777 research outputs found
A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961)
No full textRojas-Runjaic, Fernando J. M., Infante-Rivero, Edwin E., Salerno, Patricia E., Meza-Joya, Fabio Leonardo (2018): A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961). Zootaxa 4382 (1): 121-146, DOI: 10.11646/zootaxa.4382.1.
FIGURE 5 in A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961)
No full textFIGURE 5. Color variation of Hyloscirtus japreria sp. nov. in life: adults (a–e), and metamorph (f), all from Cerro Las Antenas, Zulia, Venezuela. Photos: F.J.M. Rojas-Runjaic.Published as part of Rojas-Runjaic, Fernando J. M., Infante-Rivero, Edwin E., Salerno, Patricia E. & Meza-Joya, Fabio Leonardo, 2018, A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961), pp. 121-146 in Zootaxa 4382 (1) on page 130, DOI: 10.11646/zootaxa.4382.1.4, http://zenodo.org/record/118161
FIGURE 9 in A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961)
No full textFIGURE 9. Habitat of Hyloscirtus japreria sp. nov. at Guacharaca Camp, Zulia state, Venezuela (a), and El Manantialito creek, Nuevas Ideas farm, La Guajira department, Colombia (b). Photos: F.J.M. Rojas-Runjaic (a) and F.L. Mesa-Joya (b).Published as part of Rojas-Runjaic, Fernando J. M., Infante-Rivero, Edwin E., Salerno, Patricia E. & Meza-Joya, Fabio Leonardo, 2018, A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961), pp. 121-146 in Zootaxa 4382 (1) on page 134, DOI: 10.11646/zootaxa.4382.1.4, http://zenodo.org/record/118161
FIGURE 11 in A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961)
No full textFIGURE 11. Advertisement call of Hyloscirtus callipeza. Oscillogram (a) and spectrogram (b) of a 5 s fragment of note trill. Detailed view of the oscillogram (c) and spectrogram (d) of a 0.5 s long section of the same recording.Published as part of Rojas-Runjaic, Fernando J. M., Infante-Rivero, Edwin E., Salerno, Patricia E. & Meza-Joya, Fabio Leonardo, 2018, A new species of Hyloscirtus (Anura, Hylidae) from the Colombian and Venezuelan slopes of Sierra de Perijá, and the phylogenetic position of Hyloscirtus jahni (Rivero, 1961), pp. 121-146 in Zootaxa 4382 (1) on page 138, DOI: 10.11646/zootaxa.4382.1.4, http://zenodo.org/record/118161
FIGURE 5 in A new frog of the genus Aromobates (Anura, Dendrobatidae) from Sierra de Perijá, Venezuela
No full textFIGURE 5. View of the stream of Kusare waterfall (tributary of Río Tokuko), one of the localities inhabited by Aromobates tokuko sp. nov. All other localities are similar in aspect.Published as part of Rojas-Runjaic, Fernando J. M., Infante-Rivero, Edwin E. & Barrio-Amorós, César L., 2011, A new frog of the genus Aromobates (Anura, Dendrobatidae) from Sierra de Perijá, Venezuela, pp. 37-50 in Zootaxa 2919 on page 47, DOI: 10.5281/zenodo.27790
Tachiramantis lassoalcalai Rojas-Runjaic & Echevarría & Becerra-Rondón & Infante-Rivero 2020
No full textThe evolutionary relationships of Tachiramantis within Craugastoridae are yet unclear as its position varied in all previous phylogenetic hypotheses in which samples of this genus were included (Heinicke et al. 2015; Guayasamin et al. 2017; Heinicke et al. 2018). However, all of these hypotheses were congruent in recovering Tachiramantis as monophyletic, not closely related to Pristimantis, and deeply divergent to its sister linage. Our phylogeny (Fig. 2) also differs from previous hypotheses with regard to the intergeneric relationships of Tachiramantis with other craugastorids but it is concordant with the results of Heinicke et al. (2015) and Guayasamin et al. (2017) in recovering the three species of Tachiramantis as closely related, with T. lentiginosus and T. douglasi as sister species, and T. prolixodiscus as the most external of the genus. The only difference with these two previous hypotheses corresponds to the position of the newly included P. lassoalcalai, which is well-nested within Tachiramantis as sister to T. lentiginosus + T. douglasi (rendering Tachiramantis paraphyletic). This newly discovered phylogenetic position confirms previous suspicions of a close evolutionary relationship between this species and Tachiramantis based on its overall morphological resemblance with T. lentiginosus and T. douglasi. Based on this result and in order to promote a taxonomy based on monophyletic groups, we transfer Pristimantis lassoalcalai Barrio-Amorós, Rojas-Runjaic & Barros, 2010 to Tachiramantis as Tachiramantis lassoalcalai comb. nov. With the new addition, Tachiramantis is now composed by four species distributed on the northern part of the Cordillera Oriental of Colombia, Tamá massif, Sierra de Perijá and Cordillera de Mérida. Tachiramantis remains a poorly known genus, distributed in a region undersampled by systematic studies using molecular data (Heinicke et al. 2015) and with vast areas poorly inventoried for amphibians, as the Sierra de Perijá (Rojas-Runjaic et al. 2011; Meza-Joya 2016; Rojas-Runjaic et al. 2018) and the Tamá massif (Acevedo et al. 2014; Meza-Joya et al. 2019). Thus, we expect that the inclusion of both new and already described terraranas currently allocated to Pristimantis, in future molecular phylogenies, will result in the discovery of additional members of Tachiramantis. On the other hand, although no external morphology synapomorphies are known for Tachiramantis, the occurrence of pale spots on the posterior surface of thighs (Rivero 1984) is shared by T. douglasi, T. lassoalcalai, and T. lentiginosus (absent in T. prolixodiscus). Thus, this character state can be useful to diagnose these three species from other sympatric terraranas by external morphology. The phylogenetic relevance of this phenotypic character should be objectively assessed. In addition, osteological characterization of T. lassoalcalai will be required in order to corroborate the occurrence in this species of the putative cranial synapomophies defined by Heinicke et al. (2015) for Tachiramantis. As it is known, acoustic signals play a critical role in mate recognition and mate choice, and in most cases are species-specific (Wells 2007), which make them a useful source of evidence for anuran systematics and taxonomy (Khöler et al. 2017). Until very recently there was no bioacoustic information available for any Tachiramantis, but the almost simultaneous publication of the call description of T. douglasi (Mendoza-Roldán et al. 2020) with our description of the advertisement call of T. lassoalcalai allow us to confirm that bioacoustics also constitutes a valuable tool to diagnose species in this group. These two species exhibit strikingly different advertisement calls (values for T. lassoalcalai in parentheses): T. douglasi calls last ca. 1.7 s and consist of trills of 7–9 notes, with the first being the longest (calls represented by single notes of ca. 0.7 s), with dominant frequency of ca. 5.1 kHz (dominant frequency between 2.9–3.0 kHz). Considering that the known geographic distributions of T. lassoalcalai and T. douglasi are relatively close (Mendoza-Roldán et al. 2020; Barrio-Amorós et al. 2010) and that both species are morphologically similar (see Fig. 1), their bioacoustical characterizations will be valuable to diagnose them during fieldwork and in the absence of molecular data.Published as part of Rojas-Runjaic, Fernando J. M., Echevarría, Lourdes Y., Becerra-Rondón, Adriana C. & Infante-Rivero, Edwin E., 2020, Tachiramantis lassoalcalai (Barrio-Amorós, Rojas-Runjaic & Barros, 2010) (Anura, Craugastoridae): a new combination revealed by molecular evidence, with a description of its advertisement call, pp. 372-382 in Zootaxa 4778 (2) on pages 375-376, DOI: 10.11646/zootaxa.4778.2.8, http://zenodo.org/record/397479
FIGURE 3 in A new non-sexually dichromatic species of the genus Gonatodes (Sauria: Sphaerodactylidae) from Sierra de Perijá, Venezuela
No full textFIGURE 3. Dorsal view of Gonatodes lichenosus sp. nov. (male holotype MHNLS 19116), showing clusters of enlarged, conical to spinelike scales on the flanks of neck and trunk. White arrows indicate the clusters of scales magnified in upper right and lower left frames.Published as part of Rojas-Runjaic, Fernando J. M., Infante-Rivero, Edwin E., Cabello, Pedro & Velozo, Pablo, 2010, A new non-sexually dichromatic species of the genus Gonatodes (Sauria: Sphaerodactylidae) from Sierra de Perijá, Venezuela, pp. 1-16 in Zootaxa 2671 on page 4, DOI: 10.5281/zenodo.19915
FIGURE 3 in Tachiramantis lassoalcalai (Barrio-Amorós, Rojas-Runjaic & Barros, 2010) (Anura, Craugastoridae): a new combination revealed by molecular evidence, with a description of its advertisement call
No full textFIGURE 3. Advertisement call of Tachiramantis lassoalcalai comb. nov. Oscillogram (a) and spectrogram (b) of a 5 s fragment of a call group. Detailed view of the oscillogram (c) and spectrogram (d) of a 0.5 s long section of the same recording depicting a single call (= note).Published as part of Rojas-Runjaic, Fernando J. M., Echevarría, Lourdes Y., Becerra-Rondón, Adriana C. & Infante-Rivero, Edwin E., 2020, Tachiramantis lassoalcalai (Barrio-Amorós, Rojas-Runjaic & Barros, 2010) (Anura, Craugastoridae): a new combination revealed by molecular evidence, with a description of its advertisement call, pp. 372-382 in Zootaxa 4778 (2) on page 378, DOI: 10.11646/zootaxa.4778.2.8, http://zenodo.org/record/397479
Hyloscirtus japreria Rojas-Runjaic & Infante-Rivero & Salerno & Meza-Joya 2018, sp. nov.
No full textHyloscirtus japreria sp. nov. (Figs. 2–3, 5–6, 10) Suggested common name in English: Perijá’s Stream Frog Suggested common name in Spanish: Rana Torrentícola de Perijá Hyloscirtus sp.— Rojas-Runjaic et al. (2016: 36) Holotype. Adult male, MHNLS 19236 (field number SP 294; Figs. 2–3), from Guacharaca Camp, Tetari Kopejoacha creek, Rio Negro basin, Sierra de Perijá, Machiques de Perijá municipality, Zulia state, Venezuela (10°04’21.9 N – 72°51’16.7 W; elevation 1,661 m asl), collected on 27 May 2009, by Fernando J. M. Rojas-Runjaic and Edwin E. Infante-Rivero. Paratypes. 21 specimens (16 males and five females): Two adult males, MHNLS 18837–18838 (field numbers SP 44–45) from the creek behind the house at Cerro Las Antenas, Sierra de Perijá, Rosario de Perijá municipality, Zulia state, Venezuela (10°20’37.0”N – 72°33’41.0”W; 1,430 m asl), collected on 27 March 2008, by Fernando J. M. Rojas-Runjaic; two adult males, MHNLS 18867, 18869 (field numbers SP 74, 76) and one adult female, MHNLS 18868 (field number SP 75), from the same locality, collected on 28 March 2008, by Fernando J. M. Rojas-Runjaic, Edwin E. Infante-Rivero, Pablo Velozo and Paul Granado; two adult males, MHNLS 18971–18972 (field numbers SP 117–118), from the same locality, collected on 10 July 2008, by Fernando J. M. Rojas-Runjaic, Pedro Cabello, Kripsy Herrera and Arlene Cardozo. One male, MHNLS 22568 (field number SP 218), from the same locality, collected on 26 April 2009, by Fernando J. M. Rojas-Runjaic, Adriana Becerra and Arnaldo Ferrer. Two adult males, MHNLS 18888–18889 (field numbers SP 95–96) from the creek near to second antenna, Cerro Las Antenas, Sierra de Perijá, Rosario de Perijá municipality, Zulia state, Venezuela (10°19’40.0”N – 72°35’27.0”W; 1,832 m asl), collected on 29 March 2008, by Fernando J. M. Rojas-Runjaic and Pedro Cabello; one adult male, MHNLS 18988 (field number SP 134), from the same locality, collected on 12 July 2008 by Fernando J. M. Rojas- Runjaic and Pedro Cabello. Two adult females, MHNLS 19166–19167 (field numbers SP 224–225), and one adult male, MHNLS 19168 (field number SP 226) with the same locality and collectors as the holotype, collected on 22 May 2009; two males, MHNLS 19234, 19237 (field numbers SP 292, 295), and one female, MHNLS 19235 (field number SP 293) with the same locality, date and collectors as the holotype. Three males, UIS-A 5496–5498 (field numbers NG 277–279), and one female, UIS-A 5495 (field number NG 276) from El Manantialito creek, Nuevas Ideas farm, Sierra de Perijá, vereda Barriales-Nuevas Ideas, El Molino municipality, La Guajira department, Colombia (10°35’15.87”N – 72°49’06.90”W; 1,754 m asl), collected on 19 July 2012, by Fabio L. Meza-Joya. Referred specimens. Three specimens (two juveniles and one metamorph): juvenile MHNLS 18858 (field number SP 65) from the creek behind the house at Cerro Las Antenas, Sierra de Perijá, Rosario de Perijá, municipality, Zulia state, Venezuela (10°20’37.0”N – 72°33’41.0”W; 1,449 m asl), collected on 27 March 2008 by Pedro Cabello and Edwin Infante; juvenile MHNLS 19254 (field number SP 312) with the same locality, date and collectors as the holotype; metamorph UIS-A 5499 (field number NG 280) from El Manantialito creek, Nuevas Ideas farm, Sierra de Perijá, vereda Barriales-Nuevas Ideas, El Molino municipality, La Guajira department, Colombia (10°35’16.0”N – 72°49’12.0”W; 1,962 m asl), collected on 19 July 2012, by Fabio L. Meza-Joya. Definition. A new species of Hyloscirtus, assigned to the H. bogotensis species Group (sensu Duellman 1972, and Faivovich et al. 2005), by its phylogenetic placement (Fig. 1) and morphologically, by the presence of a mental gland in males, and white parietal peritoneum. The new species is defined by the unique combination of the following characters: (1) Adult males smaller than females (males: 28.8–32.7 mm of SVL vs. females: 35.6–39.1 mm); (2) skin texture smooth on dorsum, flanks, throat, chest, forearms, dorsal and inner surfaces of thighs; areolate on ventral surface of thighs and coarsely areolate on belly; (3) body slender (2.4 to 3.0 times longer than wide); (4) snout rounded in dorsal view and profile; (5) tympanum distinct, small (td: 3.4–5.0% of SVL); tympanic membrane not differentiated; tympanic annulus visible through the skin; about 1/4 of the tympanum concealed dorsally by the supratympanic fold; (6) dentigerous processes of vomers conspicuous, straight, narrowly separated from each other; with 5–8 teeth each; (7) vocal slits longitudinal, extending from sides of tongue to the angle of mouth; (8) vocal sac bilobate, median and subgular, moderately distensible and evident externally; (9) mental gland present in males, disc-shaped, small, covering about the anterior third of throat; (10) ulnar fold present; (11) fingers with fleshy dermal fringes; (12) fingers II–IV basally webbed; (13) nuptial pads absent; (14) prepollex swollen, elliptical, not modified as a protruding spine; (15) calcar tubercle absent; (16) outer and inner tarsal folds present, inner poorly defined; (17) toes with wide lateral fringes; (18) toes extensively webbed; (19) supracloacal fold present; (20) cloacal sheath short; (21) cloaca surrounded by numerous warts; (22) in life, dorsum usually pale yellow to greenish yellow, with dense dark brown punctuation, and several reddish brown spots; (23) whitish stripes on external border of upper eyelids and supratympanic folds, longitudinally on the mid-dorsum, on supracloacal fold, outer ulnar folds, inner and outer tarsal folds, and also on dorsal internal surface of shanks; (24) white warts around the cloaca; (25) axillae bright yellow and groins ocher yellow in life; (26) interdigital membranes on hands and feet bright yellow in life; (27) white parietal peritoneum covering ventral surface of abdominal cavity almost entirely; (28) iris gray with thin black reticulation. Diagnosis. Hyloscirtus japreria sp. nov. is readily distinguishable from all other species in the H. armatus and H. larinopygion groups by the presence of mental gland in males (present in almost all species of H. bogotensis species group). Of the 16 species currently included in the H. bogotensis Group only five are known to occur on the northwestern portion of the Andes (northern Cordillera Oriental of Colombia, Sierra de Perijá and Cordillera de Mérida in Venezuela); these are: H. callipeza, H. estevesi, H. jahni, H. lascinius and H. platydactylus (Fig. 4). Hyloscirtus japreria sp. nov. differs from H. callipeza (characters of the latter in parenthesis) by having ulnar, outer and inner tarsal folds (absent), snout rounded in profile (truncate), tympanic annulus distinct (barely visible), skin of belly coarsely areolate (granular), webbing bright yellow in life (pale orange), by having a whitish ulnar stripe (absent), a whitish stripe on dorsal internal surface of shank (absent), and iris gray with thin black reticulation (bronze with thin black reticulation); also adult females of H. japreria sp. nov. are larger than females of H. callipeza (maximum SVL: 39.1 mm in H. japreria sp. nov. vs. 36.7 mm in H. callipeza). These two species also differ in temporal and spectral characteristics of advertisement calls: H. japreria sp. nov. emits much longer trills (79.72– 159.62 s vs. 4.99– 17.50 s), and more notes by trill (421–1,058 vs. 29–102 notes) than H. callipeza; the notes of H. japreria sp. nov. are also longer (0.037– 0.046 s vs. 0.016–0.019) and less spaced (time between notes: 0.101– 0.105 s vs. 0.146– 0.181 s) than those of H. callipeza. The two species also differ in the number of amplitude peaks per note, with a single peak in H. japreria sp. nov. (Fig. 7c) and three peaks of amplitude per note in H. callipeza (Fig. 11c). Finally, both peak and fundamental frequencies are lower in H. japreria sp. nov. than in H. callipeza (peak frequency: 3.08–3.15 kHz vs. 3.20–3.33 kHz; fundamental frequency: 1.54–1.59 kHz vs. 1.59– 1.65). Hyloscirtus jahni is known only from Cordillera de Mérida in the Venezuelan Andes (La Marca 1985; Barrio 2004). The new species can be readily distinguished from H. jahni by lacking spicules on the skin in males (spicules present on snout, lips, chin, chest, belly, upper arms, forearms, thighs, shanks, hands, and fingers of adult males), snout not projected beyond lip (snout projected), supratympanic fold distinct (strong), head narrower than body (head wider than body), vocal sac bilobate (single), mental gland small and discoidal (large and transversely elliptical), and adult females larger than those of H. jahni (maximum SVL 39.1 mm in H. japreria sp. nov. vs. 34.4 mm in H. jahni). These two species also differ in temporal characteristics of advertisement calls (parameter values of H. jahni taken from La Marca [1985]): The calls of H. japreria sp. nov. consist of long trills of tonal notes (79.72– 159.62 s) whereas the call of H. jahni is composed of single pulsed notes. Hyloscirtus japreria sp nov. presents a single amplitude peak per note while each note of H. jahni contains twelve peaks of amplitude. The notes of H. japreria sp. nov. are also longer (0.037– 0.046 s vs. 0.013 s) and less spaced (time between notes: 0.101– 0.105 s vs. 0.3 s) than those of H. jahni. Finally, the rate of note emission is higher in H. japreria sp. nov. than in H. jahni (5.30–6.71 vs. 3.0 notes/s). Hyloscirtus japreria and H. jahni are not sister species in our phylogeny (Fig. 1). Hyloscirtus estevesi is known only from his type locality in the Río Albarregas, Mérida state and Boconó, Trujillo state (both localities in the Cordillera de Mérida). This species was originally described as a centrolenid (Centrolenella estevesi) and recently assigned to the H. bogotensis species Group by Faivovich et al. (2005). The new species can be readily distinguished from H. estevesi by not having the snout projected beyond lip (snout projected), by having the head narrower than body (head wider than body), and by having melanophores on dorsal surface of FII–IV (melanophores only present on FIV). Hyloscirtus lascinius occurs on western versant of Cordillera Oriental in Colombia and Cordillera de Mérida in Venezuela (Rojas-Runjaic et al. 2016). Hyloscirtus japreria sp. nov. also is readily distinguishable from this species by not having calcar tubercle (present, large and triangular), by having tympanic annulus distinct (barely visible), snout rounded in profile (snout sloped in profile), skin on belly coarsely areolate (granular), dorsum not sprinkled with whitish dots (dorsum sprinkled with whitish dots), by having, in life, axillary and inguinal regions bright yellow and bright ocher, respectively (axillary and inguinal regions greenish blue), supratympanic fold delineated with a whitish stripe (supratympanic fold without a whitish stripe), and iris gray with thin black reticulation (iris golden with thin reddish reticulation). These two species also differ in temporal and spectral characteristics of advertisement calls (parameter values of H. lascinius taken from Rojas-Runjaic et al. [2016]): H. japreria sp. nov. emits longer note trills (421–1,058 notes by trill) whereas H. lascinius emits single notes or groups of two to four notes. The notes of H. japreria sp. nov. are shorter (0.037– 0.046 s vs. 0.091–0.164) and less spaced (time between notes: 0.101– 0.105 s vs. 0.700– 1.075 s) than those of H. lascinius. Finally, both peak and fundamental frequencies are higher in H. japreria sp. nov. than in H. lascinius (peak frequency: 3.08–3.15 kHz vs. 2.13–2.37 kHz; fundamental frequency: 1.54–1.59 kHz vs. 1.08–1.22). Hyloscirtus lascinius and H. japreria sp. nov. are not sister species in our phylogeny (Fig. 1). Hyloscirtus platydactylus is known from the eastern slope of Cordillera Oriental in Colombia (Acosta-Galvis 2000) and Cordillera de Mérida in Venezuela (La Marca 1985, Barrio 2004). Hyloscirtus japreria sp. nov. is similar to H. platydactylus, but can be distinguished from the latter by having ulnar, outer and inner tarsal folds (ulnar, outer and inner tarsal folds absent); mental gland small, disc-shaped, covering about the anterior third of throat (larger, rhomb-shaped, covering about the anterior half of throat); eye diameter greater than eye-to-nostril distance (eye diameter equal to eye-to-nostril distance); whitish stripe extending on outer border of the upper eyelid and on supratympanic fold (whitish stripe absent); whitish stripes on supracloacal fold, ulnar fold, outer and inner tarsal folds, and also on dorsal internal surface of shanks (whitish stripes on supracloacal, ulnar, outer and inner tarsal folds, and on dorsal internal surface of shank, absent); dorsal surfaces of fingers I–II and toes I–III immaculate bright yellow in life (ocher yellow, densely stippled with blackish brown); dorsal surfaces of finger discs and toe discs spotless (spotted with blackish brown); iris gray with thin black reticulation (golden to copper with black reticulation). These two species also differ in temporal and spectral characteristics of advertisement calls (parameter values of H. platydactylus taken from Duellman [1972] and La Marca [1985]): The notes emitted by H. japreria sp. nov. are longer (0.037– 0.046 s vs. 0.020) and less spaced (time between notes: 0.101– 0.105 s vs. 0.140 s) than those of H. platydactylus. The two species also differ in the maximum rate of note emission, that is lower in H. japreria sp. nov. than in H. platydactylus (6.71 vs. 9.00 notes/s). Peak frequency is also the fundamental in H. platydacylus (2.50–3.49 kHz) and includes in its variation the peak frequency of H. japreria sp. nov. (3.08–3.15 kHz), but fundamental frequency of this last species is lower than that of H. platydacylus (1.54– 1.59 vs. 2.50–3.49 kHz). Hyloscirtus platydactylus and H. japreria sp. nov. are not sister species in our phylogenetic analysis (Fig. 1). From the remaining species of the Hyloscirtus bogotensis species Group, H. japreria sp. nov. is also readily distinguishable by the following characters: from H. denticulentus, H. palmeri and H. piceigularis by lacking calcars (present in all these three species); from H. albopunctulatus and H. phyllognathus by lacking white spots on the dorsum (present in both species); from H. alytolylax and H. colymba by having an inner tarsal fold delineated with a whitish stripe (absent in both species); from H. lynchi by having tarsal folds and mental gland evident externally in males (tarsal folds absent and mental gland not evident in H. lynchi), from H. bogotensis by having tarsal and supracloacal whitish stripes (both stripes absent in H. bogotensis); from H. simmonsi and H. torrenticola by having ulnar and tarsal folds (absent in both); finally, from H. mashpi by having an inner tarsal fold delineated with a whitish stripe (inner tarsal fold and whitish stripe absent in H. mashpi) and by lacking a dark brown middorsal stripe (present in H. mashpi). Description of the holotype. An adult male (Figs. 2a–b) of 29.9 mm SVL. Body relatively slender (about 2.4 times longer than wide). Head slightly wider than long (HW: 35.6% of SVL; HeL: 32.9% of SVL; HeL/HW: 0.9). Snout rounded in dorsal view and profile (Figs. 3a,c). Eye-naris distance slightly shorter than eye diameter (EN/ ED: 0.9). Canthus rostralis distinct, slightly concave in dorsal view, rounded in cross section; loreal region concave and sloped; lips rounded, not flared; nostrils slightly protruded anterolaterally. Internarial region and top of head flat. Interorbital distance wider than upper eyelid. Eye prominent (ED: 10.2% of SVL). Tympanum distinct, small (TD/ED: 0.4; TD: 4% of SVL); membrane not differentiated but most of circumference of tympanic annulus visible through the skin; about 1/4 of the tympanum concealed dorsally by the supratympanic fold. Supratympanic fold developed, starting at posterior end of upper eyelid and reaching posterior margin of insertion of arm (Fig. 3c). Mental gland present, small (diameter about 11% of SVL), disc-shaped, covering the anterior third of throat (Fig. 3b). Dentigerous processes of vomers conspicuous, straight, narrowly separated from each other; each process bears 5/6 (right/left) teeth. Choanae large, elliptical, not concealed by palatal shelf of maxillary arch. Tongue oval, completely attached to the floor of mouth; vocal slits large, longitudinal, originating on sides of tongue and extending diagonally toward angle of mouth. Vocal sac moderately distensible, evident externally, bilobate, median and subgular. Forearm wider than upper arm; axillary membrane absent. Outer ulnar fold present (Fig. 3c). Fingers short and thick; discs on fingers round, slightly expanded laterally, with clearly defined circumferential groove; disc on FIII slightly wider than tympanum diameter. Relative lengths of fingers: I<II<IV<III. Fingers with fleshy dermal fringes; webbing present among fingers II–IV (Fig. 3d); webbing formula II 2 -–3+ III 2 ½–2+ IV. Subarticular tubercles swollen, round or oval; distal tubercle of FIV larger than all others and weakly bifid. Supernumerary tubercles present, small and rounded. Palmar tubercle small, flat, bifid, poorly defined. Prepollical tubercle elliptical, thick, not modified as a spine. Nuptial excrescences absent. Hind limbs robust; ThL: 46.0% SVL; SL: 50.3% SVL; FL: 40.6% SVL. Calcar tubercle absent. Outer and inner tarsal folds present (Fig. 3e), inner poorly defined. Toes short, with wide lateral fringes; toe discs small, rounded, barely expanded, slightly smaller than those on fingers. Relative length of toes: I<II<III<V<IV; extensive toe webbing, formula: I 2 -–2+ II 1 +–2½ III 1 ½–2½ IV 2 +–1+ V. Outer metatarsal tubercle small, rounded, slightly elevated; inner metatarsal tubercle elliptical, flat, about three times the size of the outer; subarticular tubercles round, moderately elevated; supernumerary tubercles present, small and rounded. Cloacal opening directed posteroventrally at mid-level of thighs; cloacal sheath short; margins of vent with numerous small folds; numerous warts around the cloaca; supracloacal fold present. Skin on dorsum, flanks, throat, chest, forearms, dorsal and inner surfaces of thighs, smooth; areolate on ventral surface of thighs and coarsely areolate on belly. White parietal peritoneum present, covering almost all ventral surface of abdominal cavity. Color in life of the holotype (based on photographs). Dorsal background color pale yellow, densely and finely punctuated with dark brown, and with scattered small reddish brown spots. Flanks paler than dorsum and almost without dark brown punctuation. Ventral surfaces immaculate yellowish cream, except margins of the lower jaw that are punctuated with dark brown. Whitish stripes present on external border of upper eyelids and supratympanic folds, longitudinally on the mid-dorsum, on supracloacal fold, ulnar fold, inner and outer tarsal folds, and on dorsal internal surface of shanks. White warts around the cloaca. Axillae bright yellow; groins bright ocher. Fingers I–II, toes I–III, and interdigital membranes of fingers and toes immaculate bright yellow. Iris gray with thin black reticulation. Color of the holotype in preservative (after eight years, May 2017). Dorsal and ventral background color pale cream (Fig. 2a–b). Head, lower jaw, and all dorsal surfaces (except fingers I–II and toes I–III), with dense and fine pale brown punctuation. Whitish stripes on border of upper eyelids, inner and outer tarsal fold, dorsal internal surface of shanks, and mid-dorsal, all barely visible; whitish and more defined stripes on the supratympanic, ulnar and supracloacal folds. Warts around the cloaca with faint whitish coloration. Measurements of the holotype (in mm). SVL: 29.9; SL: 15.0; ThL: 13.7; FL: 12.1; HaL: 9.4; HeL: 9.8; HW: 10.6; IO: 4.1; InD: 2.6; EN: 2.6; ED: 3.0; TD: 1.2; ETS: 4.5; F3D: 1.5; T4D: 1.4; F1L: 5.5: F2L: 7.0. See Table 2 for measurements of the type series. Morphological and color variation. Hyloscirtus japreria sp. nov. exhibits sexual dimorphism in size (Fig. 2), with adult females larger than males (males: 28.8–32.7 mm [30.3 ± 1.1; n = 17] of SVL vs. females: 35.6–39.1 mm [37.1 ± 1.4; n = 5]). Mental gland, vocal slits and vocal sac are secondary sexual characters present in males and absent in females. Variation of general morphometric characters of the type series is shown in Table 2. The proportions of the head (slightly wider than long to as wide as long) are very similar in the type series and do not differ between sexes (HeL/HW: 0.9–1.0; n = 22), but the relative size of the head is slightly larger in males (HeL: 29.4–33.5% of SVL [32.1 ± 1.0; n = 17]; HW: 31.8–37.0% of SVL [34.6 ± 1.3; n = 17]) than in females (HeL: 28.7–30.4% of SVL [29.7 ± 0.7; n = 5]; HW: 28.8–33.7% of SVL [32.0 ± 2.0; n = 5]). The tympanum is relatively small (3.4–5.0% of SVL [4.2 ± 0.4; n = 22]) and its ratio with respect to the eye has little variation between sexes (TD/ED: 0.3–0.4 [n = 17] in males vs. 0.4–0.5 [n = 5] in females). EN/ED ratio is also variable; however, in all but one specimen of the type series, eye-naris distance is smaller than eye diameter (0.6–1.0 [0.8 ± 0.1; n = 22]). The eye is prominent and relatively bigger in males than females (ED: 10.2–12.0% of SVL [10.8 ± 0.6; n = 17] in males vs. 9.2–10.4% of SVL [9.9 ± 0.5; n = 5] in females). ETS/ED ratio is also slightly variable but snout is always longer than eye diameter (1.1–1.5 [1.3 ± 0.1; n = 22]). Hand length ranges from 24.5–31.6% of SVL (29.0 ± 1.7; n = 22). Thigh length ranges from 43.5–48.9% of SVL (46.4 ± 1.6; n = 22); shank length ranges from 44.8–52.8% of SVL (49.2 ± 1.9; n = 22), and foot length ranges from 35.2–42.8% of SVL (38.8 ± 1.7; n = 22).
Aromobates tokuko Rojas-Runjaic, Infante-Rivero & Barrio-Amorós, 2011, sp. nov.
No full textAromobates tokuko sp. nov. English name: Perija’s Nurse Frog Spanish name: Sapito Niñera de Perijá (Figures 1–3, Table 2) Holotype. Adult male, MHNLS 18479, from the surroundings of Ipika, Yukpa indigenous community, Río Tokuko basin, Municipio Machiques de Perijá, Sierra de Perijá, Estado Zulia, Venezuela (09° 52 ’ N, 72 ° 51 ’ W; elevation 595 m), collected on 26 November 2006, by F. Rojas-Runjaic and E. Infante. Paratypes. Twenty-one specimens: four adult females (MHNL 18471, 18474, 18478, 18480) and two adult males (MHNLS 18473, 18476) with the same collection data as the holotype; four adult females (MHNLS 18490 – 91, 18493, 18495) collected on 27 December 2006, at the holotype locality; two adult males (MHNLS 18498 – 99) from Kusare waterfall, near Ipika, Río Tukuko basin, Municipio Machiques de Perijá, Sierra de Perijá, Estado Zulia, Venezuela (09° 52 ’ N, 72 ° 50 ’ W; elevation 579 m), collected on 28 December 2006 by F. Rojas-Runjaic and E. Infante; three adult females (MHNLS 18521 – 23) and one adult male (MHNLS 18520) collected on 11 April 2007, with the same collection data as the holotype; one adult female (MHNLS 18568) and three adult males (MHNLS 18566 –67, 18569), from Kiriponsa, Yukpa indigenous community, Río Tokuko basin, Municipio Machiques de Perijá, Sierra de Perijá, Estado Zulia, Venezuela (09º 52 ’ N, 72 º 52 ’ W; elevation 1005 m), collected on 12 April 2007 by F. Rojas-Runjaic and E. Infante; one adult female (MHNLS 19748) from near Terakibu, Yukpa indigenous community, Río Tokuko, Municipio Machiques de Perijá, Sierra de Perijá, Estado Zulia, Venezuela (09º 52 ’ N, 72 º 49 ’ W; elevation 419 m), collected on 16 September 2008 by F. Rojas-Runjaic and P. Cabello. Referred specimens. Eight specimens: four juveniles (MHNLS 18477, 18481– 83), with the same collection data as the holotype; three juveniles (MHNLS 18492, 18494, 18496) and one premetamorph (MHNL 18497), collected on 27 December 2006, with the same collection data as the holotype. Diagnosis. (1) Skin of posterior third of dorsum, flanks, prehumeral area, dorsum of thighs and shanks granular; (2) Paired dorsal digital scutes conspicuous and protuberant on all toes; (3) Distal subarticular tubercle under FIV circular and conspicuous; (4) FIV extending past distal subarticular tubercle of FIII; (5) FI shorter than FII; (6) Digital discs present; (7) Finger discs moderately expanded; (8) Lateral fringes absent on FI and FIV, preaxials on FII and FIII, present and conspicuous on all toes; (9) Metacarpal ridge absent or, if present, very weak and almost indistinct; (10) FIII of adult males not swollen; (11) Carpal pad absent; (12) Male excrescences on thumb absent; (13) Thenar tubercle small (approximately 1 / 3 of the palmar tubercle and equal in size and shape to the proximal subarticular tubercle of FII) elliptical and conspicuous; (14) Black arm gland in adult males absent; (15) Tarsal keel present, well-defined and slightly curved from proximal edge of inner metatarsal tubercle, transverse across tarsus, and reaching almost the mid-tarsus where it becomes slightly wider; (16) Toe discs slightly expanded, disc of TI barely wider than distal end of adjacent phalanx (including fringes), discs of TII–TV about 1.5 times wider than distal end of adjacent phalanx; (17) Toe webbing basal between TI–TIV; (18) Metatarsal fold flat and very weak, almost indistinct; (19) dorsum uniformly dark brown or with a pale brown background and irregularly-spaced dark brown spots; (20) Dorsolateral stripe present, pale golden to cream, from tip of snout to midbody or to upper level of the groin, eventually continual with paracloacal horizontal stripes; (21) Oblique lateral stripe present but diffuse, formed by small whitish spots coincident with small non-keratinized tubercles, extending from groin to midbody; (22) Paracloacal horizontal stripe present, diffuse to very well defined; (23) Ventrolateral stripe absent; (24) Throat grayish ivory (with profusion of evenly spaced melanophores when seen under magnification) on almost all males and some females, immaculate ivory on almost all females and some males; (25) Throat collar absent; (26) Chest and belly white to ivory, rarely with pale gray or pale brown spots on chest and anterior third of belly; (27) Ventral surface of arms, thighs, shanks and tarsi pale orange; (28) Palms and soles pale gray; (29) Axillae, groins and hidden parts of hind limbs weakly yellowish; (30) Iris golden and finely dotted with black, with a transversal wide blackish band and a bright golden pupil ring; (31) Large intestine unpigmented, creamish white; (32) Adult testis whitish; (33) Median lingual process absent; (34) Tympanum ill-defined, about 1 / 4 concealed posterodorsally; (35) Vocal sac structure not distinct; (36) Teeth present on the maxillary arch; (37) Small size, with males up to 23.2 mm of SVL and females up to 29.2 mm. Comparison with other species. Aromobates tokuko (characters in parentheses) differs by its small size from large species of the genus, such as A. alboguttatus (Boulenger, 1903), A. leopardalis (Rivero, 1978), A. meridensis (Dole & Durant, 1973), A. nocturnus (Myers, Paolillo & Daly, 1991), and A. capurinensis (Péfaur, 1993), all with maximum SVL> 31 mm (up to 29.2 mm). Aromobates alboguttatus has smooth dorsal skin (granular) and its venter is brown spotted with white (ivory white). Aromobates leopardalis has toes 1 / 3 webbed (basally webbed between TI–TIV), dorsal skin smooth with a few tubercles posteriorly (granular), a strongly spotted dorsum (without consistent pattern). Aromobates meridensis has FI slightly longer than FII (FI shorter than FII), subarticular distal tubercle of FIV indistinct (conspicuous), fringes absent on fingers (preaxial fringes on FII and FIII), tarsal keel straight and indistinct (slightly curved and well defined), dorsolateral stripes absent or, if present, very ill-defined (dorsolateral stripes always present, generally well-defined), and belly bright yellow with diffuse dark reticulation (belly immaculate ivory, occasionally with diffuse pale brown or grayish spots on anterior half, never reticulated). Aromobates nocturnus is the most peculiar and largest congener, with SVL reaching 62 mm (29.2 mm), fully webbed toes (basally webbed), belly coloration mottled with grayish white in life (immaculate ivory, occasionally with diffuse pale brown or grayish spots on anterior third), and unique nocturnal (diurnal), aquatic (riparian), and releasing a mercaptanlike odor when molested (absent). Aromobates capurinensis is a larger frog, mean SVL of females 33.4 mm (25.5 mm), with dorsal skin smooth (granular), dorsolateral stripes present but indistinct (distinct), oblique lateral stripe absent (formed by small whitish spots). The seven other species of Aromobates (A. duranti [Péfaur, 1985], A. haydeeae [Rivero, 1978], A. mayorgai [Rivero, 1980], A. molinarii [La Marca, 1985], A. orostoma [Rivero, 1978], A. saltuensis [Rivero, 1980], and A. serranus [Péfaur, 1985]), are similar in size to Aromobates tokuko. Aromobates duranti has smooth dorsal skin (granular), FI longer than FII (FI shorter than FII), finger discs not expanded (expanded), and a gray venter with white spots (unicolor ivory without spots). Aromobates haydeeae has no fringes on fingers (fringes on FII and FIII), FI longer than FII (FI shorter than FII), tarsal fold ill-defined (well defined), dorsal color reddish copper (dark to light brown), and ventral parts orange (ivory white). Aromobates mayorgai has smooth dorsal skin with a few tubercles posteriorly (granular), ventral parts yellow (ivory white), and the oblique lateral stripe absent (formed by small whitish spots). Aromobates molinarii has FI and II equal in length (FI shorter than FII), fringes absent on fingers (preaxial fringes on FII and FIII), tarsal fold slight (well defined), toes moderately webbed (basally webbed), and the oblique lateral stripe absent (formed by small whitish spots). Aromobates orostoma has no fringes on fingers (fringes on FII and FIII), dorsal skin smooth (granular), and venter yellow (ivory white). Aromobates saltuensis has FI and FII equal in length (FI shorter than FII), dorsal skin smooth with a few tubercles posteriorly (granular), supratympanic bulge absent (present), and no tympanum (present). Aromobates serranus has a distinct tympanum (ill-defined), dorsum brown with blotches or reticulum (usually uniform, but not reticulated), and ventral parts creamy white with brown reticulation (immaculate ivory). Description of the holotype. The holotype is an adult male of 22.6 mm SVL. Body robust, short and slightly wider than head (Fig. 1 a). Head almost as long as wide; head length 38.1 % SVL; greatest head width between angles of jaws 38.7 % SVL. Snout short and subacuminate in profile (Fig. 2 c), nearly rounded in dorsal and ventral view (Figs. 2 a–b). Distance eye-naris slightly shorter than eye diameter (EN/ED = 0.8). Nares are situated laterally to the tip of the snout; nares visible from the front, not visible dorsally; barely visible ventrally. Canthus rostralis faintly angular; loreal region flat. Interorbital region wider than upper eyelid; snout longer than eye length (ETS/ ED = 1.8). Tympanum ill-defined, about 1 / 4 of the tympanum concealed posterodorsally by low supratympanic fold formed by superficial slip of m. depressor mandibulae; tympanum is positioned closely behind eye and low, nearly touching angle of jaws. Teeth are present on maxillary arch. Vocal slits large and long, from mid-level of tongue to near angles of jaws. Tongue is slightly cordiform, 2 / 3 free posteriorly, without median lingual process. Dorsal skin, including dorsal surfaces of hind limbs, granular, moderately granular on upper eyelids, and smooth on forelimbs. Skin of the flanks moderately granular; tuberculate on the prehumeral area (inferoposteriorly to angle of mouth), on ventral portion of the flanks, external sides of shanks and tarsi. Ventral skin smooth on throat, chest and lower portion of belly, moderately granular on low chest, and upper belly; ventral skin smooth on ventral surface of the arms, and slightly granular on undersurface of thighs. Hand (Fig. 2 d) is moderate in size (26.8 % SVL). Relative lengths of adpressed fingers are III> IV> II> I; FIV reaches past the distal subarticular tubercle of FIII; discs of all fingers moderately expanded, round; disc on FIII 1.5 times wider than distal end of adjacent phalanx. FIII not swollen. Base of palm has a large, rounded palmar tubercle; small (approximately 1 / 3 of the palmar tubercle and equal in size and form to the proximal subarticular tubercle of FII), well-defined, and elliptical thenar tubercle on base of FI; one elliptical subarticular tubercle each on FI and FII, and two circular each on FIII and FIV; all tubercles very conspicuous and swollen; without supernumerary tubercles. Fringes on fingers absent on FI and FIV; preaxial fringes present on FII and FIII. Metacarpal ridge and carpal pad absent. Paired dorsal digital scutes conspicuous and protuberant in all fingers. Nuptial excrescences on thumb absent. Hind limbs of moderate length, shank 49.8 % SVL. Relative lengths of adpressed toes are IV> III> V> II> I; TI short, the tip reaching the anterior third of subarticular tubercle of TII. Toe discs slightly expanded, about 1.5 times wider than distal end of adjacent phalanx. Paired dorsal digital scutes conspicuous and protuberant in all toes. Feet (Fig. 2 e) with basal web from TI–TIV, absent between TIV and TV. One to three non-protuberant, small subarticular tubercles present (one on TI and TII, two on TIII and TV, three on TIV, the proximal one almost indistinct). Conspicuous fringes on all toes. Two metatarsal tubercles are present, including a small round outer, and an oval and protuberant inner tarsal tubercle, the latter twice the size of the former. Metatarsal fold flat and very weak, almost indistinct. With a well-defined and slightly curved tarsal keel from proximal edge of inner metatarsal tubercle, transverse across tarsus, and almost to the mid-tarsus, where it becomes slightly wider. Anal opening at upper quarter level of thighs, with a slight flap above vent. Color in life (based on field notes of FR, 11 / 26 / 2006). The dorsal background color is pale brown, with some irregular dark brown spots extending from the interorbital region to the posterior third of the dorsum; two conspicuous paravertebral dark brown stripes present, beginning in the sacral area and joining distally above the vent. A dorsolateral pale golden stripe present on each side of dorsum, starting from tip of snout and continuing through the canthus rostralis, external edge of the upper eyelid and dorsolaterally to reach the upper level of the groin; this stripe is solid and conspicuous, iridescent when seen under magnification. Sides of body with a well-defined blackish brown lateral band from tip of snout to upper level of groin, covering the naris, the two superior thirds of the eye, and the superior third of the tympanum; the upper border of this band is almost straight and in touch with the dorsolateral pale golden stripe, while the lower border becomes broader and diffuse posterior to the tympanum. A diffuse and inconspicuous oblique lateral stripe is present from groin to midbody; it is formed by small whitish spots, coincident with small non-keratinized tubercles. Ventrolateral stripe absent. Tubercles on prehumeral area and ventral portion of the flanks whitish. Paracloacal marks whitish and diffused, forming horizontal stripes from base to middle of thighs. Dorsal background of forelimbs pale brown. Shoulder and axilla slightly yellowish; anterior side of upper arm with a diffuse dark brown longitudinal stripe, posteriorly without marks. Forearm dorsally dotted with small and diffuse brown spots; with a ventrolateral row of irregular blackish brown spots extending from elbow to wrist. Fingers with alternating whitish and blackish brown bands; discs blackish brown with bluish digital scutes. Dorsal surfaces of hind limbs pale brown in background color; cross-barred with four to five well-defined blackish brown bars on thighs, shanks and tarsi; a longitudinal blackish brown stripe on anterior side of thigh, from groin to knee. Groin and hidden parts of hind limbs slightly yellowish. Toes banded as the fingers; discs blackish brown with bluish digital scutes. Ventrally the throat is grayish ivory; the chest and belly ivory (with profusion of evenly spaced melanophores when seen under magnification); ventral surfaces of arms, thighs, shanks and tarsi are pale orange. Palms and soles gray. Iris is golden and finely dotted of black, with a transversal wide blackish band and a bright golden pupil ring. Color in preservative (after three years, January 2010). Dorsal background pale brown faded to pale beige; dorsal dark brown spots and stripes turned pale brown. The golden color of dorsolateral stripe faded to pale cream (Figs. 1 a, 2 b). Blackish brown lateral band, spots of the flanks, inner longitudinal stripe of the upper arm, spots of the forearms and cross-bands of the thighs, shanks, fingers and toes turned dark brown (Figs. 1 a, 2 c–d). The ventral side pale ivory with a profusion of melanophores on the throat conferring a dirty aspect to this region (Fig. 1 b, 2 a). Palms and soles are dirty ivory, fingers and toes spotted with pale gray (Figs. 1 b, 2 d–e). Measurements of holotype (in mm). SVL: 22.6; SL: 11.3; ThL: 11.3; FL: 10.7; HaL: 6.1; HeL: 8.6; HW: 8.8; InD: 2.8; EN: 2.1; ED: 2.6; TD: 1.1; ETS: 4.5; F 3 D: 0.8; T 4 D: 1.0; 1 FiL: 3.3: 2 FiL: 3.9. See Table 2 for measurements of the type series. Variation. Aromobates tokuko is quite variable in color pattern in both sexes and it is difficult to differentiate males and females at first glance. Except by size no other sexually dimorphic characters are noticeable (Figs. 1 a– d). Males and females of similar size could only be identified by dissection or inspection of presence/absence of vocal slits. Adult females reach larger sizes than adult males; the nine adult males of the type series range in SVL from 21.7–23.2 mm (22.6 ± 0.5), while the thirteen adult females range from 21.9–29.2 mm (25.5 ± 1.8). Variation of general morphometric characters of the type series is shown in Table 2. The head length ranges from 35.0%– 39.8 % SVL (37.3 ± 1.3; n = 23), and maximum head width from 35.1 %– 39.1 % SVL (37.0 ± 1.1; n = 23). The ratio of width/length of head is somewhat variable, with the head sligthly longer than wide in 57 % of the type series, slightly wider than long in 30 %, and as wide as long in the remaining 13 %. The ratio EN/ED is also variable but every distance eye-naris is slightly shorter than eye diameter (0.7–0.9 [0.8 ± 1.1; n = 23]). The snout in males is slightly longer than the eye (ratio ETS/ED) and more variable than in females (males: 1.3–1.8 [1.4 ± 0.2; n = 10], vs. females: 1.2–1.5 [1.4 ± 0.1; n = 13]). Hand length ranges from 23.3 %– 29.4 % SVL (26.4 ± 1.3; n = 23). The male paratype MHNLS 18473 has FI slightly longer than FII, while the norm is that FI is shorter than FII. A metacarpal ridge is present but very inconspicuous in MHNLS 18498 (male) and MHNLS 18522 (female), and it is absent in all other specimens of the type series. A metatarsal fold is present but very weak and almost indistinct in MHNLS 18471, 18480, and 18490 (all females). Protuberant tubercles on the posterior third of dorsum, flanks, thighs and shanks are more evident in females (Figs. 1 a, c), very conspicuous in MHNLS 18478, 18490– 91 and 18495. Dorsal color patterns vary from uniform dark brown (males MHNLS 18520, 18498– 99; females MHNLS 18480, 18490–91, 18523) (Fig. 3 b), to pale brown with irregularly-spaced dark brown spots (Fig. 3 a), as in the holotype (Fig. 1 a). Dorsolateral stripes yellowish cream to pale golden, generally well-defined and extending to upper level of the groin (Fig. 3 a), eventually comming into contact or continuing with paracloacal light stripes (males MHNLS 18473, 18498, 18567, 18569; females MHNLS 18478, 18480); or ill-defined and ending at level of mid-dorsum (males MHNLS 18476, 18520, 18566; females MHNLS 18471, 18490–91, 18495) (Figs. 1 c, 3 b). Cross-bands on hind limbs range from four to seven, generally blackish brown and well-defined (Fig. 3 a); diffuse pale brown in MHNLS 18491, 18523 (females), and 18476 (male); hind limbs irregularly spotted with blackish brown (not cross-banded) in MHNLS 18566. Light paracloacal stripes ill-defined or absent in specimens whose dorslateral stripe is also ill-defined (males MHNLS 18476, 18566; females MHNLS 18490 – 91), and very welldefined in specimens with conspicuous dorsolateral stripe (males MHNLS 18498 –99, 18569; females MHNLS 18478, 18480). Oblique lateral stripe almost indistinct in MHNLS 18476 (lateral background uniformly pale brown), and MHNLS 18490 (lateral background uniformly dark brown); well-defined on males MHNLS 18473, 18498– 99, and 18569. Ventrally the throat, chest and belly generally are immaculate ivory in females (Fig. 1 d), and the throat is grayish ivory in males (Fig. 1 b), with a profusion of melanophores when seen under magnification (Fig. 2 a); even grayish ivory on chest in MHNLS 18499. In the males MHNLS 18473 and 18476 the throat is barely pigmented, and immaculate ivory in the male MHNLS 18499. Two females have the throat dirty ivory (MHNLS 18491 and 18495), and one female (MHNLS 18490) have weakly spotted of pale brown the throat, chest and anterior half of belly. Distribution, habitat and natural history. Aromobates tokuko is only known from four localities in the Río Tokuko basin (Kiriponsa, near Ipika, upstream of the Kusare waterfall, and Terakibu) on the eastern versant of the Sierra de Perijá, northwestern Venezuela, between elevations of 419 and 1005 m (Fig. 4). The vegetation of this altitudinal belt was described by Huber & Alarcón (1988) as seasonal semideciduous ombrofilous basimontane forest (from 100–800 m asl), and ombrofilous submontane/montane evergreen forest (from 800–2500 m asl). The region has a biseasonal climatic regime, with a dry period from December–April, and a rainy period from May– November, with maximum rain peaks in May and October, and minimum rain in January and July (Masciangioli & Febres 2000). Aromobates tokuko is a diurnal and cryptic riparian frog. Even though its presence in the streams can be readily detected by its vocalization, it is very hard to detect visually because males call from underneath rocks or logs, or from hollows and shelters, and exhibit a cryptic dorsal color pattern. The eleven specimens (three males, four females and four juveniles) collected near Ipika on November 2006 were found under rocks and rotten logs around a small quiet pool, in an area of about 4 m 2, along a small intermittent stream permanently shaded by dense canopy in the gallery forest. During the collecting effort (1400–1500 h), several males were heard calling from hidden sites along the stream, but we failed to record the vocalizations. Several tadpoles of A. tokuko in different larval stages were found in the pool but no adult specimen carrying tadpoles were observed. Several tadpoles, and two adults of Leptodactylus cf. wagneri were also found in syntopy with the tadpoles of A. tokuko. On December 2006, two males (MHNLS 18498 and 18499) were collected while vocalizing at 1140 h, hidden under rocks along a shaded section of a fast moving stream, upstream from Kusare waterfall (Fig. 5). Both males were about 1.5 m apart while calling. At this locality, A. tokuko is syntopic with Rhaebo haematiticus and Leptodactylus cf. wagneri. Finally, the specim
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