Bioacoustics of Ranitomeya yavaricola

Ranitomeya yavaricola can be distinguished from other species of Ranitomeya by the combination of irregular, pale turquoise spots and stripes on the dorsum, solid-bronze limbs, and irregular sky-blue spots on the ventral surface of the upper thighs. Ranitomeya flavovittata possesses irregular, bright yellow spots and stripes (often broken) on the dorsum (vs. pale turquoise in R. yavaricola ). Additionally, the limbs of R. flavovittata are black with light blue reticulation (vs. limbs solid bronze in R. yavaricola ), and R. flavovittata typically possesses a complete (to nearly complete) yellow median dorsal stripe and lacks conspicuous spots on the ventral surface of the upper thighs (vs. median dorsal stripe absent and ventral thigh spots present in R. yavaricola ). Ranitomeya vanzolinii possesses bright yellow dorsal spotting, light blue reticulation on a ground color of black on limbs, and lacks conspicuous spots on the ventral surface of the upper thighs. Ranitomeya yavaricola is similar in appearance to the nominal morph of Excidobates captivus and Adelphobates castaneoticus . The nominal morph of Excidobates captivus has dull brown limbs (vs. bronze in R. yavaricola ), bears yellow spots on the flanks (vs. flank spots absent in R. yavaricola ), and has paired reddishorange dorsolateral dashes (vs. dorsolateral spots and stripes pale turquoise to spring green in R. yavaricola ). Adelphobates castaneoticus is larger in size (SVL up 23 mm ), lacks the inner metacarpal tubercle, and lacks distinct bronze limbs. Furthermore, it has conspicuous red flash-marks on the upper surface of the forearms, thighs, and calves. Measurements (in mm) of holotype . The male holotype ( Fig. 3 ) has SVL 15.2; FL 7.1; TL 7.2; KK 13.1; FoL 4.9; HaL3.4; HL 5.5; HW 5.0; BW 4.3; UEW 2.3; IOD 2.2; IND 1.8; TD 0.9; ED 1.9; DET 0.5; L1F 1.9; L2F 2.2; W3D 0.9; W3F 0.3. For paratype measurements see Table 1 . Description of holotype . Widest part of head at jaw articulations. Head width slightly wider than body. Tongue ovoid; teeth absent. Snout sloping and rounded in lateral profile, slightly blunted in dorsal profile. Nares situated at tip of snout and directed laterally; both nares visible from ventral and anterior view but not from dorsal view. Canthus rostralis rounded, loreal region flat and nearly vertical. Upper eyelid approximately equal in width to interorbital distance; internarial distance roughly equal to horizontal eye diameter. Tympanum round, partially concealed posterodorsally. In life, skin texture nearly smooth on dorsal surfaces of body and head; limbs and rump weakly granular. Venter weakly granular on limbs and body, ventral surface of head nearly smooth. Hands ( Fig. 4 ) relatively large, length 22 % of SVL. Relative length of appressed fingers III> IV> II> I; first finger 90 % length of second; finger discs greatly expanded, width of disc on finger III 2.6 times width of adjacent phalanx. Unpigmented median metacarpal tubercle present on base of palm; inner metacarpal tubercle present near base of finger I; unpigmented proximal subarticular tubercles present on base of each digit, except on finger I, where tubercle is part-way up the digit; distal subarticular tubercle visible only on finger III. All tubercles raised above level of hands; scutes present on dorsal surface of fingers. MZUNAP -01-518 MZUNAP -01-520 MZUNAP -01-519 1 2 3 4 5 6 7 8 9 10 11 12 Mean (Max-Min) SVL FL TL 17.2 7.0 7.5 15.2 7.1 7.2 16.8 7.0 7.3 15.5 7.2 6.8 16.0 7.7 7.5 16.2 7.4 7.1 16.2 7.5 7.2 15.9 7.0 7.4 17.1 8.0 6.9 16.7 7.6 7.6 16.6 7.5 7.6 17.7 7.9 7.5 16.7 8.0 7.2 15.9 7.2 7.2 16.7 7.0 7.3 16.4 (17.7 15.2) 7.4 (8.0 7.0) 7.3 (7.6 6.8) KK FoL HaL HL HW 14.8 6.1 3.8 6.1 5.3 13.1 4.9 3.4 5.5 5.0 14.0 5.5 3.8 5.9 5.1 13.8 5.7 3.8 5.8 5.4 14.8 6.2 4.0 6.2 5.4 14.2 6.4 4.0 6.6 5.9 13.7 5.9 3.9 6.2 5.4 14.3 5.6 4.1 6.2 5.5 14.2 6.0 4.0 6.4 5.6 14.4 6.4 4.0 5.9 5.4 14.9 5.9 4.2 5.8 5.6 14.7 6.0 4.2 5.9 5.7 14.5 6.6 3.9 6.3 5.7 14.0 6.0 4.0 6.2 5.5 14.1 6.5 4.2 6.5 5.4 14.2 (14.9 13.1) 6.0 (6.6 4.9) 4.0 (4.2 3.4) 6.1 (6.6 5.5) 5.5 (5.9 5.0) BW UEW IOD IND TD ED DET 4.3 2.6 2.3 1.9 0.9 1.9 0.3 4.3 2.3 2.2 1.8 0.9 1.9 0.5 5.4 2.4 2.3 2.1 0.9 2.2 0.4 5.2 2.5 2.2 2.0 0.7 2.0 0.5 5.9 2.7 2.5 2.3 0.5 2.0 0.7 4.9 2.9 2.3 2.2 1.1 2.0 0.6 5.5 2.6 2.2 2.2 0.8 2.0 0.3 5.3 2.8 2.2 2.2 0.7 2.1 0.4 5.7 2.6 2.2 2.3 0.7 1.9 0.6 5.4 2.6 2.5 2.3 0.5 2.1 0.7 5.6 2.4 2.0 2.2 0.7 1.9 0.5 6.1 2.4 2.1 2.4 0.9 2.0 0.6 5.6 2.8 2.3 2.0 0.9 2.1 0.8 5.1 2.7 2.6 2.3 0.9 1.8 0.6 5.8 2.0 2.6 2.2 1.3 2.0 0.3 5.3 (6.1 4.3) 2.6 (2.9 2.0) 2.3 (2.6 2.0) 2.2 (2.4 1.8) 0.8 (1.3 0.5) 2.0 (2.2 1.8) 0.5 (0.8 0.3) L1F L2F W3D W3F 1.8 2.5 0.8 0.4 1.9 2.2 0.9 0.3 2.1 2.2 0.9 0.3 2.0 2.6 1.0 0.4 2.1 2.8 1.0 0.3 2.0 2.8 1.1 0.3 2.0 2.6 1.0 0.4 1.9 2.9 1.0 0.4 2.2 3.0 1.1 0.4 2.0 2.5 1.0 0.4 2.1 2.9 1.0 0.5 2.1 2.9 1.0 0.3 2.0 2.8 1.1 0.4 2.0 2.5 1.1 0.3 2.1 2.8 1.1 0.5 2.0 (2.2 1.8) 2.7 (3.0 2.2) 1.1 (2.0 0.8) 0.4 (0.5 0.3) BM SEX - 3 - Ƥ - 3 0.3 * 0.3 3* 0.4 3* 0.3 3* 0.3 3* 0.4 3* 0.4 3* 0.5 3* 0.5 Ƥ ** 0.4 3* 0.3 _ 0.4 _ 0.4 (0.5 0.3) _ Hind limbs moderate length, with heel of appressed hind limbs reaching level of eye. Femur and tibia roughly equal in length, femur 99 % length of tibia; knee–knee distance 86% of SVL. Relative lengths of appressed toes IV> III> V> II> I ( Fig. 4 ); first toe short with unexpanded disc; second toe with slightly expanded disc, discs on toes III–V moderately expanded. Two unpigmented metatarsal tubercles on base of foot, one situated medially near base of toe I, the other situated laterally at the base of the fifth metatarsal. Proximal subarticular tubercles present at base of each toe but most notable on toes I, II, III due to lack of pigmentation. Toes III and V with two large subarticular tubercles, toe IV with three subarticular tubercles. Tarsal keel extends from below knee to medial metatarsal tubercle at foot. Tarsal tubercle absent; feet and hands lack webbing and lateral fringing. Color in life. In life, body black with metallic sage-green dashes anterior and posterior to each eye, a single spot between eyes; metallic-sage dorsolateral stripes extend from upper thighs to shoulders where they each form a single dash. Broad sage oblique-lateral lines extend from the axillae to groin, where they fuse with a large spot on upper surfaces of each thigh. Broad labial stripe present, continues posteriorly to upper surfaces of arms. Limbs and digits solid bronze. Paired sage dashes on underside of thighs. Underside of head sage with two pairs of gular spots, creating appearance of an hourglass. Venter black with coarse, irregular sage marbling. Iris black. Color in preservative. In preservative, turquoise/sky-blue coloration turns grey and bronze limb coloration turns brown. Variation (based on 15 adults ). Adults 15.2–17.7 mm SVL (mean 16.4 mm ). Head about as wide as body except in single gravid female whose body was wider than the head. Head width 98 % of body width (range 81–109 %). Head width 30–36 % of SVL in adults. No apparent sexual dimorphism in external morphology except that males possess faint vocal slits on the floor of the mouth and have a slightly expanded subgular pouch. The completeness of dorsolateral and medial lateral stripes varies considerably between individuals ( Fig. 5 ). Line/dash/spot coloration varies subtly between a sky-blue to a very light bluish-yellow to pale sage-green, with a majority of individuals being light turquoise. Venter coloration is typically darker than dorsal coloration and is less variable, being predominantly skyblue. Ventral ground coloration varies from dark brown to black. Venter reticulation is largely symmetrical and often forms a broad, irregular, black medial-stripe in half of the specimens ( Fig. 5 ). Most individuals possess a large gular spot in the center of the throat (80 %). In all individuals, sky-blue dashes are present on the ventral surface of the upper thighs, though their size varies considerably between individuals. Hands relatively large, length 22–26 % of SVL. First finger 65–95 % length of second; finger discs moderately expanded in both males and females, width of disc on finger III 2 –3.6 times width of adjacent phalanx. Tibia 86–107 % length of femur (mean 99 %); knee–knee distance 82–92 % of SVL (mean 87 %). Tadpole measurements (in mm). A stage 25 tadpole (MZUNAP-01-521) was used for the description ( Fig. 6 ). Total length 12.5; body length 4.7; internarial distance 0.9; eye to nares distance 0.8; eye diameter 0.5; interorbital distance 0.7; tail length 7.8. Tadpole description. Snout rounded when viewed from above; body ovoid in dorsal view. Eyes dorsal, angled laterally, pupils white in preservative. Nares not forming tube, situated half-way between eye and tip of snout, directed dorsolaterally. Spiracle sinistral; vent dextral. Ventral tail fin begins at tail base, dorsal tail fin begins just posterior to plane of vent opening, ventral and dorsal fins relatively uniform in thickness throughout tail, tapering towards tip. Musculature depth uniform throughout, tapering toward tip. Mouth directed anteroventrally. Oral disc emarginate, anterior and posterior labia forming flaps free from body wall, 1.4 mm in width. Marginal papillae absent on anterior labium, present in one complete row on posterior labium. Papillae rounded; submarginal papillae absent. Jaw sheaths deep in longitudinal width, serrate, lacking indentations. Lateral processes short, extending barely past lower jaw. Labial tooth row formula is 2(2)/3[1]. A-1 complete (62 teeth), A-2 with medial gap (34 teeth), same width as A-1. P-1 with medial gap (46 teeth), P-2 (44 teeth), and P-3 complete (44 teeth); P-1 and P-2 equal width, P-3 slightly shorter. Color in Life. In life, the head appears light grey. Pigmentation on dorsum mottled brown, ground color weakly transparent. Eyes black, papillae white. Ventral coloration is transparent (most internal organs are visible) with irregular faint red flecking that is dense around mouth and nares. Tail musculature white with abundant brown mottling, fins almost transparent. Color in Preservative. In preservative, coloration is identical, though red pigmentation turns to brown. Vocalizations . The following values are presented as: min-max (average ± SD, number of individuals). The advertisement call is a short trilled note ( Fig. 7 ) with duration between 630–880 ms (760 ± 140 ms, 7) and is repeated at irregular intervals of 2–7 notes per minute (3.14 ± 1.80 notes per minute, 6). Each note consists of 20–27 pulses (mean = 24). Calling activity is sporadic and continues throughout daylight, but peaks in the early morning and late afternoon. Dominant frequency is 5400–6000 Hz (5600 ± 2000 Hz, 5) at temperatures between 24.5–26 °C. The call of R. yavaricola sounds similar to the calls of other species in the vanzolinii group, although there are some slight differences within this group. Ranitomeya flavovittata has slightly longer notes (1 sec vs. 760 ms in R. yavaricola ). Ranitomeya imitator ( n = 25) has a call that is slightly shorter in duration (686 ms vs. 760 ms in R. yavaricola ) and has a slightly lower dominant frequency (5200 Hz vs 5600 Hz in R. yavaricola ).