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The genus Pseudis (Anura: Pseudidae) in Rio Grande do Sul, southern Brazil, with description of a new species

Axel Kwet
Universitt Tbingen, Zoologisches Institut, Auf der Morgenstelle 28, D-72076 Tbingen, Germany e-mail: axel.kwet@uni-tuebingen.de Abstract. A new species of Pseudis is described from So Francisco de Paula, Rio Grande do Sul, Brazil. It is known from the southern parts of the Serra Geral where it occurs in grassland, inhabiting permanent ponds and still-water zones of slow owing creeks. It is characterized by a paired vocal sac and a bulbous thumb, considerably widened at the base. It is distinguished from P. minutus by its different call, body coloration, rounded snout, and a more robust body with shorter hindlimbs. Advertisement call, tadpole, and life history are described. The taxonomy of the family Pseudidae in Rio Grande do Sul is discussed, supporting the synonymy of Lysapsus mantidactylus and P. meridionalis with P. minutus .

Introduction The small neotropical family Pseudidae currently contains six species: Lysapsus limellus Cope, 1862, Pseudis bolbodactylus Lutz, 1925, P. fuscus Garman, 1883, P. minutus Gnther, 1859 1858, P. paradoxus (Linnaeus, 1758), and P. tocantins Caramaschi and Cruz, 1998. Lysapsus mantidactylus (Cope, 1862) has been considered a junior synonym of P. minutus by Klappenbach (1985), and Caramaschi and Cruz (1998) revalidated the taxonomic status of P. bolbodactylus and P. fuscus, previously treated as subspecies of P. paradoxus . According to Savage and Carvalho (1953) the generic name Pseudis is of masculine gender, even though Caramaschi and Cruz (1998) and many other authors (Gnther, 1859 1858, Cope, 1862; Boulenger, 1882, 1886; Garman, 1883; Lutz, 1925; Miranda-Ribeiro, 1926; Frost, 1985; Cei, 1987; Emerson, 1988; Duellman, 1993; Dixon et al., 1995; Bosch et al., 1996; S and Lavilla, 1997) treated it as feminine. A general revision of the Pseudidae may show that other subspecies of P. paradoxus and the different lineages of L. limellus also represent valid species (Klappenbach, 1985; Caramaschi and Cruz, 1996; S and Lavilla, 1997).
c Koninklijke Brill NV, Leiden, 2000 Amphibia-Reptilia 21: 39-55

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Axel Kwet

During eld studies in the Brazilian state of Rio Grande do Sul (Kwet, 1997, 1999; Kwet and Di-Bernardo, 1999) two different species of harlequin frogs were found. Due to their morphological similarities, both were considered a single species in the amphibian list of Braun and Braun (1980). However, the pseudid occurring on the Araucaria plateau of southern Brazil differs considerably from P. minutus of the lowland regions of Rio Grande do Sul, Uruguay and Argentina, and will be described here as a new species. In order to con rm the proposed inclusion of L. mantidactylus and Pseudis meridionalis MirandaRibeiro, 1926 in P. minutus (Klappenbach, 1985), advertisement calls and morphological characters of pseudids from several regions of Rio Grande do Sul and Argentina will be compared.

Methods The type material was collected at the Centro de Pesquisas e Conservao da Natureza Pr-Mata, which is located at the Serra Geral in Rio Grande do Sul, Brazil, approximately 150 km northeast of the city of Porto Alegre. Specimens examined are deposited in the Museu de Cincias e Tecnologia da PUCRS (MCP, Porto Alegre, Brazil), Staatliches Museum fr Naturkunde Stuttgart (SMNS, Stuttgart, Germany), Zoologisches Museum Hamburg (ZMH, Hamburg, Germany), Zoologisches Museum Berlin (ZMB, Berlin, Germany), and Natural History Museum (formerly British Museum of Natural History (BM) London, U.K.). The following measurements were taken to the nearest 0.1 mm with dial calipers (if not described specially, positions of measurement after Cei, 1980): snoutvent length (SVL); head length (HL); head width (HW); horizontal tympanum diameter (TD); horizontal eye diameter (ED); eyelid length (EL); eyelid width (EW); eye-nostril distance, from center of the nostril to the anterior edge of eye (EN); nostril-snout distance, from center of nostril to tip of snout (NS); internarial distance between centers of nostrils (IN); thigh length (THL); tibia length (TL); foot length, from proximal edge of inner metatarsal elevation to tip of fourth toe (FL); hand length, from proximal edge of thumb articulation to tip of the third nger (HAL); thumb width, taken at widest point on thumb base (TW). All specimens collected were xed in formalin 6% and after one or two days transferred into 70% ethanol. Drawings were made with a camera lucida attached to a stereomicroscope (Wild Heerbrugg). Call recordings from six Pseudis populations were obtained with a Sony WM-D6C tape recorder and a Sennheiser K6 microphone. Three to six calls from each place were analyzed on a Kay DSP Sonagraph 5500, using a wide band display. The following voucher specimens were collected, referring to localities where the sound recordings were obtained: MCP 2590, MCP 3351, MCP 3491, SMNS 9001:3, and SMNS 9124.

A new species of Pseudis

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Results Pseudis cardosoi sp. nov. ( gs. 1A, B) Holotype. MCP 3354, adult male, collected at the Centro de Pesquisas e Conservao da Natureza Pr-Mata, 29 30 S, 50 10 W, Municipality of So Francisco de Paula, Serra Geral, 960 m a.s.l., State of Rio Grande do Sul, Brazil, on 20 January 1998. Paratypes. MCP 2579, adult female and MCP 2580, adult male, collected on 18 December 1996. MCP 2577, MCP 2590, adult males and MCP 2581, adult female, collected on 16 January 1997. MCP 2588, adult male, collected on 8 February 1997. MCP 3193, adult female and MCP 3194, adult male, collected on 3 December 1997. MCP 3350, adult male, collected on 28 October 1997. All paratypes are from the type locality.
Table 1. Characteristics for distinguishing adults of Pseudis cardosoi and P. minutus . Pseudis cardosoi sp. nov. Snout-vent length Males 33-46 mm (mean 42 mm), females 45-56 mm (mean 51 mm). Robust and short; when hindlimb adpressed to body, metatarsal articulation not reaching tip of snout. Pseudis minutus Males 24-39 mm (mean 32 mm), females 39-51 mm (mean 46 mm). More slender and longer; when hindlimb adpressed to body, metatarsal articulation reaching tip of snout or beyond. Slender; base of rst nger slightly widened (thumb width in males 1.62.4 mm, in females 2.0-2.4 mm).

Hindlimbs

Fingers

Robust; base of bulbous rst nger considerably widened (thumb width in males 2.5-3.3 mm, in females 2.73.4 mm). Olive, dark to light green, or brown. Large dark blotches present or not; lateral surface of body without light lateral stripe; no light middorsal band. Only one distinct stripe. First stripe at level of vent opening indistinct; second below cloaca most conspicuous and extending across the entire thigh; remaining lines incomplete, irregular and much fainter. Broadly rounded. Not dilated.

Dorsal coloration in life Dorsal pattern

Light green or brownish. Usually with small dark blotches; lateral surface of body mostly with light longitudinal stripe; sometimes with distinct light middorsal band. Usually three distinct stripes. First, second and third stripes extending across the entire thigh; remaining one or two lines incomplete and indistinct.

Pattern of dark longitudinal stripes on lower side of thigh

Lateral snout pro le Tips of toes

Truncate. Slightly dilated.

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Axel Kwet

(A)
Figure 1. Pseudis cardosoi sp. nov. (A) Holotype (MCP 3354), adult male. (B) Calling male at breeding pond in Pr-Mata (Municipality So Francisco de Paula).

Diagnosis (see also table 1). A medium sized species of Pseudis with paired vocal sac in males. This distinguishes it from P. paradoxus which has a single vocal sac and a larger body size. The new species ( gs. 1A, B) differs from P. minutus ( g. 2) by having a more robust body, metatarsal articulation not reaching tip of snout when hindlimb adpressed to body (articulation reaching beyond tip of snout in P. minutus), a rounded snout ( g. 3A) (truncate in P. minutus; g. 3B), bulbous and robust ngers with thumb considerably widened at base ( g. 4A) (more slender in P. minutus; g. 4B), and by absence of noticeably dilated tips of toes ( g. 5A) (dilated in P. minutus; g. 5B). Description of holotype. Adult male, MCP 3354 ( g. 1A). Body robust. Head broad, slightly attened, approximately as long as wide ( g. 3A). Snout rounded in dorsal and lateral views, upper jaw protruding in pro le. Eye to nostril distance about equal the distance from nostril to tip of snout. Nostrils very scarcely protuberant, directed dorsally; distance from nostril to tip of snout as wide as internarial distance. Loreal region gently sloping. Canthus rostralis indistinct, rounded. Eyes large, slightly projecting; interorbital region at, wider than internarial distance. Tympanum distinct, round, its diameter sligthly smaller than eye diameter; supratympanic fold very weak. Vomerine teeth in two short, rounded, well-separated patches between the choanae; choanae small

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(B)
Figure 1. (Continued).

and widely separated; tongue large, cordiform. Vocal sac of males paired, conspicuous as wrinkled, dark areas in the lateral gular region. Forearms robust; upperarms short and more slender. Fingers robust, bulbous at base, without webbing; tips of ngers not attened or expanded ( g. 4A). Relative lengths of ngers 2 < 1 = 4 < 3. Thumb considerably widened at base. Metacarpal tubercles extremely reduced; subarticular tubercles small, conical, and raised, very distinct on thumb. Hindlimbs robust, short; when adpressed to body, metatarsal articulation not reaching tip of snout. Toes extensively webbed; web reaching the base of the tips of toes; tips of toes not dilated ( g. 5A). Relative lengths of toes 1 < 2 < 5 < 3 < 4. Inner metatarsal tubercle sharp, spurlike, elliptical; outer metatarsal tubercle absent; subarticular tubercles small, conical and raised; tarsal fold present. Skin on dorsum and hindlimbs closely, coarsely granulated; longitudinal rows of tubercles on upper surface of thigh, tibia and tarsus; skin on head and venter smooth. For measurements see table 2. Weight of the holotype in life 10.7 g. Coloration of holotype in preservative. Dorsum dark gray to olive-gray, with irregularly formed, blackish blotches. Dorsal surface of limbs with coloration like dorsum; dark blotches on each forelimb, thigh, tibia, and foot. Dark line from tip of snout across nostril to eye. Ventral surface white, scarcely dotted on venter. Ventral design of thigh with one distinct dark longitudinal line, and three or four weaker, interrupted stripes.

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Figure 2. Pseudis minutus . Adult male from Guaritas (Municipality Caapava do Sul).

Coloration in life and variation. Dorsal surface of body and limbs light to dark green, olive, or brownish, usually with some darker spots or blotches of irregular form. Dorsum without light middorsal line. Black or dark green stripe along canthus rostralis narrow, in some specimens broad and blotlike. Tympanum olive, tan or light brown. Coloration around anal region and at lateral surface of head, body and thigh light yellow. Ventral surface white, on venter usually scattered with dark brown dots. Ventral surface of thigh with several dark brown longitudinal lines, varying in number, form, and extension. First line at the level of vent opening usually indistinct, due to the dark dorsal surface. Second line below cloaca most conspicious, broad and extended, the third one incomplete and interrupted. The remaining two or three lines very irregular and much fainter. In some specimens the lines are partially con uent, forming a net-like pattern. For variation in measurements of males and females, compared with the data of P. minutus, see table 2. Geographical distribution. The new species occurs in the southern regions of the Serra Geral. It is known from several localities in northeastern Rio Grande do Sul, for example from the Municipalities Canela, Gramado, So Francisco de Paula, Cambar do Sul, Bom Jesus and Vacaria. It could be assumed that it also occurs in the neighbouring State of Santa Catarina, as some collecting places are situated only a few kilometers from the border. Sympatry with P. minutus has not been observed.

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Figure 3. Lateral view of the head of (A) the holotype of Pseudis cardosoi sp. nov. (MCP 3354), and (B) Pseudis minutus (MCP 3492). Scale bar = 5 mm.

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Table 2. Measurements of Pseudis cardosoi and P. minutus (mm). Pseudis minutus Gnther, 1859 1858 Females n = 15 Range s 2.763 1.363 0.789 0.431 0.456 0.478 0.335 0.300 0.346 0.313 1.455 0.816 1.232 0.607 0.232 24.5-38.4 9.5-13.7 8.5-13.6 2.7-4.5 2.3-4.1 3.5-5.3 1.6-2.9 1.7-2.8 1.6-2.7 1.3-2.2 13.7-24.4 13.3-22.5 13.1-22.9 8.8-12.9 1.6-2.4 32.05 11.48 11.19 3.29 3.16 4.29 2.28 2.30 2.16 1.77 19.60 18.06 18.86 11.19 1.99 4.168 1.267 1.387 0.501 0.494 0.518 0.313 0.289 0.289 0.224 3.302 2.587 2.747 1.155 0.224 s 45.4-55.9 14.9-20.1 16.0-18.9 3.5-5.3 4.0-5.6 4.9-6.6 2.5-3.8 2.1-3.3 3.0-4.0 2.0-3.2 26.1-30.6 23.4-26.9 23.9-28.6 14.4-16.6 2.7-3.4 51.09 17.85 17.39 4.41 4.63 5.75 3.07 2.96 3.36 2.59 28.27 24.59 25.45 15.41 2.97 Mean Range Mean Males n = 23 Females n = 5 Range 39.5-50.5 14.8-18.1 13.4-17.4 4.1-5.1 3.5-5.3 4.5-6.6 2.8-3.9 2.8-3.4 2.2-3.6 1.8-2.6 25.3-30.4 22.9-27.4 22.8-27.7 11.8-14.9 2.0-2.6 Mean 45.80 16.22 15.78 4.55 4.38 5.27 3.10 3.15 2.92 2.32 27.65 25.23 24.98 13.88 2.30 s 3.751 1.141 1.420 0.418 0.627 0.703 0.400 0.217 0.458 0.286 2.077 1.628 1.726 1.118 0.253

Pseudis cardosoi sp. nov.

Males n = 22

Range

Mean

SVL HL HW TD ED EL EW EN NS IN THL TL FL HAL TW

36.3-45.9 12.7-16.3 12.3-16.8 3.2-4.4 3.4-4.6 4.7-6.0 2.2-3.4 2.1-3.2 2.3-3.7 2.0-2.9 20.3-26.5 18.6-24.7 18.5-24.2 10.6-14.6 2.5-3.3

42.20 14.71 14.87 3.68 4.06 5.45 2.92 2.72 2.89 2.44 24.22 21.42 21.82 12.72 2.89

2.580 0.937 1.025 0.325 0.350 0.354 0.265 0.309 0.306 0.220 1.600 1.643 1.633 1.029 0.232

Axel Kwet

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Figure 4. Ventral view of the right hand of (A) the holotype of Pseudis cardosoi sp. nov. (MCP 3354), and (B) Pseudis minutus (MCP 3379). Scale bar = 5 mm.

Etymology. The species name honors the late Ado Jos Cardoso, an outstanding herpetologist and personality, in recognition of his contributions to the knowledge of the Brazilian anurofauna. Egg description. The egg of Pseudis cardosoi is characterized by a considerable size (egg diameter 2.5-3.0 mm, mean 2.7 mm, n = 20), being much larger than that of P. minutus (egg diameter 1.5-2.0 mm, mean 1.7 mm, n = 20). It shows a dark brown and white coloration, being surrounded by a large, transparent, greenish egg capsule. The spawn is deposited separately or in small clumps, xed to submerged plants. One complete egg clutch does not exceed 200 rather sticky eggs. Four collected amplectant pairs deposited 70, 113, 125, and 189 eggs (mean 124 eggs, n = 4). In contrast, one pair of P. minutus collected in Taim (Municipality of Rio Grande, RS) deposited 272 small black and white eggs. Tadpole description. SMNS 9235-37, larvae in developmental stages 34-37 (Gosner, 1960) ( g. 6A). Typical pond tadpole, suspension feeder. Total length between 79 and 87 mm (n = 3). Body proportions in per cent of total length: body length 0.34-0.39; body width 0.19-0.21; body heigth 0.23-0.24. Body ovoid in dorsal and lateral views; widest at midbody, highest at posterior half of body. Eyes lateral, medium sized. Nostrils small, dorsolateral, nearer to snout than to eyes (fronto-nasal distance/naso-ocular distance 0.70.8). Snout broadly rounded in dorsal and lateral views. Mouth small, terminal ( g. 6B);

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Figure 5. Ventral view of the right foot of (A) the holotype of Pseudis cardosoi sp. nov. (MCP 3354), and (B) Pseudis minutus (MCP 3379). Scale bar = 5 mm.

multiple rows of well-developed marginal papillae, dorsally interrupted by a wide gap, being 50-60% of width of oral disc. Labial tooth row formula 1(1-1)/(1-1)2. Median gap in rst tooth row of posterior labium distinct but very small and, due to the xation, slightly overlapping in some specimens (also observed in g. 6B). Median gap in second row of anterior labium large, about equal the lenght of each half of the tooth row (gap considerably smaller in P. minutus). All tooth rows approximately equal in lenght, except third row in posterior labium which is shorter. Vent opening large, triangular, midventral. Spiracle single, sinistral, laterally located, dorsally visible. Tail length about two thirds of total length (60-67%). Tail height greater than body height (tail height/body height 1.1-1.2). In lateral view, dorsal n, ventral n, and tail musculature about equal in height. Dorsal n extending from midbody (at eye level) to tip of tail, with margin uniformly curved. Ventral

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Figure 6. Tadpole of Pseudis cardosoi sp. nov., (A) lateral view at stage 37 (Gosner, 1960) (SMNS 9237), scale bar = 10 mm; (B) oral disc (SMNS 9237), due to the xation process, the median gap in the rst tooth row of the posterior labium is slightly overlapping, scale bar = 1 mm.

n extending from vent to tip of tail, with margin at anterior portion convexe, at posterior portion slightly concave. Tip of tail agelliform. Coloration of body in preservative pale brown or grey with some light reticulations. Posterior portion of tail dark brown to black. Coloration generally much darker and more uniform than in P. minutus, without clearly de ned whitish stripes on body or tail.

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Table 3. Call parameters of Pseudis cardosoi and Pseudis minutus . n Air temperature (time) 19 C (0:20 h) 10 C (1:00 h) 22 C (0:30 h) 14.5 C (20:30 h) 17 C (23:30 h) Call duration in msec (mean) 180-300 (230) 250-450 (380) 80-100 (90) 110-130 (120) 100-150 (130) Notes per call 11-22 Notes per 100 msec 7-8

Axel Kwet

Dominant frequency (kHz) 1.7-2.3

Pseudis cardosoi So Francisco de Paula 4 Nov 1995; SMNS 9001:3 Pseudis cardosoi So Francisco de Paula 7 Oct 1997; MCP 2590 Pseudis minutus Caapava do Sul 21 Mar 1996; SMNS 9124 Pseudis minutus Rio Grande 2 Oct 1997; MCP 3351 Pseudis minutus Isla Talabera, Argentina 17 Dec 1997; MCP 3491

13-20

3-6

1.5-2.3

10-13

11-12

2.5-2.7

12-14

10-12

1.9-2.6

10-16

10-12

2.6-3.1

n indicates number of calls analyzed from the same individual.

Call descriptions. Advertisement calls of Pseudis cardosoi ( g. 7A) were recorded at the Pr-Mata reserve in Rio Grande do Sul, at air temperatures of 10 C and 19 C (table 3). The loud, modulated, variable call resembled the grunting of a pig. One call consisted on average of 11 to 22 notes and had a uctuating dominant frequency at peak amplitudes ranging from 1.5 to 2.3 kHz. The call duration was highly variable but at lower temperatures the calls were more prolongued and had lower rates of notes per msec. The advertisement calls of P. minutus ( gs. 7B, C) were super cially similar to those of P. cardosoi . However, they were much shorter with higher dominant frequencies, fewer notes per call, and higher note rates (table 3). Recordings were analyzed from two Municipalities of Rio Grande do Sul, Caapava do Sul and Rio Grande, and from one place near the type locality of Lysapsus mantidactylus in Argentina (Isla Talabera, 100 km north of Buenos Aires). Furthermore, calling specimens of Pseudis were heard at Osrio, 80 km from the type locality of P. meridionalis, on 21 December 1997. Their calls sounded equal to those of P. minutus. However, the bad quality of the recordings did not allow the analyzing of these calls. Natural history. Pseudis cardosoi lives on open grassland (campos) of the Araucaria plateau, at about 700 to 1100 m a.s.l., not occurring in forested areas. It inhabits mainly permanent pasture ponds and still-water pools of slowly owing creeks. The new pseudid is a strictly aquatic species which I have never seen leaving the water. Its migration success seems to be very limited, as two formerly inhabited ponds next to other Pseudis

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Figure 7. Audiospectrograms of the advertisement calls of (A) Pseudis cardosoi ; air temperature 19 C; PrMata, So Francisco de Paula, Brazil; 4 November 1995, 0:20; voucher specimen SMNS 9001:3. (B) P. minutus ; air temperature 22 C; Guaritas, Caapava do Sul, Brazil; 21 March 1996; 0:30; voucher specimen SMNS 9124. (C) P. minutus from near the type locality of Lysapsus mantidactylus ; air temperature 17 C; Isla Talabera, 100 km north of Buenos Aires, Argentina; 17 December 1997; 23:30; voucher specimen MCP 3491.

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populations which dried out in the hot November 1995 did not become repopulated in the following three years. The occurrence of P. cardosoi was correlated with aquatic vegetation. Of two comparable ponds at the same location, only differing in the presence or absence of aquatic plants, the species was only found in the vegetated pond. The breeding activity of P. cardosoi extended throughout the entire warm season which lasts on the Araucaria plateau from about September to May. Activity depended on air temperatures higher than 8-10 C. Tadpoles were found throughout the year since they develop slowly and often overwinter. Recently metamorphosed juveniles of P. cardosoi were frequently observed from February through April. They measured 30 to 34 mm (n = 4; SMNS 9098, 9213, 9233-34), whereas juveniles of P. minutus were much smaller, measuring 21 to 25 mm (n = 3; MCP 2578, 3349, ZMH A03368). Males of P. cardosoi called by day and night, generally with higher frequencies at night. Other anurans that called simultaneously in permanent ponds were Hyla minuta, H. faber, H. pulchella, Scinax eringiophilus, S. squalirostris, Bufo ictericus, and Leptodactylus ocellatus. In slowly owing creeks I observed the new species in sympatry with Hyla leptolineata. While calling, the males assumed a oating position on the water surface, only the eyes, nostrils, and the paired in ated vocal sac protruding ( g. 1B). If plants were present, they grasped them with their ngers to x themselves. Males called from all places within the pond although they generally preferred deeper and more vegetated sites. Pseudis cardosoi is an opportunistic feeder, catching any small object moving around its head, mainly insects falling into the water. The excrements of two captured specimens contained remains of one large orthopteran, one large cicada, four coleopterans (two Staphylinidae, two Scarabaeidae), one small dragon y (Zygoptera), and one water bug (Naucoridae).

Discussion Pseudis minutus was described from a single specimen collected by Charles Darwin in South America but remained unknown to most herpetologists for more than a century. Even authors who reviewed the genus (Miranda-Ribeiro, 1926; Savage and Carvalho, 1953; Gallardo, 1961) based their knowledge about P. minutus on the original description without examination of specimens. Gallardo (1961) considered the species as dif cult to allocate within the Pseudidae. Part of his doubts was founded by Boulenger (1882) who erroneously reported the holotype with a single vocal sac whereas Gnther (1859 1858) in his original description stated correctly a paired one (con rmed by personal examination of the holotype in April 1998). Hence, referring to Boulenger (1882), Gallardo (1961) mentioned the possibility that P. minutus is only a southern race of P. paradoxus which possesses a single vocal sac. Gallardo (1961) also revalidated Lysapsus mantidactylus which is characterized by a paired vocal sac and was synonymized with P. paradoxus by Savage and Carvalho (1953). Gallardo (1961) treated L. mantidactylus as a widely distributed pseudid and remarked that much material previously identi ed as P. minutus

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or P. meridionalis actually belongs to this species. Other authors (Braun and Braun, 1980; Cei, 1980) adopted this view, without resolving the status of P. minutus. Another pseudid, P. brasiliensis, distinguished from three Brazilian specimens (ZMB 3184-85) by Wiegmann (cited in Peters, 1863), is not valid as no description exists (nomen nudum, Klappenbach, 1985). Peters (1863) considered these specimens as P. minutus. Later, Klappenbach (1985) clari ed this taxonomic confusion within the Pseudidae, showing that in fact L. mantidactylus (type locality Buenos Aires; Cope, 1862), P. meridionalis (type locality restricted to Torres, northeastern Rio Grande do Sul by Bokermann, 1966), and P. minutus (type locality restricted to Maldonado, Uruguay by Klappenbach, 1985) belong to the same species. As P. minutus is the most ancient name it has priority over the others. This synonymization was supported by S and Lavilla (1997) who showed that the description of the P. minutus tadpole agrees with that of L. mantidactylus (Fernndez and Fernndez, 1921; Gallardo, 1964). My own data con rm that the mentioned taxa belong to the same species whereas P. cardosoi represents a new one. Sonagrams of L. mantidactylus from near the type locality ( g. 7C; Barrio, 1970) match with those of P. minutus from Rio Grande do Sul ( g. 7B), whereas the sonagrams of P. cardosoi ( g. 7A) are different, resembling those of P. paradoxus from Argentina (Barrio, 1970) and Bolivia (Bosch et al., 1996). The comparison of eggs, larvae, and recently metamorphosed juveniles also revealed clear differences. Unlike the tadpoles of P. minutus (S and Lavilla, 1997) those of P. cardosoi ( gs. 6A, B) showed a dark pigmentation on the posterior portion of the tail, resembling L. limellus (Kehr and Basso, 1990) and P. paradoxus (Dixon et al., 1995). This pattern functions as antipredator mechanism (Caldwell, 1982, 1986). Finally, all examined specimens from near the type localities of P. minutus, P. meridionalis, and L. mantidactylus agreed among each other in external morphology, including the holotype of P. minutus (BM 1947.2.25.96). Besides P. minutus, P. cardosoi is the second known pseudid from Rio Grande do Sul. After the synonymization by Klappenbach (1985), most distributional data of L. mantidactylus in the amphibian list of Braun and Braun (1980) must be treated as those of P. minutus. However, their citations from the Araucaria plateau (regions 8 and 10) presumably represent P. cardosoi . The latter species seems to be restricted to the Serra Geral, whereas P. minutus occupies a large area extending from the Argentine provinces of Buenos Aires, Entre Ros, Santa F and Corrientes (Gallardo, 1964), to Uruguay (Klappenbach, 1985), the southern, central and northeastern coastal Rio Grande do Sul (Braun and Braun, 1980), up to southeastern Santa Catarina (Boulenger, 1888). A recently collected specimen from Tenente Portela (MCP 3159) extends the distributional range of P. minutus to the extreme north of Rio Grande do Sul and implies an occurrence in the close Argentine province of Misiones, as previously reported by Berg (1896) but doubted by Gallardo (1964). In addition to P. cardosoi and P. minutus, a third pseudid species should occur in Rio Grande do Sul. Records for Lysapsus limellus exist from either the neighbouring Argentine province Corrientes (directly at the border; Cspedez et al., 1995) and Uruguay (some kilometers from the border; Klappenbach and Langone, 1992). Furthermore, the occurrence of Pseudis

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paradoxus platensis was suggested by P.C. Braun in an unpublished draft of his amphibian list. However, the nearest place from which this species is known lies in Corrientes (Cspedez et al., 1995), 150 km from the border to Rio Grande do Sul.

Acknowledgements. I thank W. Maier and W. Engels (Tbingen) for providing the opportunity for my eldwork. I am grateful to M. Di-Bernardo, P.C.A. Garcia and G.M.F. Pontes (Porto Alegre) for the support in Brazil, and to B.T. Clarke (London) for the loan of type material collected by Charles Darwin. A. Schlter kindly made the sonagrams. For commenting on the manuscript I acknowledge J. Hallermann (Hamburg), Christiane Weirauch (Tbingen), and three anonymous reviewers. The work was supported by grants of the LGFG, SHIFT, and DAAD.

References
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A new species of Pseudis

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Appendix: additional specimens examined


Pseudis cardosoi sp. nov. Brazil: Rio Grande do Sul: Bom Jesus: Ausentes (MCP 3347-48, 3353; SMNS 9213); Cambar do Sul (MCP 469, 482); So Francisco de Paula: Pr-Mata (MCP 1792-93, 1809, 1848, 3371, 3373-75, 3378; SMNS 9001:1-4, 9099-9100, 9107, 9163; ZMH A03317, A03366-67); Vacaria (MCP 3538). Pseudis minutus Brazil: Rio Grande do Sul: Caapava do Sul: Guaritas (MCP 2582-87, 2589, 3368-69, 3376-77, 3379; SMNS 9118-24; ZMH A03318); Mostardas (MCP 1120, 1123, 1125-26); Palmares do Sul: Capivari (MCP 3352, SMNS 9166); Porto Alegre (MCP 1561-62; ZMH A03369, ZMB 31092); Rio Grande: Taim (MCP 3195-96, 3351, SMNS 9152); Santo Antnio da Patrulha (MCP 1603-07); Tenente Portela (MCP 3159); Viamo (MCP 9981003). Santa Catarina: Sombrio (MCP 191). Argentina: Buenos Aires: Isla Talabera (MCP 3491-92). Pseudis paradoxus Brazil: Mato Grosso do Sul: Aquidauana (MCP 1448). Argentina: Corrientes: Bella Vista (SMNS 8636). Received: July 20, 1998. Accepted: April 7, 1999.