Bioacoustics of Anomaloglossus verbeeksnyderorum

Description of holotype . Dorsal and ventral skin smooth in preservative; without tubercles on the posterior part of the dorsum. Dorsal skin forming a well-defined rounded, posteriorly projecting flap above vent, opened at the upper level of thighs; no anal tubercles. Head longer than wide, HW 33.6 % of SVL. Snout subacuminate in profile ( Fig 1 A), truncate in dorsal and ventral views ( Figs 1 B and 1C respectively). Nares situated near the tip of the snout and directed slightly anterolaterally, visible from the front, and barely visible from the dorsum and below. Canthus rostralis rounded; loreal region slightly concave. Interorbital region width slightly wider than upper eyelid width. Snout slightly longer than eye diameter; EN 64% of ED. Tympanum large, 40% of ED, distinct only its inferior half; its posterodorsal half hidden beneath the superficial slip of the M. depressor mandibulae ; positioned close behind eye and low, close to the angle of jaws. Maxillary teeth present. Tongue longer than wide, triangular, notched posteriorly, posterior half free; vocal slits present, large. MLP very small, round, as long as wide, positioned on the anterior third of the tongue. Hand length small, 27.4 % of SVL. Relative lengths of adpressed fingers III>IV>I>II ( Fig 2 A); F I slightly longer than F II. Discs of all fingers barely expanded; F III disc 1.2 times width of distal end of adjacent phalanx. Base of palm with large, nearly round, median palmar tubercle; oval inner thenar tubercle on base of F I; one subarticular tubercle on F I and F II, and two on F III and F IV, distal tubercles small; all tubercles low, round. Fringes only appreciable on F II, thick and low. Hind limbs of moderate length; shank 46.6 % of SVL. Relative lengths of adpressed toes IV>III>V>II>I ( Fig 2 C); TI reaching, when adpressed, half subarticular tubercle of TII. Toe discs moderately expanded; T IV disc 1.6 times width of distal end of adjacent phalanx. Feet moderately webbed, the web distally continuous with a thick fringe on all toes, including external edges of T I and T V. Toes moderately webbed; webbing formula I 1 ½-2½ II 1 2/3-3 III 2 ½-3 2/ 3 IV 4-2½ V. One to three non protuberant subarticular tubercles on toes as follows: one on T I and T II, two on T III and T V, and three on T IV. Sole with two distinct metatarsal tubercles, a round outer metatarsal tubercle, and a 1/3 longer elliptical inner metatarsal tubercle. A thick tarsal keel, straight and weakly curved distally, ending at mid length of tarsus, proximally continuous with the narrow fringe on free (preaxial) edge of T I. Metatarsal fold present, low. Color in preservative. dorsally dark brown with two conspicuous parotoid pale brown elongate marks (comma shape), and two symmetrical scapular pale brown round marks ( Fig 1 E). Rostral area pale brown. Both flanks are dark brown, with long white oblique lateral stripes, reaching the posterior edge of the arm. Upper lips dark brown with a few irregular small whitish spots; tympanic area pale brown. Arms and limbs are pale brown, bar crossed by dark brown. Two symmetrical white paracloacal marks surrounded by black. Ventral surfaces are white at simple view; under magnification, melanophores are concentrated on the throat, especially on the chin and along the edges of the lower mandible; melanophores are more scarce on the chest, and missing on the belly ( Fig 1 F). Palms and soles are dark brown. Color in life . The only available photo of a living male is that of an uncollected specimen. In all individuals seen was evident, however, the characteristic color described herein. Anomaloglossus verbeeksnyderorum males ( Fig 3 A) differ from all known species in the genus by showing some bright marks, mostly on the parotoid area (orange), upper arm (orange), post tympanic area (white and orange); paracloacal area (orange), inguinal region and oblique lateral stripe (yellow), and some bluish small spots on the flanks, upper lip, and fingers. Also the upper surfaces of hind limbs are dull orange, barred with dark brown. The rest of dorsum is dark brown with darker brown canthal, supratympanic and interorbital stripes; ventrally males are bright yellow. Females ( Fig 3 B) are more according to dull colored Anomaloglossus , as they are dorsally light brown to greyish, with dark brown marks; the parotoid and paracloacal areas are dull orange, and the axilar region can be yellowish. The fingers have bright bluish white rings beside the finger discs; ventrally they are white. Measurements of holotype (in mm). SVL: 19.3; SL: 9.0; FL: 8.8; HeL: 8.0; HW: 6.5; EN: 1.6; ED: 2.5; TD: 1.0; F3D: 0.6; T4D: 0.6 1FiL: 3.0; 2FiL: 2.7. See Table 1 for measurements of the type series. Character Males (n= 4) Females (n= 9) SVL 18.8±0.7 (17.8–19.5) 19.3±1.8 (16.9–22.3) SL FL 8.5±1.1 (7–9.5) 8±0.8 (7.2–8.9) 9.1±0.6 (8–9.8) 8.4±0.8 (6.8–9.5) HeL 8.1±0.5 (7.6–8.8) 8.3±1.0 (7–9.6) HW ED 6.7±0.5 (6–7.3) 2.2±0.1 (2.1–2.4) 7.0±0.5 (6.2–7.6) 2.3±0.3 (1.9–2.7) TD 1±0.1 (0.9–1.1) 1.1±0.2 (0.8–1.4) F3D T4D 0.5±0.1 (0.4–0.6) 0.6±0.1 (0.5–0.8) 0.6±0.1 (0.5–0.8) 0.7±0.1 (0.5–0.8) 1FiL 3±0.2 (2.8–3.3) 3±0.3 (2.5–3.4) 2FiL 2.7±0.3 (2.4–3) 2.8±0.3 (2.5–3.4) Variation . The only difference between males and females in preservative is the throat coloration, immaculate white on females, with few melanophores on the chin edge ( Fig 1 D), and dirty white to greyish on males, with a profusion of melanophores observed with magnification ( Fig 1 C). Size is not very significant (mean SVL of males 18.8 ± 0.7 mm ; mean SVL of females 19.3 ± 1.8 mm ). The shape of the snout is also variable; truncate ( Fig 1 C) to round ( Fig 1 D). The shape of the fingers can be slender ( Fig 2 A) or robust ( Fig 2 B). Although in the type series there is not an individual that presents warts or flat tubercles on the posterior part of the body or on the thighs, as is usual for Anomaloglossus , a few flat warts can be appreciated in Fig 3 A. The paracloacal marks are most patterned and bright on males ( Fig 3 A) than on females ( Fig 3 B). Little variation is observed in dorsal coloration of females. The back can be light to dark brown, with irregular darker marks, barely distinct on dark specimens (MHNLS 19646, MHNLS 19648, CVULA 7139, for instance), and conspicuous on lighter specimens (CVULA 7137-38). The marks are usually a dark interorbital bar pointing backwards, two symmetric dark and round spots on the occipital area, and an inverted W behind the head (CVULA 7140). All specimens have the hind limbs bar-crossed, being better marked on males. On the left foot of MHNLS 19644 there is a median metatarsal tubercle, half size of inner metatarsal tubercle; the outer metatarsal tubercle is much bigger than on right side; and there is another smaller median tubercle just below the median one. The left foot lacks one toe naturally, apparently T IV) (see remarks; Fig 2 D). Vocalization . A single call was recorded at 24.5 ºC the morning of the 19th of June, 2007. The complete train call has 152 pulsed notes ( Fig 4 A, 4B), lasting 17.14 sec. Nine notes are produced per second ( Fig 4 C, 4D). The dominant frequency is at 4405 Hz while the fundamental is at 3921 Hz. 25 consecutive notes were chosen at random on the spectrogram to determine note duration and inter-note interval. Note duration and inter-note interval were measured, respectively, 0.067±0.003; 0.064–0.072 sec, and 0.044±0.003; 0.0042– 0.0048 sec. (mean±SD and range, respectively). Call comparisons . Only five Anomaloglossus advertisement calls are known. Grant et al. (1997) describe the call of Anomaloglossus atopoglossus ; Myers & Donnelly (1997) that of A. tamacuarensis ; Myers & Donnelly (2008) A. tepuyensis , and Kok et al. (2006a , 2006b ) A. kaiei and A. beebei . For comparison we note (in parentheses) the pertinent call characters of A. verbeeksnyderorum . Anomaloglossus atopoglossus produces calls with 12–14 notes (152), with note call duration of 0.7–0.9 sec (0.06 sec), and a frequency from 4160 to 4240 Hz (3921-4405 Hz). Anomaloglossus tamacuarensis call consists of a long train of double notes (single notes) given at three per second, resulting in six notes per second (9), with a frequency of 3840 Hz (3921–4405 Hz). The call of A. tepuyensis is a short trill of 14–22 notes (152), with 4–7 notes per second (9), with frequency varying from 3270 to 3580 Hz (3921–4405 Hz). Anomaloglossus kaiei does not have a trill, but instead a repetitive and unmodulated call with a note duration of 0.03 and inter-note duration of 0.1 sec (0.06; 0.04 sec) and a dominant frequency of 4850 Hz (3921–4405 Hz). Finally, the described call of A. beebei is a series of whistled trills made from 2 to 5 modulated notes (continuous trill), with a frequency of 4640 Hz (3921–4405 Hz), and 7 (10) calls per second.

Distribution . Known from its type locality and a nearby site (Cerro el Tigre, near Tobogán de la Selva, together a single spot in the map, Fig 5 ), where the species was heard calling in sympatry with Dendrobates leucomelas . There is no reason to believe that the new species does not occur throughout lowland rainforest on granitic substrate at least through the north-western Amazonas and Bolívar states in Venezuela . The area has been little explored, existing a continuum of habitat both to the NE and S. Natural history . Anomaloglossus verbeeksnyderorum has only been seen active on rainy days. Of the many times that the stream at Tobogán de la Selva was visited (from 1995 to 2007 , at least once a year), only in two instances was the species apparent, in both cases during and after rains. The type series was found active on the ground of an immense sandstone slab under rainforest on 22 April 2005 ( Fig 6 ). Around midday, after a rain in the morning, males were calling, and most individuals were seen along the slab, hiding rapidly under boulders when we approached. The animals were also seen and heard on 19 and 20 of June 2007 inside the forest although never more than 20 m away from the stream. Other herpetofauna observed along the creek inside the forest were (during the day) Dendrobates leucomelas, Synapturanus salseri, Gymnophtalmus sp., Plica pansticta, Corallus hortulanus, Bothrops atrox ; and (at night) Pristimantis vilarsi, Leptodactylus lithonaetes, Leptodeira annulata, Oxyrhopus melanogenys, Sibon nebulata and Paleosuchus palpebrosus .