SHORTER COMMUNICATIONS Journal of Herpetology, Vol. 43, No. 1, pp. 159–164, 2009 Copyright 2009 Society for the Study of Amphibians and Reptiles
Tadpole of Telmatobius mayoloi (Anura: Ceratophryidae) CE´SAR AGUILAR1,2 1
AND
EDGAR LEHR3
Museo de Historia Natural, Departamento de Herpetologı´a, Universidad Nacional Mayor de San Marcos, Av. Arenales 1256, Jesus Marı´a, Ap. 14-0434, Lima, Peru´; E-mail:
[email protected] 3 Staatliche Naturhistorische Sammlungen Dresden, Museum fu¨r Tierkunde, Ko¨nigsbru¨cker Landstrasse 159, D-01109 Dresden, Germany
ABSTRACT.—The tadpole of Telmatobius mayoloi from central Peru (Departamento de Ancash) is described based on specimens ranging from Gosner Stage 28–41. The tadpole of T. mayoloi is similar to other Telmatobius larvae with pond type morphology. Geographically closest to T. mayoloi are Telmatobius carrillae and Telmatobius rimac. The tadpole of T. mayoloi differs from the latter in reaching a maximum total length of 102.1 mm at Stage 38 whereas T. carrillae (maximum total length = 99.20 mm at Stage 39) and T. rimac (maximum total length = 77. 70 mm at Stage 36) are smaller. Furthermore, the snout of the tadpole of T. mayoloi is slightly narrower between the nares, and its tip is pointed compared to the broader snouts of T. carrillae and T. rimac. The tadpole of T. mayoloi is the fourth largest in Telmatobiinae; data comparing tadpole total lengths and adult snout–vent lengths of Peruvian Telmatobiinae are presented. RESUMEN.—Se describe el renacuajo de Telmatobius mayoloi procedente del Peru´ central (Departamento de Ancash) en base a especimenes con Estadı´os de Gosner que van del 28 al 41. El renacuajo de T. mayoloi es similar a las otras larvas de Telmatobius con morfologı´a de tipo poza. Especies geogra´ficamente ma´s cercanas a T. mayoloi son Telmatobius carrillae y Telmatobius rimac. El renacuajo de T. mayoloi se diferencia de estos u´ltimos al alcanzar una longitud total ma´xima de 102.1 mm en el Estadı´o 38 mientras que T. carrillae (longitud total ma´xima = 99.20 mm en el Estadı´o 39) y T. rimac (longitud total ma´xima = 77. 70 mm en el Estadio 36) son ma´s pequen˜os. Adema´s, el hocico del renacuajo de T. mayoloi es ligeramente ma´s angosto al nivel de las narinas y su extremo es puntiagudo en comparacio´n con los hocicos ma´s anchos de T. carrillae y T. rimac. El renacuajo de T. mayoloi es el cuarto ma´s grande dentro de Telmatobiinae y se presentan datos sobre la longitud total de los renacuajos y la longitud hocico-cloaca de los adultos de Telmatobiinae que habitan en Peru´. Frogs of the genus Telmatobius are distributed along the Andes from Ecuador to Chile and Argentina between 1,300 and 5,400 m elevation. They comprise 57 mostly aquatic or semiaquatic species of which 23 (40%) occur in Peru (Cei, 1986; De la Riva et al., 2005; Lehr, 2005; Seimon et al., 2007). Knowledge on reproduction and larval phase is limited, and tadpole descriptions are only available for 10 (43%) of Peruvian species of Telmatobius. The tadpoles that are unknown or not described account for 13 species: Telmatobius arequipensis Vellard, 1955; Telmatobius brevirostris Vellard, 1955; Telmatobius colanensis Wiens, 1993; Telmatobius degener Wiens, 1993; Telmatobius hockingi Salas and Sinsch, 1996; Telmatobius intermedius Vellard, 1951; Telmatobius latirostris Vellard, 1951; Telmatobius mayoloi Salas and Sinsch, 1996, Telmatobius necopinus Wiens, 1993; Telmatobius punctatus Vellard, 1955; Telmatobius sanborni Schmidt 1954; Telmatobius thompsoni Wiens, 1993; and Telmatobius timens De la Riva, Aparicio and Rı´os, 2005 (De la Riva et al., 2005; Lehr, 2005). Telmatobius mayoloi is an aquatic species whose distribution is restricted to the Rı´o Santa and its tributary streams in Departamento de Ancash between 3,515 and 4,050 m elevation (Lehr 2002, 2005; Lehr et al., 2002). Telmatobius mayoloi is an endangered species (IUCN et al., 2006), and both adults and larvae are used locally for food and for traditional medicine (Lehr, 2000). Although the tadpole of T. mayoloi was 2
Corresponding Author.
mentioned in the original species description by Salas and Sinsch (1996), it was never described. In 1997 and 1998, Aguilar and Lehr did extensive fieldwork in the Rı´o Santa Valley, collecting additional specimens and tadpoles of T. mayoloi. Distributional data of T. mayoloi were summarized by Lehr (2002) and Lehr et al. (2002), and Lehr (2005) presented a photo of a live tadpole at Gosner Stage 38 (MUSM 20550) with a total length of 102.1 mm. Description of some features of the oral cavity and chondrocranium are shown in Aguilar (2006). Herein, we describe the tadpole of T. mayoloi. MATERIALS AND METHODS For tadpoles examined see Appendix 1. Tadpoles were found along with adult specimens of T. mayoloi. Tadpoles were fixed in 10% formalin and were staged according to Gosner (1960). The format used for tadpole description follows that of Aguilar et al. (2007a). Terminology of external larval features follows Lavilla (1988) and Altig and Johnston (1989). The following measurements were taken to the nearest 0.1 mm: body length (distance from the tip of the snout to the body terminus, which is the junction of the posterior body wall with the tail axis); tail length (distance from the body terminus to the absolute tip of tail); total length (sum of body length and tail length); body width (measured at the widest point right behind the eyes); eye diameter; interorbital distance (measured between the centers of the pupils); internarial distance (measured between the centers of the nares indicated by less pigmentation when
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FIG. 2. Oral disc of Telmatobius mayoloi (MUSM 20550, Gosner Stage 28). Drawing by E. Lehr.
FIG. 1. Tadpole of Telmatobius mayoloi (MUSM 20550, Gosner Stage 28, total length 57.05 mm) in lateral (A), dorsal (B), and ventral (C) views. Photos by E. Lehr. closed); distance between tip of snout and naris (from center of naris to middle of snout); distance between naris and eye (from the center of naris to the anterior edge of the eye); tail height (greatest); tail muscle height (at its beginning and at tail midlength); tail muscle width (at its beginning); and oral disc width. Measurements taken with an ocular micrometer include eye diameter, interorbital distance, internarial distance, distance between tip of snout and naris, distance between naris and eye, tail muscle height, tail muscle width, and oral disc width; all others were taken with digital calipers. Drawings were made using a stereomicroscope with a camera lucida. Coloration in life description is based on a photo taken by E. Lehr. Museum acronyms are as follows: MUSM 5 Museo de Historia Natural Universidad Nacional Mayor de San Marcos, and SMF 5 Forschungsinstitut und Naturmuseum Senckenberg. RESULTS The description is based on one specimen at Gosner Stage 28 (MUSM 20550, see Figs. 1, 2). For morphometric data of the tadpole series (Gosner stages 28–41) see Table 1. Description.—The tadpole of T. mayoloi belongs to the exotrophic ecomorphological guild, section I, group B7 as defined by Altig and Johnson (1989). At stage 28, the total length is 57.05 mm. Body is slightly depressed (height / body width 5 0.94) and oval in dorsal view. Maximum body width is at about midbody, anterior to the spiracle tube. The snout is pointed in dorsal and lateral views (Fig. 1). Lateral line organs are not visible in dorsal and lateral views. The mouth is located anteroventral and is surrounded by a small oral disc (width of oral disc / body width 5 0.43; Fig. 2). Marginal papillae are interrupted rostrally. One row of intramarginal papillae is present continuously in the suprangular and infrangular but absent in the mental area. Papillae are simple and conical. The suprarostrodont is wider than long and concave. The infrarostrodont is convex laterally and concave medially. Both are black and keratinized with
small triangular serrations. Keratodonts are small and the Labial Tooth Row Formula (LTRF) is 2(2)/3(1). Nares are oval, small, without projections and inflexions and dorsolaterally orientated, closer to the eyes than to the snout (snout–naris distance / naris– eye distance 5 1.54). Eyes are small (eye diameter / body width 5 0.13) and dorsolaterally orientated. The spiracle is single, sinistral, and located in the posterior third of the body (snout–spiracle distance / body length 5 0.63). The spiracular opening is oval and the inner wall is present as a slight ridge. The vent tube is dextral and attached to the lower fin. Fins are concave, and the posterior end is rounded; dorsal fin does not extend into the body. Maximum tail height at about midlength and is as high as body height (maximum height tail / body height 5 0.91). The muscular tail begins at the tail-body junction and posteriorly tapers to a tip. Muscular tail is as high as its width at the tailbody junction (muscular height / width 5 1.00). Color in Preservative.—Dorsal body grayish-tan with brown flecks, more abundant in the posterior half; ventral body translucent, intestine visible; muscular tail with brown flecks concentrated in the anterior half; dorsal fin with smaller brown flecks anteriorly and translucent medially, ventral fin translucent anterior and medially; muscular tail, dorsal and ventral fin with dark brown spots on the third posterior end. Color in Life.—Based on a photo taken by E. Lehr of a specimen at Stage 38 (MHNSM 20550). Body and muscular tail dull brown with greyish-olive flecks; fins translucent with dull brown flecks anteriorly and spots on the posterior third; iris bronze. Variation.—Variation from the above description is noted in the coloration of body and tail. Specimens at Stages 29, 31, 32, 36 (MUSM 20550 series) and Stages 28, 38 (MUSM 7417) have scattered brown flecks on dorsal body and anterior two-thirds of muscular tail. At Stage 38 a specimen (MUSM 7417) has brown flecks on ventral body; a specimen at Stage 28 (MUSM 7417) has brown flecks on the anterior two-thirds of dorsal fin and medially on ventral fin; MUSM 20550 specimens at Stages 29, 31, 32, and 36, and MUSM 7417 specimen at Stage 38 have scattered brown flecks on the first two-thirds of both dorsal and ventral fin. See Table 1 for morphometric data. Habitat.—Tadpoles (MUSM 20550) were found in slow currents in the Rı´o Santa and in its tributaries, between stones and vegetation. It seems that reproduction of T. mayoloi is not seasonal because tadpoles at Stages 28, 38, and a metamorph (Stage 45, SVL 5
Character Body length Tail length Total length Body width Eye diameter Interorbital distance Internarial distance Distance snout–naris Distance naris–eye Distance snout– spiraculum Tail heigth (greatest) Tail muscle height (at its beginning) Tail muscle height (at tail midlength) Tail muscle width (at its beginning) Oral disc width
Lot
Gosner Stage (1960)
21.9 35.1 57.1 12.0 1.5
5.1
3.3
3.7
2.4
13.8
10.3
6.9
5.0
5.0 5.2
22.0 32.0 54.0 11.6 1.4
4.6
3.5
3.7
2.2
13.3
10.1
5.5
4.1
5.5 4.5
28
MUSM 20550
28
MUSM 7417
5.3 4.3
4.5
6.0
9.9
13.6
2.4
3.2
3.4
5.1
22.2 35.2 57.3 10.6 1.5
MUSM 20550
29
5.3 4.3
4.3
5.0
10.1
12.3
2.3
2.4
3.4
5.5
18.4 33.3 51.7 10.7 1.5
MUSM 20550
29
6.0 4.9
5.0
7.3
13.5
16.0
2.5
3.3
4.0
6.5
27.7 40.1 67.8 14.0 1.9
MUSM 20550
31
5.7 5.9
9.0
8.0
13.2
15.0
2.2
4.8
3.2
5.9
24.8 42.9 67.7 11.8 1.9
SMF 80592
31
5.0 6.8
5.7
7.0
11.8
13.6
2.3
5.3
3.4
5.6
22.0 39.1 61.1 14.0 2.4
SMF 80799
31
6.2 6.5
6.0
7.1
13.6
16.7
2.7
4.2
4.2
6.9
28.3 46.2 74.5 15.3 1.9
MUSM 20550
32
34
6.8 5.8
9.0
8.0
12.8
16.2
2.6
4.7
3.7
4.1
26.5 44.7 71.1 12.6 2.1
SMF 80592
TABLE 1. Measurements (in mm) of selected characters of Telmatobius mayoloi tadpoles. 35
7.3 6.9
8.9
8.6
14.7
18.1
2.8
6.2
3.9
4.6
32.2 54.5 86.6 15.7 2.1
SMF 80592
36
8.3 7.7
10.8
9.7
17.0
20.2
3.2
6.0
3.7
7.5
32.2 51.5 83.6 17.9 2.4
SMF 80592
36
6.6 7.1
6.1
8.0
14.3
16.6
3.1
3.4
4.3
7.3
28.1 46.3 74.4 13.4 2.1
MUSM 20550
36
7.5 7.6
7.0
8.5
12.2
19.5
4.1
4.8
3.8
7.0
30.7 43.7 74.4 16.7 1.7
MUSM 20551
38
9.8 8.4
10.9
10.7
18.3
20.5
3.4
6.3
4.4
7.6
34.1 50.2 84.2 18.1 2.4
SMF 80592
38
8.0 8.0
9.0
10.0
19.3
20.5
3.0
4.0
4.8
8.5
34.9 50.7 85.6 20.8 2.1
MUSM 7417
40
6.7 7.6
8.2
7.8
13.8
17.7
2.4
6.8
3.6
6.7
27.9 52.7 80.6 16.7 2.6
SMF 80799
40
7.6 8.2
6.6
8.3
12.3
19.1
4.2
3.8
3.8
7.3
30.3 46.8 77.1 16.7 2.1
MUSM 20551
41
8.6 8.5
10.8
10.0
18.5
21.9
3.3
8.0
4.4
7.7
33.8 46.5 80.3 21.1 3.4
SMF 80594
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TABLE 2. Comparison of maximum snout–vent length (SVL) of adults and total length of tadpoles (in mm) of Peruvian Telmatobiinae. Species
Batrachoahrynus brachydactylus Batrachophrynus macrostomus Telmatobius atahualpai Telmatobius brevipes Vellard, 1951 Telmatobius carrillae Telmatobius culeus (Garman, 1876) Telmatobius ignavus Barbour and Noble, 1920 Telmatobius jelskii (Peters, 1873) Telmatobius marmoratus (Dume´ril and Bibron, 1841) Telmatobius mayoloi Telmatobius peruvianus Wiegmann, 1834 Telmatobius rimac Telmatobius truebae Wiens, 1993
Adult Tadpole
Gosner Stage
Source
66.6
41 40 36 38 39 ? (hind limbs present) 34
Sinsch (1990); Sinsch et al. (1995) Sinsch (1990); Sinsch et al. (1995) Aguilar et al. (2007a) Wiens (1993) Morales (1988); Aguilar (pers. obs.) Benavides et al. (2002); Vellard (1951) Wiens (1993)
71.2
110.0
39
74.6
76.9
36–38
108.1 54.4 86.9
102.1 94.0 77.7
38 36–38 36
82.0
75.2
35
Sinsch (1990); Salas and Sinsch (1996) Benavides et al. (2002); Formas et al. (2005) Lehr (2005) Formas et al. (2005) Sinsch et al. (1995); Aguilar et al. (2007b) Wiens (1993)
71.9 170.3 63.4 71.3 51.0 137.9
89.0 181.0 74.1 73.4 99.2 94.0
78.9
37.70, MUSM 7417) were collected on 9 August 1989 in the dry season and other tadpoles at Stages 28, 29, 31, 32, and 36 (MUSM 20550) were collected on 28 March 1997 at the end of the rainy season. DISCUSSION The tadpole of T. mayoloi is similar to other pond type Peruvian Telmatobius larvae and shares with them the presence of a small oral disc with transangular lateral margins, a rostral gap and intramarginal lateral papillae present in angular area, rostrodonts wider than large, LTRF 5 2(2)/3(1), level of nares aperture not raised, spiracle position sinistral, vent tube present and opening dextrally, and tail fins normal or not reduced (Lavilla, 1988). Telmatobius species geographically closest to T. mayoloi and with a pond type larva are T. carrillae Morales, 1988 and T. rimac Schmidt, 1954. Tadpoles of both species are similar to T. mayoloi but differ in the shape of the snout in dorsal view (Aguilar et al., 2007b). The tadpole of T. mayoloi has a snout which is slightly narrower between the level of the nares and its tip is pointed compared to the broader snouts of T. carrillae and T. rimac. Despite the wider distribution of tadpoles with pond type morphology in Telmatobius, rheophilous tadpoles are present in some species of Bolivia and Peru (Lavilla and De la Riva, 1993; De la Riva, 2005; Aguilar et al., 2007a). So far known, the only rheophilous tadpole of Telmatobius present in Peru is the larva of Telmatobius atahualpai. This tadpole differs from the larva of T. mayoloi in having a wider oral disc, a LTRF 5 3/7(1), and in lacking a rostral gap (Aguilar et al., 2007a). Knowledge of phylogenetic relationships between Telmatobius species with other related genera is partial, in the best of cases. Aguilar and Pacheco (2005) proposed that Batrachophrynus and Telmatobius are closely related based on internal oral features. Moreover, Aguilar (2006) found in a phylogenetic
analysis of Batrachophrynus Peters, 1873, and 15 species of Telmatobius based on adult and larval morphological characters, that T. mayoloi forms a clade with T. carrillae and Batrachophrynus species. The tadpole of T. mayoloi differs from both Batrachophrynus species in having dark brown spots on the posterior third of the tail (Batrachophrynus brachydactylus Peters, 1873, lacks spots on the posterior third and Batrachophrynus macrostomus Peters, 1873, has a uniform dark brown tail [Sinsch, 1986]). Moreover, the tadpole of B. macrostomus is much larger than T. mayoloi at similar stages. At Gosner Stage 36 and 38, total length of T. mayoloi tadpole is 74.38 and 102.1 mm, respectively (Table 1; Lehr, 2005), but in two individuals of B. macrostomus at the same stages, total length is 131.10 and 135.95 mm, respectively (CA, pers. obs.). Within Telmatobiinae, the tadpole of T. mayoloi is the fourth largest preceded by B. macrostomus, Telmatobius jelskii, and Telmatobius gigas (Sinsch, 1990; De la Riva, 2002; Table 2). Although, T. mayoloi is one of the largest Peruvian Telmatobius species, adult size of Telmatobius seems not to parallel tadpole total length (Table 2). For instance, maximum adult size recorded for T. jelskii is 71.20 mm in contrast to 108.1 mm of T. mayoloi (SMF 80500), but at Stage 39 T. jelskii tadpole can reach a total length of 110.00 mm (Sinsch, 1990; Salas and Sinsch, 1996). Further detailed comparisons and a comprehensive phylogenetic analysis are needed to asses the evolution of body size in Telmatobiinae. Acknowledgments.—We thank J. H. Co´rdova for permitting access to specimens and providing a working space, G. Ko¨hler for loan of material. We thank G. R. Smith, I. De la Riva, U. Sinsch, K. Siu Ting and anonymous reviewer for comments on the manuscript. Fieldwork conducted in 1997 was kindly financed by the Forschungsinstitut und Naturmuseum Senckenberg (SMF), Frankfurt, Germany. We thank INRENA for collecting and export permits.
SHORTER COMMUNICATIONS LITERATURE CITED AGUILAR, C. 2006. Relaciones filogene´ticas entre algunos Telmatobinidos (Anura, Leptodactylidae, Telmatobiinae) de Peru´ basado en la morfologı´a de los estados larval y adulto. Unpubl. Master’s thesis, Universidad Nacional Mayor de San Marcos, Lima, Peru. Available at http://www.cybertesis.edu.pe/ sisbib/2006/aguilar_pc/html/index-frames.html. AGUILAR, C., AND V. PACHECO. 2005. Contribucio´n de la morfologı´a bucofarı´ngea larval a la filogenia de Batrachophrynus y Telmatobius. In E. O. Lavilla and I. De la Riva (eds.), Estudios sobre las ranas andinas de los ge´neros Telmatobius y Batrachophrynus (Anura: Leptodactylidae), pp. 219–238. Monografı´as de Herpetologı´a Vol. 7. Asociacio´ n Herpetolo´ gica Espan˜ola, Valencia, Spain. AGUILAR, C., K. SIU TING, AND P. VENEGAS. 2007a. The rheophilous tadpole of Telmatobius atahualpai Wiens, 1993 (Anura: Ceratophryidae). South American Journal of Herpetology 2:165–174. AGUILAR, C., M. LUNDBERG, K. SIU TING, AND M. JIMENEZ. 2007b. Nuevos registros para la herpetofauna del departamento de Lima, descripcio´n del renacuajo de Telmatobius rimac Schmidt, 1954 (Anura: Ceratophrydae) y una clave de los anfibios. Revista Peruana de Biologı´a 14:209–216. ALTIG, R., AND G. F. JOHNSON. 1989. Guilds of anuran larvae: relationships among developmental modes, morphologies, and habitats. Herpetological Monographs 3:81–109. BENAVIDES, E., J. C. ORTIZ, AND J. W. SITES. 2002. Species boundaries among the Telmatobius (Anura: Leptodactylidae) of the Lake Titicaca basin: allozyme and morphological evidence. Herpetologica 58:31–55. CEI, J. M. 1986. Speciation and adaptative radiation in Andean Telmatobius frogs. In F. Vuilleumier and M. Monasterio (eds.), High Altitude Tropical Biogeography, pp. 374–386. Oxford University Press, New York. DE LA RIVA, I. 2002. Rediscovery and taxonomic status of Telmatobius marmoratus gigas Vellard, 1969 ‘‘1968’’ (Anura: Leptodactylidae). Herpetologica 58:220–228. ———. 2005. Synopsis of Bolivian Telmatobius. In E. O. Lavilla and I. De la Riva (eds.), Estudios sobre las ranas andinas de los ge´neros Telmatobius y Batrachophrynus (Anura: Leptodactylidae), pp. 65–101. Monografı´as de Herpetologı´a Vol. 7, Asociacio´n Herpetolo´gia Espan˜ola, Valencia, Spain. DE LA RIVA, I., J. APARICIO, AND N. RIOS. 2005. New Species of Telmatobius (Anura: Leptodactylidae) from Humid Paramo of Peru´ and Bolivia. Journal of Herpetology 39:409–416. FORMAS, R., A. VELOSO, AND J. C. ORTIZ. 2005. Sinopsis de los Telmatobius de Chile. In E. O. Lavilla and I. De la Riva (eds.), Estudios sobre las ranas andinas de los ge´neros Telmatobius y Batrachophrynus (Anura: Leptodactylidae), pp. 103–114. Monografı´as de Herpetologı´a Vol. 7, Asociacio´n Herpetolo´gica Espan˜ola, Valencia, Spain. GOSNER, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpetologica 16:183–190.
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IUCN, CONSERVATION INTERNATIONAL, AND NATURESERVE. 2006. Global Amphibian Assessment [Internet]. Accessed 2 March 2008. Available at www. globalamphibians.org. LAVILLA, E. O. 1988. Lower Telmatobiinae (Anura: Leptodactylidae): generic diagnosis based on larval characters. Occasional Papers, Museum of Natural History, University of Kansas, Lawrence 124:1–19. LAVILLA, E. O., AND DE LA RIVA. 1993. La larva de Telmatobius bolivianus (Anura: Leptodactylidae). Alytes 11:37–46. LEHR, E. 2000. Zur Nutzung einiger Amphibien- und Reptilienarten in Peru. Reptilia 5:40–46. ———. 2002. Amphibien und Reptilien in Peru: Die Herpetofauna entlang des 10. Breitengrades von Peru: Arterfassung, Taxonomie, o¨kologische Bemerkungen und biogeographische Beziehungen. Dissertation, Natur- und Tier-Verlag, Naturwissenschaft, Mu¨nster, Germany. ———. 2005. The Telmatobius (Anura: Leptodactylidae) species of Peru´. In E. O. Lavilla and I. De la Riva (eds.), Estudios sobre las ranas andinas de los ge´neros Telmatobius y Batrachophrynus (Anura: Leptodactylidae), pp. 39–64. Monografı´as de Herpetologı´a Vol. 7, Asociacio´n Herpetolo´gica Espan˜ola, Valencia, Spain. LEHR, E., G. KO¨HLER, AND B. STREIT. 2002. Die Herpetofauna von Mittelperu entlang eines Transektes von der pazifischen Ku¨ste bis in die Hochanden. Faunistische Abhandlungen Museum fu¨r Tierkunde Dresden 22:361–392. MORALES, V. R. 1988. Una nueva especie de Telmatobius (Anura, Leptodactylidae), de Ancash, Peru´. Revista Brasileira de Zoologia 5:603–608. SALAS, A., AND U. SINSCH. 1996. Two new Telmatobius species (Leptodactylidae, Telmatobiinae) of Ancash, Peru. Alytes 14:1–26. SEIMON, T. A., A. SEIMON, P. DASZAK, S. R. P. HALLOY, L. M. SCHLOEGEL, C. AGUILAR, P. SOWELL, A. D. HYATT, B. KONECKY, AND J. E. SIMMONS. 2007. Upward range extension of Andean anurans and chytridiomycosis to extreme elevations in response to tropical deglaciation. Global Change Biology 13:288–299. SINSCH, U. 1986. Anfibios de la sierra central de Peru´. Una clave de identificacio´n para adultos y larvas. Boletı´n de Lima 45:23–33. ———. 1990. Froschlurche (Anura) der zentral-peruanischen Anden: Artdiagnose, Taxonomie, Habitate, Verahltenso¨kologie. Salamandra 26:177–214. SINSCH, U., A. W. SALAS, AND V. CANALES. 1995. Reassessment of central peruvian Telmatobiinae (genera Batrachophrynus and Telmatobius). I. Morphometry and classification. Alytes 13:14–44. VELLARD, J. 1951. Estudios sobre batracios andinos. I. El grupo Telmatobius y formas afines. Memorias del Museo de Historia Natural ‘‘Javier Prado’’ 1:1–89. WIENS, J. J. 1993. Systematics of the leptodactylid frog genus Telmatobius in the Andes of northern Peru´. Occasional Papers, Museum of Natural History, University of Kansas, Lawrence 162:1–76. Accepted: 24 June 2008.
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SHORTER COMMUNICATIONS APPENDIX 1 Larval Specimens Examined
Telmatobius mayoloi: Peru: ANCASH: Catac (09u4892309S, 77u2692309W, 3,520 m above sea level), on 31 March 1997 by E. Lehr and C. Aguilar: SMF 80799 (series of two: Stages 31, 40), MUSM 20551 (series of two: Stages 36, 40); between Conococha and Carpa in the Rı´o Santa (10u019100S, 77u199400W, 3,890 m above sea level), on 25 May 1998 by E. Lehr and C. Aguilar: SMF 80592 (series
of eight: Stages 31, 34, 35, 36, 38, 40 [33]): SMF 80594 (Stage 41); Rı´o Santa near km 143 on road from Lima to Huaraz, Provincia de Huaraz, Departamento de Ancash, on 9 August 1989 by H. Ortega: MUSM 7417 (series of two: Stages 28, 38); Conococha, from Rı´o Santa north of Conococha (10u0291409S, 77u199230W, 3,860 m above sea level), on 28 March 1997 by E. Lehr and C. Aguilar: MUSM 20550 (series of six: Stages 28, 29 [23], 31, 32, 36, 38 [used for condrocranium preparation, measurements except for total length not available]).