (live specimens). The oral disc is translucent, in an anteroventral position. The width of the disc is 1.4 times the interocular distance. The oral disc is intraangular, with a rostral gap present and a mental gap absent (Ž g. 4C). A line of marginal papillae is seen along the periphery of the disc, except in the rostral region. Scarce intramarginal papillae are found in the mental area. Three to four papillae are found at supra-angular regions. Rostrodonts are wider than they are high, and are well keratinized. Suprarostrodont s and infrarostrodonts possess serrations. Keratodont formula is normal (1) (1-1)/(1-1) (2) (Ž g. 4C). The spiracular tube is sinistral and laterally located, and the aperture is oval with a well-deŽ ned border, visible on dorsal view (Ž g. 4A, B). The proctodeal tube is transparent, wide, and the vent opening dextrally with the distal opening diameter shorter than the internarial distance (Ž gs. 4B, D). Tail length is 1.5 times the body length, with a straight axis. The dorsal Ž n begins in a  eshy crest on the posterior third of the body. The ventral Ž n begins at the end of the proctodeal tube. The tail tip is rounded. Fin depth does not exceed the body depth (Ž g. 4A). Dorsal and ventral Ž ns are transparent with numerous minute melanophores. The dorsal surface appears 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 (Ž g. 4D).

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

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 biarmed chromosomes (Ž g. 5). The fundamental number (FN) was 52. When chromosomes are arranged in pairs of decreasing length, pairs 1-3 are large (> 100 units), pairs 4-6 are intermediate (between 80 to 100 units), and 7-13 small (< 80 units). A remarkable secondary constriction can be observed on the short arm of one homologue of pair 1. The chromosome measurements are shown in table 3.

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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 D metacentric, sm D submetacentric, st D subtelocentric. a Chromosome s with secondary constriction. Relative length was calculated according to Bogart (1970). Arm ratio D long arm / short arm. Pair

Relative length

Ratio

Type

1a

147:50 § 11:92 122:10 § 6:33 103:90 § 4:25 98:15 § 6:57 91:83 § 6:20 85:31 § 3:84 64:78 § 4:42 61:48 § 3:47 54:96 § 3:08 50:16 § 2:42 43:50 § 9:84 39:61 § 9:41 31:96 § 9:28

1:27 § 0:11 3:83 § 0:13 2:00 § 0:77 1:32 § 0:09 5:98 § 0:79 4:94 § 1:77 1:84 § 0:15 1:28 § 0:20 1:71 § 0:12 1:56 § 0:10 1:35 § 0:12 1:22 § 0:18 1:53 § 0:44

m st sm m st st sm m sm m m m m

2 3 4 5 6 7 8 9 10 11 12 13

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

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Lircay river (5-7 m wide, 10-12 ±C) during the southern summer (January 1998). Adults and tadpoles were collected in the water, among the rocks. The juvenile was captured out of the water, under a stone close to the river. The holotype (IZUA 3119) and paratype (IZUA 3120) had nuptial asperities on Ž ngers and chest and two females (paratypes IZUA 3118-3121 ) presented mature oocites (2.4-2.6 mm diameter) with the animal pole black and the vegetal pole creamy-white. Etymology. This new species is named in memory of Professor Dr. Hugo Campos Cereceda, for his remarkable contributions to the development of the Natural Sciences in Chile.

Discussion Among the leptodactylid frogs, the species of the genus Alsodes are characterized by an arciferal 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 larval characteristics. The morphological features of adults and tadpoles of Alsodes hugoi are appropriate reasons to include this species in the genus Alsodes. Cei (1980) considered that two morphologically different evolutionary lines exist in the genus Alsodes. This results in one stock having a rather simple horny ornamentation on the thumb pad (e.g. A. nodosus and A. tumultuosus), and another showing a more complex spiny structure on the Ž rst Ž ngers (e.g. A. australis, A. kaweshkari) (Formas et al., 1997, 1998). The characteristics of the nuptial pads of A. hugoi suggest that this species can be included in the stock with complex spiny ornamentation. Formas and Vera (1983), based on chromosome number, recognized three groups within the genus: the barrioi group (2n D 34); the monticola group (2n D 26) and the nodosus group (2n D 22). The presence of 26 chromosomes in A. hugoi allows its inclusion in the monticola group. The maximum SVL (70.7 mm) of Alsodes hugoi is larger than that of the other congeneric species. Among the larger frogs, the external morphology of the foot and chromosome numbers provides useful characteristics to identify each taxon. One example of this is A. nodosus, a species characterized by the absence of lateral fringes (present on the toes in A. barrioi, A. hugoi, A. kaweshkari, and A. tumultuosus) and a karyotype with 22 chromosomes (Formas and Vera, 1983). While foot morphology of A. hugoi and A. barrioi are similar, these taxa can be distinguished because A. barrioi has 34 chromosomes (Formas and Vera, 1983) and A. hugoi has 26. Alsodes hugoi differs from A. australis, A. kaweshkari, and A. tumultuosus in the development of the tarsal fold, 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 tadpole of A. hugoi is characteristic of this species and it is an appropriate feature to distinguish this larva from the tadpoles of other congeneric species.

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Specimens examined Abbreviations CU: 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 verrucosu s: 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. Valladares who helped us in the Ž 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 de Cuyo, Argentina), and Carmen Ubeda (Universidad del Comahue, Argentina) kindly provided specimens for comparisons . Marcos Navarro expertly prepared the drawings. The photograph of the paratype was taken by René Navarro. This work was supported by Fondo Nacional de Ciencia y Tecnología (FONDECYT), Project 1000426.

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Duellman, W.E. (1970): The hylid frogs of the Middle America. Monogr. Mus. Nat. Hist. Univ. Kansas 1: 1-753. Duméril, A.M., Bibron, G. (1841): Erpétologie générale, ou histoire naturelle des Reptiles. Vol. 8, p. 1-784. París, Impr. de Fain. Formas, J.R. (1981): The identity of the frog Eupsophus vanzolinii from Ramadillas, Nahuelbuta Range, southern Chile. Proc. Biol. Soc. Wash. 93: 920-927. Formas, J.R. (1989): Sinonimia e identidad de la rana austral Chilena Eupsophus vittatus (Philippi, 1902) (Anura, Leptodactylidae) . Bol. Soc. Biol. Concepción 60: 123-127. Formas, J.R. (1992): The tadpole of Eupsophus vertebralis (Anura: Leptodactylidae) . Herpetologica 48: 115-119. Formas, J.R., Cuevas, C.C., Núñez, J. (1998): A new species of Alsodes (Amphibia: Anura: Leptodactylidae ) from Southern Chile. Proc. Biol. Soc. Wash. 111: 521-530. Formas, J.R., Ubeda, C., Cuevas, C.C., Núñez, J. (1997): Alsodes australis, a new species of leptodactylid frog from the temperate Nothofagus forest of Southern Chile and Argentina. Stud. Neotrop. Fauna Environm. 32: 200-211. Formas, J.R., Vera, M.I. (1983): Karyological relationships among frogs of the genus Alsodes, with description of the karyotype s of A. vanzolinii and A. verrucosus. Copeia 1983: 1101-1107 . Gallardo, J.M. (1970): A propósito de los Telmatobiinae (Anura, Leptodactylidae ) patagónicos. Neotropica 16: 73-85. Gosner, K.L. (1960): A simpliŽ ed table for staging anuran embryos and larvae with notes on identiŽ cation. Herpetologica 16: 183-190. Lavilla, E.O. (1988): Lower Telmatobiinae (Anura: Leptodactylidae) : generic diagnoses based on larval characters. Occas. Papers Mus. Nat. Hist. Univ. Kansas 124: 1-19. Levan, A., Fredga, A., Sandberg, A. (1964): Nomenclature for centromeric positions on chromosomes . Hereditas 52: 201-220. Lynch, J.D. (1978): A re-assessment of the telmatobiine leptodactylid frogs of Patagonia. Occas. Papers Mus. Nat. Hist. Univ. Kansas 72: 1-57. Philippi, R.A. (1902): Suplemento de los Batraquios Chilenos descritos en la Historia Física y Política de Chile de don Claudio Gay. Santiago de Chile, Librería José Ivens. Veloso, A.M., Díaz, N.P., Iturra, P., Penna, M. (1981): Descripción de una nueva especie de telmatobino del género Alsodes (Amphibia, Leptodactylidae ) de la Cordillera de Nahuelbuta (Sur de Chile). Med. Amb. 51: 72-77. Veloso, A.M., Iturra, P., Galleguillos, G. (1979): Evidencias cromosómicas en el género Alsodes (Amphibia, Leptodactylidae) , con la descripción de una nueva especie. Physis 94: 91-98. Veloso, A., Sallaberry, M., Navarro, J., Iturra, P., Valencia, J., Penna, M., Díaz, N. (1982): Contribución sistemática al conocimiento de la herpetofaun a del extremo norte de Chile. En: El hombre y los ecosistemas de montaña, p. 1-268. Montevideo, Uruguay, Impreso por la OŽ cina Regional de Ciencia y Tecnología de la Unesco para América Latina y el Caribe (ROSTLAC). Wiens, J.J. (1993): Systematic of the Leptodactylid frog genus Telmatobius in the Andes of northern Perú. Occas. Papers Mus. Nat. Hist. Univ. Kansas 162: 1-76.

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

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