Pseudopaludicola murundu Toledo, Siqueira, Duarte, Veiga-Menoncello, Recco-Pimentel & Haddad, 2010, sp. nov.
Creators: name: Toledo, Luís Felipe
name: Siqueira, Sérgio
name: Duarte, Thiago C.
name: Veiga-Menoncello, Ana Cristina P.
name: Recco-Pimentel, Shirlei M.
name: Haddad, Célio F. B.
Publisher: name: Zenodo
Year: 2010
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Amphibia
Anura
Leiuperidae
Pseudopaludicola
Pseudopaludicola murundu

description: Pseudopaludicola murundu sp. nov.
Holotype. CFBH 8241, an adult male (Fig. 1) collected in a temporary pond in the “Sítio do Carlão”, district of Itapé, Municipality of Rio Claro, state of São Paulo, southeastern Brazil (22°20'41.57"S; 47°40'1.52"W; approximately 600 m above sea level), by L. F. Toledo, J. G. R. Giovanelli, and F. B. Britto, on 0 9 January 2005.
Paratopotypes. Eleven males: CFBH 8235-40; 8242, collected on 0 9 January 2005 by L. F. Toledo, J. G. R. Giovanelli, and F. B. Britto; ZUEC 14286-87; 14289-90, collected on 31 January 2007 by L. F. Toledo, S. Siqueira, and E. F. Favero; and four females: CFBH 8243, collected on 0 9 January 2005 by L. F. Toledo, J. G. R. Giovanelli, and F. B. Britto; ZUEC 14284-85, 14288, collected on 31 January 2007 by L. F. Toledo, S. Siqueira, and E. F. Favero.
Diagnosis. Pseudopaludicola murundu is assigned to the genus Pseudopaludicola due to the presence of hypertrophied antebrachial tubercle (see Lynch, 1989). Pseudopaludicola murundu is a medium-sized species for the genus and is characterized by the following combination of characters: (1) single, dark, and subgular vocal sac with dark longitudinal folds in males; (2) immaculate to light brown, nuptial pads in males that cover the external part of finger I, from the wrist to the second phalanx; and (3) presence of 11 pairs of chromosomes without heteromorphic sex chromosomes (XX/XY).
Comparison with other species. Pseudopaludicola murundu is distinguished from all other species of the genus (except from P. saltica) by having very long hindlimbs, with tibio-tarsal articulation reaching beyond the end of the snout (in other species, except P. saltica, the tibio-tarsal articulation reach, at most, the nostrils). From species of the P. pusilla species group it is also distinguished by the absence of T-shaped terminal phalanges. From P. saltica it is distinguished by having the nostrils closer to the apical snout region (in P. saltica this distance is larger: mean snout-nostril distance = 1.1 ± 0.25 in males of P. murundu and 1.35 ± 0.21 in males of P. saltica), larger and clearer nuptial pads in adult males collected during peak of the breeding season (smaller and darker in adult males P. saltica collected during peak of the breeding season), longitudinal folds in the vocal sac (absent in P. saltica), and a darker vocal sac. Furthermore, it can be distinguished from P. saltica by having the chromosome pair 8 homomorphic between females and males, whereas in P. saltica this chromosome pair is heteromorphic in males, which can be considered a XX/XY sex chromosome system (Duarte et al., 2010). In addition, the sequence divergence computed with MEGA Ver. 4.0 (Tamura et al. 2007) based on mitochondrial DNA (a fragment of Cyt-b) showed a higher degree of nucleotide divergence between P. m u r u n d u and P. saltica (8%) when compared to other close related species, P. ameghini – sensu Cope, 1887 and P. ternetzi (5%) (Veiga-Menoncello et al. unpublished data).
Description of the holotype. Body elliptic and broad. Head triangular, longer than wide. Snout subelliptical in dorsal view and rounded in profile (Figure 2 A, 2B). Nostrils slightly protuberant, directed anterolaterally. Mouth opening ventral. Canthus rostralis rounded. Loreal region slightly concave. Choanae rounded. Eye protuberant, its diameter larger than interorbital distance. Interorbital area flat. Tympanum indistinct. Vocal sac single, externally expanded, large, and with longitudinal folds; vocal slits present. Vomerine teeth absent. Tongue elliptical, longer than wide. Finger length I<II<IV<III. Toe length I<II<V<III<IV. Finger and toe tips without disks (Figure 2 C, 2D). Thumb with keratinized beige nuptial pad. Finger webbing absent and toe webbing reduced I-II 2- 3 III 3-4 IV-V. Finger and toe subarticular tubercles conical and single. A large subconical ulnar tubercle. Few rounded supernumerary tubercles in the hand in the area delimited by the first subarticular tubercles, the elliptical internal metacarpal tubercle, and the ovoid external metacarpal tubercle. Hindlimbs very robust and long. Thigh shorter than tibia; foot slightly longer than thigh and slightly shorter than tibia. Supernumerary tubercles absent in the foot. Metatarsal tubercles present, elliptical; internal larger than the external; external more protuberant than the internal. A well developed fold from the internal metatarsal tubercle to the mid-ventral tarsus. Skin of belly smooth; ventral surface of thigh granular. Dorsum of head, body, and limbs smooth with scattered tubercles; the skin on the scapula region has two arc shaped granular folds. Flanks with scattered tubercles. Cloacal region granular. Measurements of the holotype are presented in table 1.
Holotype Paratypes
In life, dorsum and limbs are brown with dark brown spots or stripes; vertebral line yellowish brown from the snout to vent; belly white; ventral view of legs yellowish white and vocal sac beige with dark dots; iris golden superiorly and brownish inferiorly (see Figure 1). In preservative, dorsum and dorsal part of limbs brown with dark brown spots or stripes; vertebral line beige from the snout to vent; belly light beige; legs yellowish white; vocal sac beige with dark dots and with dark longitudinal folds.
Variation. Females slightly larger than males, lacking vocal sac and nuptial pads (present in males). The nuptial pad can be brownish to light cream color in males. The snout shape in lateral view varies from rounded to slightly truncate. In 75% of the individuals (12 out of 16 examined) the vertebral line is absent. In life this line can be white, yellow, or red. In life, the two arc shaped granular folds on the scapula region are generally darker than the general coloration of dorsum (see also Figure 1). Dorsal pigmentation varies among individuals, from brown to grayish. The number and form of the dorsal spots also varies, which can be small scattered brown dots up to large brown stripes over a lighter general dorsal coloration.
Natural history notes. Males were observed calling in open areas of a private farm. The region was previously a mosaic composed of Cerrado (Brazilian savannah) and semidesciduous Atlantic forest. Males called from the ground near shallow (about 2–3 cm of depth) slow-flowing water bodies, on the “murundus”, which are small elevations on the ground (about 10 cm height), covered by grassy and/or arboreal vegetation. Calling males were spaced for at least 5 cm in distance, and it was possible to observe silent males even closer to the calling ones (however, we could not confirm if these males were performing satellite behavior). Males were observed calling at dawn and during the first hours of the night. One amplectant pair was observed and collected in the same microhabitat of the calling site. Egg laying was not observed, so we could not attribute any reproductive mode to P. murundu. Tadpoles were active at night in the shallow and slow-flowing water bodies, where males and the amplectant pair were observed. No other species was observed in syntopy.
Geographic distribution. The new species is known only for the type locality, which is an open grassland field of anthropogene origin used for cattle ranching in a particular farm.
Etymology. The specific name is a Tupi word that means small elevations on the ground, covered by grassy and/or arboreal vegetation. This is the most common calling site of this species.
Advertisement call. The advertisement call of Pseudopaludicola murundu is composed of groups of notes (each with three to six pulses) that are emitted in mean intervals of 92 ms (Figure 3 A and 3B). The calls are generally long, with more than 10 seconds of duration (Figure 3 A) and high pitched, with most of the frequencies higher than 5 kHz. The mean rate of notes emitted per minute is 225. Males can call in antiphony as the interval between notes are similar to the duration of notes. The first pulse of the note generally occupies lower frequencies than the last one. The intermediary pulses occupy the intermediary frequencies between the first and the last pulses. Further information of the numerical characteristics of the advertisement call is provided in Table 2.
Parameter Values Dominant frequency (kHz) 5.72 ± 0.32 (n = 10) (5.17 – 6.37) Minimum frequency (kHz) 5.03 ± 0.20 (n = 10) (4.71 – 5.33) Maximum frequency(kHz) 6.82 ± 0.26 (n = 10) (6.44 – 7.25) Notes per call 45.60 ± 27.39 (n = 14) (15 – 82)
Pulses per note 5.07 ± 0.49 (n = 16) (3 – 6)
Interval between notes of the same call (s) 0.09 ± 0.03 (n = 14) (0.03 – 0.13) Interval between pulses of the same note (ms) 3.36 ± 4.41 (n = 14) (00 – 12) Tadpoles. Larvae were obtained in shallow ponds at the type locality. As there was no other amphibian species in syntopy with P. murundu we assigned these tadpoles to this species (this tadpole confirm with general morphology of Pseudopaludicola tadpoles), however this identification is preliminary and not verified. The following description is based on seven tadpoles (CFBH 22670) in developmental stages 37-39 (Gosner 1960). Body elliptical in dorsal and ventral views (Figure 4 A,C), depressed/globular with flattened venter (Figure 4 B); body wider than high; snout rounded; eyes medium-sized, dorsolateral; nostrils dorsal, small, and reniform; nostrils closer to the eyes than to the tip of snout; spiracle sinistral, its opening in the middle of body; cloacal tube large, medial; caudal musculature robust; dorsal fin originating on the anterior part of the tail; dorsal fin wider than ventral fin. Oral disc directed ventrally, emarginated, and bordered by one row of well developed marginal papillae, interrupted along a large area on the anterior labium; tooth row formula 2(2)/2 or 2(2)/2(1); upper and lower jaw sheathes strongly developed and serrate (Figure 4 D). In preservative, dorsum pale brown; throat and belly transparent; caudal musculature with very scattered pale brown pigmentation; fins transparent with scattered pale brown pigmentation.
Three tadpoles in developmental stage 37 measured (range, average, SD) total length 24.79–27.10 (25.65, 1.26); body length 8.05–8.90 (8.41, 0.44); maximum body height 3.67–4.36 (3.97, 0.36); maximum body width 5.02–5.68 (5.27, 0.36); internarial distance 1.24–1.27 (1.26, 0.02); interorbital distance 1.05–1.06 (1.06, 0.01); eye-nostril distance 0.39–0.49 (0.45, 0.05); eye diameter 1.26–1.33 (1.28, 0.04).
Karyotype description. The karyotype of the P. murundu has 2n = 22 chromosomes. Pairs 1, 2, 5, 7, 9, 10, and 11 are metacentrics, pairs 3, 4, and 6 submetacentrics, and pair 8 telocentric (Figure 5 A–C). Besides the centromeric heterochromatic blocks, positive C-bands were detected in the interstitial regions on the long arms of the pairs 1, 2, 3, 4, and 6, and on the short arms of the pairs 1 and 2 (Figure 5 B). The nucleolar organizing region (NOR) was detected by silver staining (Ag-NOR) in the telomeric region on the long arm of the chromosome 8 (Figure 5 C; Table 3. The homologous of the pair 8, telocentric in both sexes, showed a notable size heteromorphism, and the Ag-NOR site was observed only in the larger homologous of this pair (Figure 5 C) which were identified as 8 and 8’ (Table 3).
Number of chromosomes
description: Published as part of Toledo, Luís Felipe, Siqueira, Sérgio, Duarte, Thiago C., Veiga-Menoncello, Ana Cristina P., Recco-Pimentel, Shirlei M. & Haddad, Célio F. B., 2010, Description of a new species of Pseudopaludicola Miranda-Ribeiro, 1926 from the state of São Paulo, Southeastern Brazil (Anura, Leiuperidae), pp. 38-48 in Zootaxa 2496 on pages 40-46, DOI: 10.5281/zenodo.195717