A new species of Telmatobius (Anura, Ceratophryidae) from Northern Jujuy Province, Argentina Author Barrionuevo, Sebastián Author Baldo, Diego text Zootaxa 2009 2030 1 20 journal article 10.5281/zenodo.186251 032f2475-dcbc-4fe6-ae7e-f41b07ab6185 1175-5326 186251 Telmatobius rubigo sp. nov. ( Fig. 1–2 ) , 1874 ): Fernández , 1926 : 277 [part.]. , 1841 ): Cei , 1980 : 255 [part.]. Holotype . Fundación Miguel Lillo, Tucumán, Argentina . ( FML 20828), adult male, from Argentina : Jujuy: Departamento Santa Catalina: El Queñoal ( 21º52'17.5''S , 66º06'09,2''W , 3966 m a.s.l.), collected on 15 January 2005 by Y . Arzamendia, D. Baldo, J. Baldo, S. Barrionuevo, D. Casagranda and A. Dallagnol. Paratopotypes. FML 20829 (male, cleared and stained), MACN 39092 (male), FML 20827, MACN 39093, MLP DB 3615, 3621 (females) and FML 20830-5, MACN 39094, MLP DB 3622, 3648 (juveniles), with same data as the Holotype . Referred specimens. FML 4391 ( 1 male , 3 females , 1 sub-adult, 2 juveniles ), from Argentina : Jujuy: Departamento Rinconada: Cuesta de Fundiciones, northeastern slop ( 22º29'S , 66º16', 3800 m a.s.l.); FML 2813 ( 4 males and 2 females ); FML 2814 ( 2 males , 4 females and 1 juvenile ); FML 2818 ( 2 males , 5 females and 1 juvenile ), from Argentina : Jujuy: Departamento Santa Catalina: Timón Cruz ( 22º12'S , 66º10'W , 4200 m a.s.l.), 25 April 1980 ; FML 4517 ( 1 male , 3 females and 2 sub-adults), from Argentina : Jujuy: Departamento Santa Catalina: Arroyo Farallón , north of Laguna de Los Pozuelos, 20 January 1989 ; MLP A 4710 (juvenil), MLP DB 4755-6, 4765, from Argentina : Jujuy: Departamento Santa Catalina: Santa Catalina, Santa Catalina River ( 21º55'00''S , 66º03'00''W , 3810 m a.s.l.), 6 January 2006 ; MLP DB 3457-9, 3576-8 (juveniles), from Argentina : Jujuy: Departamento Santa Catalina: Santa Catalina, Santa Catalina River ( 21º55'00''S , 66º03'00''W , 3810 m a.s.l.), 21 November 2004 ; 6 January 2006 ; 4766 (male), 4812 (tadpoles), from Santa Catalina, Provincial Road Nº 5, 2 Km eastearn from Santa Catalina ( 21º56'58,2''S , 66º02'21,6''W , 3802 m a.s.l.), 6 and 26 January respectively. MLP DB 4760 (tadpoles), from Argentina : Jujuy Province: Santa Catalina Departament : El Queñoal ( 21º52'17.5''S , 66º06'09,2''W , 3966 m a.s.l.), 7 January 2006 . FIGURE 1. (A) Dorsal and (B) ventral view of the holotype of Telmatobius rubigo (FML 20828). FIGURE 2. General dorsal view (A, bar = 10 mm); dorsal (B, bar = 5mm) and (C, bar = 5mm) lateral view of head; right hand (D, bar = 5mm), detail of nuptial excrescences (E, bar = 1mm), and right foot (F, bar = 5mm) of the holotype of Telmatobius rubigo (FML 20828). Etymology. The word rubigo is a Latin noun meaning rust, in allusion to the characteristic lichenous rusty blotches in the dorsal skin of this new species. The species name is used as noun in apposition to the genus name. Diagnosis and comparison with other species. We assign this new taxon to Telmatobius genus based on the presence of presumptive synapomorphies as nuptial pad in the thumb only, frontoparietals fused posteriorly (Wiens, 1993) and fanglike teeth embedded in the labial mucosa (Trueb, 1979). Telmatobius rubigo can be distinguished from any other species of Telmatobius by the following combination of characters: (1) snout–vent length (SVL) of males to 50.4 mm , females to 52.2 mm ; (2) head in lateral profile variable (depressed or high) with sloping snout; (3) head subacuminate in dorsal view; (4) upper lips flared, notched medially; (5) postcommisural gland absent or a set of small, separated glands; (6) tympanic annulus visible under the skin in the most of specimens; (7) forelimb of males moderately robust, without humeral spine; (8) dermal fringes of finger variable; base of thumb bearing nuptial pad in contact with the inner palmar tubercle; nuptial spicules small, closely arranged; (9) toes moderately webbed; plantar surface without spicules; (10) tarsal fold present; (11) skin smooth with pustules in the flanks and dorsal surface of thighs; (12) in life greenish-grey dorsum, uniform or with darker gray blotches, always scattered with rusty lichenous blotches; (13) belly pale orange, ventral surfaces of limbs bright orange, yellow and reddish; and (14) iris light grey without flecks; (15) skull well ossified; (16) frontoparietal fenestra bottle shape; (17) zygomatic ramus of squamosal long, slim and bended medially; (18) anterior end of cultriform process irregularly truncate and reaching the level of vomers; (19) posterior half of maxilla slim and straight in lateral view; (20) premaxilar alae inclined posteriorly in lateral view; (21) pars media plectri (columella) very thin; (22) hyoid plate extensively calcified; (23) clavicle and scapula fused; (24) tadpole without intramarginal mental papillae in the oral disc; (25) color in life of larvae brownish with characteristic bright yellow spots. Telmatobius rubigo differs from all the other species analyzed here by having dorsal skin scattered with rusty orange lichenous blotches, although the color in life of T. halli is unknown (Noble, 1938; Formas et al., 2003). Telmatobius rubigo differs from T. atacamensis , T. aff. marmoratus, T. huayra , T. marmoratus , T. chusmisensis , T. fronteriensis , T. phillipi and T. vilamensis in exhibiting bright orange and yellow coloration on the ventral skin of limbs. Tympanic annulus is visible externally in the most of the specimens of T. rubigo (never visible externally in T. huayra , T. dankoi , T. fronteriensis , T. halli , T. phillippi , and T. vilamensis ). In Telmatobius rubigo the spicules in the chest are present only in males, while in T. atacamensis , T. chusmisensis , T. hypselocephalus and T. marmoratus they can be present in both sexes. These spicules are absent in both sexes in T. halli and T. phillippi . The upper lip of Telmatobius rubigo is notched medially (unnotched in T. phillippi ), and the nuptial pad is in contact with the inner palmar tubercle (not in contact in T. phillippi ). Telmatobius rubigo has pointed, curved and well developed maxillar and premaxillar teeth which are absent in T. dankoi and T. vilamensis ; premaxillar teeth are absent and maxillar ones are vestigial in T halli ). Telmatobius rubigo has a well ossified skull (poorly ossified in T. chusmisensis , T. platycephalus , T. phillippi and T. vilamensis ) and a well ossified carpus (which is cartilaginous in T. chusmisensis ). The vomer in T. rubigo is not well developed, as it is in T. marmoratus , T. atacamensis , T. platycephalus , T. hypselocephalus , and T. phillippi . The species from subpáramo valleys or cloud forests from Argentina ( T. ceiorum , T. contrerasi , T. hauthali , T. laticeps , T. oxycephalus , T. pinguiculus , T. pisanoi , T. schreiteri , T. scrocchii and T. stephani ) differ from T. rubigo in exhibiting the following characters: (1) higher heads, (2) eyes more laterally positioned, and closer to the mouth, (3) shorter snout, with the nostrils closer to the tip of snout than to the eyes, (4) superior lip not flared, unnotched medially, (5) nuptial spines larger. Description of the Holotype ( Fig. 1 , 2 ). An adult male, medium-sized, SVL 50.4 ( Fig. 2 A). Head length (measured from commissural angle to tip of snout) 33.6% of SVL, wider than long; head width (measured at level of commissural angle) 35.8% of SVL; head length 93.9% of head width; head subacuminate in dorsal view ( Fig. 2 B), slightly depressed with sloping snout in lateral profile ( Fig. 2 C); nostrils small, not protruding, oriented dorsolaterally, approximately at the same distance to tip of the snout and to the eye; internarial region convex; canthus rostralis indistinct, loreal region slightly concave; upper lips flared, notched medially; maxilla and premaxillae with small curved and pointed teeth, embedded in the labial mucosa; teeth of vomers not visible; small, oval choanae; tongue rounded, attached anteriorly to floor of mouth, free posteriorly for about one-fourth of its length; vocal slits absent; tympanum and tympanic annulus absent; supratympanic fold extending obliquely from behind eye to posterior margin of postcommisural gland and in continuity with suprahumeral fold. Forelimbs moderately robust; relative length of fingers II <I <IV <III ( Fig. 2 D); webbing absent between fingers; tips of fingers swollen; fingers with almost unnoticeable lateral fringes; thenar tubercle large, elliptical, depressed; palmar tubercle rounded, approximately the same size as thenar tubercle; one subarticular tubercle on the base of each finger and at penultimate phalangeal articulation on Fingers II and IV; one round supernumerary tubercle between inner and outer palmar tubercles; palmar surface smooth, without keratinized spicules; base of thumb broadened proximally, bearing a nuptial pad composed of small keratinized, black and conical spicules closely arranged, reaching posterior margin of thenar tubercle, and extending dorsally from the base of thumb to the base of the distal phalange of thumb ( Fig. 2 E). Hind limbs moderately robust; tibia length 44.7% of SVL; tibia length 88.8% of foot length (measured from the proximal border of the inner metatarsal tubercle to the tip of the fourth toe); relative length of toes I <II <III <V <IV ( Fig. 2 F); webbing formula I (1 1/2) – (--) II (1 1/3) – (--) III (2 1/3) – (3 3/4) IV (3 1/2) – (2+)V; webbing extending as lateral fringes to tips of toes; tips of toes swollen, approximately of the same size as those of fingers; outer metatarsal tubercles rounded, slighthly smaller than one-fourth size of the elliptical inner metatarsal tubercle; subarticular tubercles rounded and well developed; supernumerary tubercles absent; plantar surface smooth; tarsal fold developed. Skin of the dorsum and dorsal surface of the head relatively smooth; skin of flanks with pustules, some of them with keratinized tip; some small tubercles are present on the lateral region of the head; dorsal surface of limbs with flat pustules; skin of venter and ventral surfaces of limbs smooth except for keratinized spicules in the chest; cloacal opening unornamented, located just below the dorsal level of thighs. Coloration of the Holotype ( Fig. 1 ). In life, grey dorsum, with darker gray solid blotches and scattered with orange lichenous blotches; venter pale orange, throat pale pink, ventral surfaces of limbs bright orange, yellow and reddish; iris light grey without flecks. In preservative brownish-gray dorsally with solid darker grey blotches; the belly, throat and the ventral surfaces of limbs are pale cream. The palmar and plantar surfaces are dark gray with the tubercles pale gray. Measurements of the Holotype . SVL 50.4; HW 18.0; HL 16.9; ED 4.5; IOD 6.6; EW 2.9; END 3.1; IND 3.0; SL 7.5; EMD 3.9; NMD 4.3; BW 19.2; BH 12.4; HAL 12.4; RUL 10.9; FEL 22.0; TL 22.5; FOL 37.6; WL 14.1; IMTL 2.0; OMTL 0.8. Variation. Range of measurements of 20 adult specimens (average given in parentheses): SVL 39.9–54.0 (47.7); HW 13.9–19.8 (16.9); HL 12.3–17.5 (15.1); ED 4.0–4.9 (4.5); IOD 5.2–7.0 (6.4); EW 1.8–3.2 (2.5); END 2.7–4.5 (3.2); IND 2.5–3.6 (2.9); SL 4.1–7.6 (6.4); EMD 2.8–4.8 (3.8); NMD 3.6–4.9 (4.2); BW 14.5–20.6 (17.9); BH 10.4–16.2 (11.8); HAL 10.4–13.5 (11.7); RUL 8.3–11.2 (10.0); RUW 3.3–5.8 (4.3); FEL 17.4–25.1 (20.8); TL 18.2–24.0 (21.0); FOL 29.6–39.4 (34.3); WL 10.7–14.5 (12.6); IMTL 1.4–2.6 (1.9); OMTL 0.4–1.4 (0.8). As in other Telmatobius species head shape in Telmatobius rubigo is extremely variable, from moderately depressed with a long snout to a high head with a short snout. The supratympanic fold is strong to almost unnoticeable. The number of vomeral teeth range from 0 to no more than 3, sometimes completely covered by the palatal mucosa. The skin on the flanks is more tuberculate in some specimens, especially those from Cuesta de Fundiciones and in large females from El Queñoal. The skin of specimens from Arroyo Farrallón is similar to those of El Queñoal but whit dorsal skin of thighs being more warty and spiny. The skin in most specimens from Timón Cruz is smooth, occasionally with a few scattered spicules. The tympanic annulus is evident in most of analyzed specimens except in those from Timón Cruz, where the majority has a concealed annulus. The development of webbing varies intra and interpopulationally. The studied adult specimens present some differences in color pattern in life, the ground color can be grey with darker blotches or uniformly greenish. All examined adults specimens share the pattern of rusty lichenous blotches that is also present in some juveniles and metamorphs ( Fig. 3 ). However, a few juvenile individuals are completely uniform. In preservative the design of dorsum skin is uniformly grayish or grey with darker blotches. The venter is uniformly cream, but in some specimens darker blotches are present in thighs. FIGURE 3. Paratopotype (juvenile) of Telmatobius rubigo showing the characteristic color pattern of lichenous rusty blotches. The paratopotype MLP DB 3622 has a malformation in the left foot; fingers I, IV and V are malformed and the tip of finger III is atrophied. Sexual dimorphism is evident, mature males have a nuptial pad in the dorsal and ventral surface of thumbs that reach the inner metacarpal tubercles and also have scattered spines in the chest and arms. Those spines are absent in females. The males are smaller than females (SVL: males 39.9–50.4 mm ; females: 43.6–54.0 mm), and have more robust forearms (RUW: males 4.2–5.8 mm ; females 3.3–5.5 mm .), and more developed webbing (WL/FOL: males up to 58%, females up to 56%). Osteology ( Fig. 4 ). Skull moderately depressed, well ossified ( Fig. 4 A–B); sphenethmoid well ossified, broadly exposed dorsally, anterior portion reaching anterior border of nasals; prootic well ossified, fused with exoccipital dorsally and ventrally, crista parotica consists in a narrow strip of cartilage articulated with otic ramus of squamosal; exoccipital ossified dorso and ventromedially; frontoparietals fused posteriorly; frontoparietal fenestra bottle shaped, extends more than half the length of the orbit. Anterior margin of fontanelle formed by sphenetmoid at a level approximately ¼ posterior on length of orbit; posterior margin of fontanelle at a level approximately ¼ anterior on length of orbit; nasals subtriangular, elongate, close to each other medially, lateral vertex bended backwards, in contact with sphenethmoid; maxillary process of nasal not reaching pars facialis of maxilla; pars facialis of maxilla lacking preorbital process; neopalatines long, straight, slightly tapered medially; in contact laterally with pterygoid and medially with sphenethmoid, broadly separated and not reaching anterior terminus of cultriform process of parasphenoid; parasphenoid robust, cultriform process irregularly truncate and reaching the level of vomers; alae of parasphenoid large, almost perpendicular to cultriform process and not in contact with medial ramus of pterygoids anterolaterally; pterygoid well developed and robust, anterior ramus extending to level of palatine; inner margins of rami describing a nonsinuous curve between maxilla and otic capsule. Outer margins of anterior and posterior rami forming a slightly curve profile between maxilla and squamosal; medial ramus of pterygoid short, barely contacting prootic by pseudobasal cartilague; maxillary arch complete; quadratojugal moderately long, contacting maxilla but not fused; basal portion of palatoquadrate cartilage mineralized; palatoquadrate cartilage invested by the quadratojugal, posterior ramus of pterygoid, and ventral ramus of squamosal; otic ramus of squamosal shorter and broader than zygomatic ramus; zygomatic ramus long, slim and bended medially; plectral apparatus with pars interna plectri well developed, pars media plectri thin and pars externa plectri cartilaginous; tympanic ring incomplete dorsally; maxilla and premaxilla dentate; teeth pedicellate, monocuspid, fang-like; number of teeth in premaxilla (11/10) and in maxilla (33/34); posterior half of maxilla slim and straight in lateral view; alary process of premaxilla wider at the base, acuminate and bifurcate, the distal half slightly bended outwards in frontal view; disposed with more than 66º with respect to the maxillary in lateral view; vomers small, with slender prechoanal and postchoanal process; dentigerous process of vomers without teeth in the cleared specimen. FIGURE 4. Dorsal (A) and ventral (B) view of skull, pectoral girdle (C), right humerus (D) and hyoid apparatus (E) of a paratopotype of Telmatobius rubigo (FML 20829). Hyoid plate wider than long ( Fig. 4 E), calcified around the bases of the posteromedial processes, which originate inside the plate, and are long and well ossified. Other calcified areas inside the plate are also present; hyale long, sinuous, bearing well developed anterior processes; anterolateral or alar processes moderately long, expanded distally; posterolateral processes straight, divergent respect to midline, approximately as long as anterolateral processes, not expanded distally; esophagic process of aritenoid cartilage broad. Pectoral girdle arciferal ( Fig. 4 C); omosternum cartilaginous, slightly calcified basally; sternum completely ossified, xiphisternon bilobed; clavicle fused with scapula; coracoids well developed, not juxtaposed medially; epicoracoids calcified; scapula bicapitate; suprascapula with cleithrum bifurcate and cartilaginous portion calcified. Eight procelous presacral vertebrae; transverse processes of presacrals II, III, and IV moderately large, relative width of those transverse processes: III> IV> II; sacral diapophyses moderately expanded, oriented slightly posterodorsal to longitudinal axis of vertebral column; articulation of sacrum with urostyle bicondylar; urostyle bearing small, proximal dorsal crest; ilia cylindrical, long; pubis completely ossified. Humerus of male robust ( Fig. 4 D), flattened, with well developed anteroproximal, medial and ventral crests. The two last crests continue anteriorly as one crest to almost reach the head of the humerus; phalangeal formula of hand 2-2-3-3; terminal phalanges knobbed; prepollex with one basal and four distal elements; basal and three distal elements ossified; phalangeal formula of foot 2-2-3-4-3; prehallux with four elements, basal and one distal elements ossified. Skull measurements. Skull length 16.5, skull width 17.4; orbit length 5.8; dorsal length sphenetmoid 3.4; ventral length sphenetmoid 4.4; frontoparietal length 9.0; slope angle of alar process of premaxillary in lateral view with respect maxillary 66.55º. Tadpole external morphology ( Fig. 5 ). The larvae of Telmatobius rubigo belongs to the benthic ecomorphological guild (section II: A: 1) of McDiarmid and Altig (1999), as revised from Altig and Johnston (1989). Body length about 40% of total length (BL/TL= 0.38 ± 0.02) ( Fig. 5 B); body shape oval in dorsal view ( Fig. 5 D–E); with the maximum body width placed at the posterior portion of the head, sometimes at abdominal region. Body depressed in lateral view (BH/BW=0.78 ± 0.06); ventral contour flat to slightly convex, dorsal contour convex from the anterior edge of the oral disc to the posterior border of the eyes, and almost straight from behind the eyes to the origin of the dorsal fin. Snout rounded or slightly truncated in dorsal view, and rounded in lateral view. Nostrils not protuberant, rounded or oval, and situated dorsally, closer to eye than snout (NS/EN= 1.66 ± 0.16). Eyes relatively small, situated dorsolaterally. Spiracle single, lateral, sinistral, directed posterodorsally; almost as long as wide, and placed approximately halfway between snout and posterior margin of body, with its inner wall present as a slight ridge; its opening is oval, placed below body midline, being its diameter smaller than the tube diameter, and visible in dorsal and lateral view. Intestinal assa central. Vent tube dextral, with opening not visible laterally, concealed by a fold of the vent tube wall; right wall attached anteriorly to the ventral fin. Tail large (TAL/TL= 0.62 ± 0.02), with well developed musculature, not reaching tail tip. Tail axis straight, and tail tip rounded. Dorsal and ventral fins of similar depth; dorsal fin not extending onto body; ventral fin starting at the end of vent tube; both fins almost as high as body height. Edge of both fins sub-parallel for the first and second thirds, and convex in the posterior third. Oral disc ( Fig. 5 C) anteroventral, transversally elliptical; medium sized, not emarginated, and with a large dorsal gap in the marginal papillae, with no mental gap. Marginal papillae simple, small and longer than wide, sub-conical; arranged in a single or single alternated row in the anterior (upper) labium, and arranged in a single alternate or occasionally double row in the posterior (lower) labium. Sub-marginal papillae larger than marginal ones; few and scattered in supra-angular region, but denser in the angular and infra-angular regions. Sub-marginal infra-angular papillae arranged in short bilateral flaps which run close to the labial teeth ridges, and converge towards the mental region. Some additional sub-marginal papillae are scattered in the mental area, but not forming a complete row. Jaw sheaths robust, wider than deep, FIGURE 5. Dorsal view of a living tadpole of Telmatobius rubigo showing the characteristic yellow blotches (A). Lateral (B, bar = 10mm), dorsal (D, bar = 5mm) and ventral (E, bar = 5mm) view of a fixed tadpole (FML 21157). Detail of the oral disc (C, bar = 5mm). finely serrated, and heavily pigmented distally for about 2/3 to 1/2 their length. Free margin of the upper jaw sheath widely arch shaped, with lateral processes well developed. Lower jaw sheath V-shaped. Labial tooth row formula (LTRF) 2(2)/3(1). Gap in A2 clearly visible. Medial gap in P1 very narrow. In one specimen (stage 34) there is a small ridge with labial teeth between A1 and A2. Tadpole coloration ( Fig 5 A): In life ground color brown-greenish with rounded bright yellow blotches scattered in the dorsum of body, tail musculature and fins. Iris golden. In preservative, body brown-grayish, tail tan, dorsal and ventral fins translucent. Skin with small darker flecks. In some specimens those flecks are tiny, more numerous and evenly distributed. Large dark blotches are present in the fins at the tip of the tail, more intensely pigmented in some individuals; venter transparent, internal organ visible; eyes black; perinasal region darker than surrounding areas. Tadpole measurements. Range of measurements of 9 tadpoles (average given in parentheses): TL: 64.4–77.7 (72.0); BL: 25.2–29.7 (27.5); TAL: 39.2–49.6 (44.5); BW: 13.8–17.5 (16.3); BH: 11.9–13.8 (12.7); IOD: 3.8–4.3 (4.0); IN: 2.8–3.2 (3.0); EN: 2.5–3.0 (2.7); NS: 3.9–4.9 (4.5). In Table 1 tadpole measurements are discriminated according to Gosner (1960) developmental stages. TABLE 1. Measurements (mm) of the tadpoles of Telmatobius rubigo , providing mean ± SD and range.

Stage 33 (n=1)	Stage 34 (n=5)	Stage 35 (n=1)	Stage 36 (n=1)	Stage 37 (n=1)
Total length	72.5	72.0 ± 4.9 (64.4–77.7)	73.6	72.6	69.6
Body length	28.7	27.6 ± 1.6 (25.2–29.7)	26.6	26.8	27.0
Tail length	43.8	44.3 ± 4.0 (39.2–49.6)	47.0	45.8	42.6
Body width	16.0	16.2 ± 1.5 (13.8–17.5)	16.4	15.9	17.0
Body height	13.2	12.7 ± 0.7 (11.9–13.8)	12.3	12.6	12.7
Interorbital distance	3.8	4.07 ± 0.15 (3.9–4.3)	3.8	4.0	4.0
Internarial distance	2.8	3.1 ± 0.1 (2.8–3.2)	3.2	2.9	2.9
Eye-narial distance	3.0	2.7 ± 0.2 (2.6–3.0)	2.5	2.7	2.6
Narial-snout distance	4.4	4.5 ± 0.4 (3.9–4.7)	4.9	4.2	4.6

Karyotype ( Fig 6 ). Chromosome counting from all the samples revealed that Telmatobius rubigo has a karyotype composed by 13 biarmed chromosomes pairs (2n = 2x =26) without identifiable sex chromosomes (Fig, 6 A). The first six pairs of the chromosome complement are large and the others seven are small. Pairs 1, 3, 8, 10, 11, 12 and 13 are metacentric, pairs 2, 4, 6, 7 and 9 are submetacentrics, and pair 5 is subtelocentric. The secondary constrictions and Ag-NOR sites were observed on the short arms of both homologues of chromosome pair 6 ( Fig. 6 B). All normal stages were observed in meiotic preparations. A notorious bouquet polarization (Rabl orientation) was observed in Zygotene and early Pachytene, with telomeres grouped in a restricted cluster of the inner membrane of the nuclear envelope ( Fig. 7 A). Zigotene cells showed the telomeric regions paired and condensed, and internal regions unpaired and highly discondensed, indicating early stages of synapsis. Thus, the initiation of chromosome pairing was apparently distal and bidirectional (which is also supported by the observation of internal asynaptic loops in bivalentes IIs), give rise to ring IIs with terminal chiasmata in Diakinesis and Metaphase I ( Fig. 7 B). In Metaphase II, 13 chromosomes were always observed ( Fig. 7 C). FIGURE 6. Karyotype of paratopotype of Telmatobius rubigo (MLP DB 3622) (A). Sixth chromosome pair bearing the NORs (B). FIGURE 7. Meiotic cells of paratopotype of Telmatobius rubigo (MACN 39093): Zygotene showed a bouquet polarization (Rabl orientation) (A); Metaphase I with 13 ring IIs (B); Metaphase II with 13 chromosomes (C). Geographic distribution and natural history ( Fig. 8 ). This species is only known from a few localities, from the endorheic basin of Laguna de Los Pozuelos ( Fig. 8 ) in the Central Andean Puna Ecoregion (Dinerstein, et al. 1995), Jujuy province, Argentina . This basin comprises an area of about 3800 Km2 between 3600 and 4700 m a.s.l. and collects water of streams of Sierras de Escaya from the east, Sierra de Cochinoca from east and southeast, Sierra de Rinconada from the west, and Sierra de Carahuasi from the west and southwest ( Fig. 8 ) (Cajal et al . 1998). The climate of the Puna is cold and dry with average annual temperatures that range between 0–9º C (depending on the altitude), and large daily thermal amplitude (around 30º C) (Tecchi, 1991). Precipitations oscillate between 100 and 500 mm /year, that fall as snow and hail in the winter, and in the summer rains are more common (Cabrera, 1968). At the type locality (El Queñoal) T. rubigo inhabits a high altitude permanent stream that flows through a gorge. The riparian vegetation is predominantly characterized by some scattered trees (“queñoas”, Polylepis tomentella ), arborescence cactuses (“cardones” Thrichocereus spp.), and tall grasses ( Cortaderia speciosa ) ( Fig. 9 ). This stream crosses the Puna steppe and drains to Santa Catalina river , a tributary of Laguna de Los Pozuelos. Specimens of T. rubigo were always found underwater, on permanent high altitude streams. Some specimens were collected under submerged rocks, others while were resting at the bottom of small natural ditches of quiet waters and therefore they were easily detected from the edge of the streams. The few people that live at El Queñoal raise corn, beans and flower crops by the margins of the stream. This locality is about 10 km (straight-line distance) NW from Santa Catalina town. At the type locality and Santa Catalina river Telmatobius rubigo is sympatric with Hypsiboas sp. ( pulchellus group) ( Hylidae Rafinesque ) and Rhinella spinulosa (Wiegmann) ( Bufonidae Gray ).