Bioacoustics of Diasporus darienensis

FIGS 10 , 14A, B ELEUTHERODACTYLUS DIASTEMA – MYERS 1969: FIG. 19B Holotype SMF 97304 (original field number AB 1144 ), an adult female ( Fig. 10 ), collected by Abel Batista & Konrad Mebert on the ridge of Pirre mountain range , ~ 3 km north from the peak of Cerro Pirre , Parque Nacional Darien ( PND ), Distrito de Pinogana , Darien , Panama , on 8 December 2012 at 20:35 h ( 7.97312 N , 77.70785 W ; 1143 m a.s.l. ). Paratypes MHCH 2840–41 , 2844–47 , 2862 , SMF 97303, 97314 , 97306–08 , 97661–62 , with same collecting data as holotype ; MHCH 2850 – 52 , SMF 97309 – 10 , 97312 , collected by Abel Batista & Milan Vesely in the Jingurudo mountain range , on a ridge between Aldo Creek and Sambu River , between the Comarca Embera- Wounaan and the PND, Distrito de Sambu , Darien , Panama , 26 – 30 September 2011 ( 7.69271 N , 78.04200 W ; 869 m a.s.l. ) ; SMF 97313, collected by Abel Batista & Gustavo Dojirama at the top of Cerro Sapo , PND, Distrito de Garachine , Darien , Panama , on 4 December 2011 , at 22:06 h ( 7.97618 N , 78.36263 W ; 1169 m a.s.l. ) . Diagnosis Diasporus darienensis sp. nov. is characterized by the following combination of traits (see Table 1 ): (1) dorsal skin texture smooth and/or with rounded or pointed scattered tubercles; (2) tympanic annulus concealed by skin, tympanic membrane absent; (3) snout acuminated in dorsal view and rounded in profile; (4) usually with a slightly enlarged and conical supraocular tubercle, cranial crests absent; (5) dentigerous processes of vomers triangular, diagonal to the eyes, near to the middle of the mouth and posteriorly separated about three-quarters of their total length from each other; (6) vocal sac moderately developed, with longitudinal gular folds evident, vocal slits present on the posterior part of the jaw, halfway under the tongue and ending at the midlevel of the eyes, no nuptial pads; (7) finger II longer than finger I, ungual flap mostly expanded, rounded, and more evident on fingers II – IV; (8) no fringes or webbing on fingers; (9) palmar tubercle ovoid, flattened, and slightly larger than thenar tubercle; thenar tubercle low and elongate; subarticular tubercles rounded and globular; no supernumerary tubercles, palmar accessory tubercles small, rounded, and almost indistinguishable; (10) heel smooth; (11) no fringes or webbing on toes, ungual flap slightly expanded to rounded, more evident on toes II – V; (12) plantar tubercle indistinguishable, between one and three non-protuberant subarticular tubercles present (one on toes I and II, two on toes III and V, and three on toe IV); inner metatarsal tubercle elongated; outer metatarsal tubercles conical and smaller than inner; tarsal ridge absent; (13) dorsal ground color in life brown to reddish, some specimens with pale reticulations on a dark background color; usually with a pair of red or pale dorsolateral lines, venter translucent or suffused with dark color, vocal sac yellow; (14) SVL 18.1 ƚ 21.3 (14.9—22.9, N = 21), males 17.1 ƚ 1.11 (14.9—18.5, N = 15), females 20.7 ƚ 1.86 (18.5—22.9, N = 6); (15) advertisement call composed of a single, amplitude-modulated short note (49.1 – 51.7 ms) with harmonic structure. The dominant frequency is also the fundamental frequency, with most energy emitted at 3.34 – 3.81 kHz. Description of the holotype An adult female (SVL 17.40, Fig. 10 ) with slender body; dorsal skin smooth with small scattered tubercles, ventral skin smooth, discoidal fold not evident, low anal warts present; one small conical supraocular tubercle; eye twice as long as snout; tympanum of moderate size, ratio TD/EL 34%; tympanum indistinguishable, annulus tympanicus concealed by skin, tympanic membrane absent, positioned above the junction of jaws and behind the orbit; head as long as wide (HL/HW 1.03), greatest head width between angles of jaw 40% of SVL; snout subacuminate from above and rounded in profile; nares situated near tip of snout and slightly dorsolaterally directed, clearly visible in frontal view, also visible dorsally but not ventrally; canthus rostralis rounded; loreal region feebly concave; dentigerous processes of vomer clearly visible, orbit in an oblique outline in frontal of eyes, each with five teeth; vocal slits absent; tongue long (25% of SVL) and knobbed at the end, first third attached to floor of mouth; hands moderate in size, 20% of SVL; relative lengths of adpressed fingers I <II <IV <III; finger II subequal in size to finger VI, finger II reaching the disc on finger IV when adpressed; finger III disc 1.6 times wider than distal end of adjacent phalanx; palmar tubercle low and rounded, larger than thenar tubercle; thenar tubercle low and elongate; subarticular tubercles rounded and globular; no supernumerary tubercles; Mean ƚ SD (range); see Material and methods for abbreviations. Numbers in parenthesis next to the species names represents the number of specimens analysed. Species ( N ) SVL HW HL D. anthrax (2)* 16.5 – 18.7 – – D. tigrillo (2) * 16.8 – 17.5 D. citrinobapheus (7) 19 ƚ 1.41 (17.3 – 21.8) 7 ƚ 0.47 (6.5 – 7.8) 7.6 ƚ 0.54 (7 – 8.7) D. darienensis sp. nov. (21) 18.1 ƚ 2.13 (14.9 – 22.9) 6.5 ƚ 0.82 (5.2 – 8.4) 6.5 ƚ 0.64 (5.6 – 7.9) D. aff. diastema , CP (49) 19.2 ƚ 2.78 (14.6 – 27.7) 7 ƚ 1.2 (5.4 – 10.9) 7.6 ƚ 0.99 (6 – 10.6) D. aff. diastema , MM (5) 19.8 ƚ 3.11 (16 – 24.5) 7.3 ƚ 0.93 (6.1 – 8.7) 7.3 ƚ 1.04 (6 – 8.9) D. aff. diastema , EPL (20) 21.3 ƚ 1.82 (18.3 – 25.2) 7.9 ƚ 0.83 (6.1 – 9.6) 7.4 ƚ 0.63 (6.7 – 8.4) D. gularis (3) 22.1 ƚ 1.75 (20.4 – 23.9) – – D. hylaeformis (28) 20.3 ƚ 1.51 (16.9 – 23.1) 7.4 ƚ 0.74 (5.9 – 8.8) 8 ƚ 0.71 (6.8 – 9.5) D. igneus (4) 26.1 ƚ 0.5 (25.5 – 26.6) 9.9 ƚ 0.17 (9.7 – 10.1) 8.5 ƚ 0.31 (8.2 – 8.8) D. majeensis sp. nov. (15) 21.5 ƚ 2.64 (15.3 – 25.5) 8.1 ƚ 0.92 (6.2 – 9.7) 7.8 ƚ 0.84 (6 – 9.4) D. pequeno sp. nov. (13) 19.3 ƚ 2.38 (16.9 – 24.8) 7.2 ƚ 0.96 (6.2 – 8.9) 7.2 ƚ 1.04 (5.9 – 9.1) D. aff. quidditus (51) 14.5 ƚ 1.44 (11.5 – 17.9) 5.4 ƚ 0.48 (4 – 6.5) 5.6 ƚ 0.48 (4.5 – 6.3) D. sapo sp. nov. (11) 22.6 ƚ 2.86 (18.8 – 29.1) 8 ƚ 0.82 (7.1 – 9.7) 8.2 ƚ 1.05 (6.6 – 10.2) D. tinker (39) 17.3 ƚ 1.55 (14.6 – 20.4) 6.4 ƚ 0.42 (5.6 – 7.3) 6.2 ƚ 0.58 (5.3 – 7.2) D. ventrimaculatus (4) 22.8 ƚ 1.89 (20.2 – 24.7) – – D. vocator (12) 14.4 ƚ 1.46 (12 – 17.2) 4.9 ƚ 0.41 (4 – 5.4) 5.4 ƚ 0.66 (4.4 – 6.3) Species TL HAL HW/SVL D. anthrax – – – D. tigrillo – 0.36 (0.34 – 0.37) D. citrinobapheus 7.9 ƚ 0.68 (7 – 9.2) – – D. darienensis sp. nov. 7.7 ƚ 0.73 (6.5 – 9.5) 6.8 ƚ 0.89 (5.4 – 9) 4.4 ƚ 0.39 (3.7 – 5.1) D. aff. diastema , CP 8 ƚ 1.32 (5.5 – 12) 4.5 ƚ 0.93 (3.2 – 6) 0.4 ƚ 0.02 (0.3 – 0.4) D. aff. diastema , MM 8.4 ƚ 0.96 (7.3 – 9.9) 4 ƚ 0.58 (3.5 – 5) 0.4 ƚ 0.01 (0.4 – 0.4) D. aff. diastema , EPL 8.3 ƚ 0.77 (7.3 – 9.4) 3.9 ƚ 0.5 (3.4 – 4.8) 0.4 ƚ 0.02 (0.3 – 0.4) D. gularis – – – D. hylaeformis 8 ƚ 0.75 (6.6 – 10) – – D. igneus 11.7 ƚ 0.5 (11.3 – 12.4) 11.5 ƚ 0.27 (11.1 – 11.7) 11.9 ƚ 0.36 (11.5 – 12.2) D. majeensis sp. nov. 9.1 ƚ 1.14 (6.8 – 10.9) 8.3 ƚ 1.01 (5.9 – 9.6) 5.1 ƚ 0.63 (3.5 – 6.1) D. peque ~ no 8.4 ƚ 0.71 (7.5 – 10) 7.1 ƚ 0.69 (6.1 – 8.5) 4.4 ƚ 0.51 (3.7 – 5.5) D. aff. quidditus 6.9 ƚ 0.56 (5.9 – 8) 5.5 ƚ 0.54 (4.3 – 6.3) 3.3 ƚ 0.32 (2.7 – 4) D. sapo sp. nov. 10.4 ƚ 1.02 (8.3 – 12.1) 9.5 ƚ 1.07 (7.3 – 11.5) 5.5 ƚ 0.6 (4.2 – 6.4) D. tinker 7.3 ƚ 0.66 (5.3 – 8.8) 6.2 ƚ 0.42 (5.3 – 7.4) 3.8 ƚ 0.31 (3.2 – 4.6) D. ventrimaculatus – – – D. vocator 5.8 ƚ 0.43 (5.2 – 6.6) 4.9 ƚ 0.88 (3.9 – 5.5) 2.9 ƚ 0.38 (2.5 – 3.2) Species HW/HL HL/SVL TL/SVL D. anthrax – – – D. aff. tigrillo 0.92 (0.85 – 0.99) 0.39 (0.38 – 0.40) 0.48 (0.46 – 0.50) D. citrinobapheus 0.9 ƚ 0.04 (0.9 – 1) 0.4 ƚ 0.02 (0.4 – 0.4) 0.4 ƚ 0.01 (0.4 – 0.4) D. darienensis sp. nov. 1 ƚ 0.06 (0.9 – 1.1) 0.4 ƚ 0.02 (0.3 – 0.4) 0.4 ƚ 0.02 (0.4 – 0.5) D. aff. diastema , CP 0.9 ƚ 0.07 (0.8 – 1) 0.4 ƚ 0.02 (0.3 – 0.4) 0.4 ƚ 0.04 (0.4 – 0.6) D. aff. diastema , MM 1 ƚ 0.02 (1 – 1) 0.4 ƚ 0.01 (0.4 – 0.4) 0.4 ƚ 0.03 (0.4 – 0.5) D. aff. diastema , EPL 1 ƚ 0.05 (0.9 – 1.1) 0.4 ƚ 0.01 (0.3 – 0.4) 0.4 ƚ 0.02 (0.4 – 0.4) D. gularis – – – D. hylaeformis 0.9 ƚ 0.05 (0.8 – 1) 0.4 ƚ 0.02 (0.3 – 0.4) 0.4 ƚ 0.03 (0.3 – 0.5) D. igneus 1.2 ƚ 0.03 (1.1 – 1.2) 0.3 ƚ 0.01 (0.3 – 0.3) – D. majeensis sp. nov. 1 ƚ 0.03 (1 – 1.1) 0.4 ƚ 0.02 (0.3 – 0.4) 0.4 ƚ 0.02 (0.4 – 0.5) D. peque ~ no 1 ƚ 0.05 (1 – 1.1) 0.4 ƚ 0.02 (0.3 – 0.4) 0.4 ƚ 0.02 (0.4 – 0.5) D. aff. quidditus 0.9 ƚ 0.07 (0.8 – 1.2) 0.2 ƚ 0.19 (0 – 0.4) 0.5 ƚ 0.03 (0.4 – 0.5) D. sapo sp. nov. 1 ƚ 0.05 (0.9 – 1.1) 0.4 ƚ 0.02 (0.3 – 0.4) 0.5 ƚ 0.02 (0.4 – 0.5) D. tinker 1 ƚ 0.06 (0.9 – 1.2) 0.4 ƚ 0.02 (0.3 – 0.4) 0.4 ƚ 0.03 (0.3 – 0.5) D. ventrimaculatus – – – D. vocator 0.9 ƚ 0.07 (0.8 – 1.1) 0.4 ƚ 0.03 (0.3 – 0.4) 0.4 ƚ 0.04 (0.4 – 0.5) *Measurements taken from original descriptions and literature. Traits Species DF (kHz) Low freq. (Hz) High freq. (Hz) Delta freq. (Hz) Note duration (s) D. anthrax * 3.81 3.19 ƚ 0.35 (2.94 – 3.44) 4.45 ƚ 0.29 (4.25 – 4.65) 1.31 – 1.22 0.06 D. citrinobapheus (2) 2.86 – 3.04 2.77 – 2.95 2.95 – 3.42 0.2 – 0.5 0.11 – 0.17 D. darienensis 3.57 ƚ 0.33 (3.34 – 3.81) 3 ƚ 0.29 (2.79 – 3.21) 4.07 ƚ 0.33 (3.83 – 4.3) 1.07 ƚ 0.04 0.05 ƚ 0 (0.05 – 0.05) sp. nov. (2) (1.04 – 1.09) D. aff. diastema , EPL 3.3 ƚ 0.12 (3.2 – 3.5) 2.9 ƚ 0.07 (2.8 – 3) 3.8 ƚ 0.11 (3.6 – 3.9) 0.9 ƚ 0.05 (0.8 – 0.9) 0.1 ƚ 0.01 (0.1 – 0.1) (7) D. aff. diastema , MM 3.1 ƚ 0.2 (3 – 3.4) 2.7 ƚ 0.1 (2.7 – 2.9) 3.5 ƚ 0.15 (3.4 – 3.7) 0.8 ƚ 0.06 (0.7 – 0.9) 0.1 ƚ 0 (0.1 – 0.1) (4) D. aff. diastema , CP (7) 3.3 ƚ 0.16 (3.2 – 3.5) 2.97 3.82 0.86 0.09 D. igneus * (1) 2.4 2 2.7 0.7 0.05 – 0.10 D. majeensis sp. nov. 2.47 – 2.71 2.38 – 3.03 2.85 – 3.14 0.50 – 0.93 0.01 – 0.02 (1) D. peque ~ no (1) 3.44 – 3.48 3.20 – 3.23 3.67 – 3.63 0.39 – 0.46 0.09 – 0.15 D. aff. quidditus (22) 4.81 ƚ 0.14 (4.55 – 5.08) 4.56 ƚ 0.18 (4.35 – 4.84) 4.97 ƚ 0.19 (4.77 – 5.29) 0.41 ƚ 0.02 (0.39 – 0.45) 0.34 ƚ 0.04 (0.25 – 0.38) D. tinker (9) 3.5 ƚ 0.19 (3.14 – 3.71) 3.16 ƚ 0.16 (2.84 – 3.32) 3.8 ƚ 0.2 (3.42 – 4.07) 0.64 ƚ 0.06 (0.56 – 0.75) 0.17 ƚ 0.02 (0.14 – 0.19) D. ventrimaculatus * 2.50 – 2.61 2.14 2.9 0.76 0.07 D. vocator (5) 4.6 ƚ 0.3 (4.35 – 5.1) 3.83 ƚ 0.17 (3.71 – 3.94) 4.94 ƚ 0.25 (4.77 – 5.12) 1.12 ƚ 0.08 (1.06 – 1.18) 0.02 ƚ 0 (0.01 – 0.02) Species Note interval (s) Call rate (calls/min) Notes/bouts Bout duration Interbout duration D. anthrax 0.55 – 5.77 23.4 – 44.2 D. citrinobapheus 16.58 ƚ 0.47 (16.25 – 16.91) 3.61 ƚ 0.1 (3.54 – 3.68) 19 – 30 19.30 – 55.50 38.47 – 156.43 D. darienensis sp. nov. 3.08 ƚ 0.9 (1.65 – 4.84) 20.29 ƚ 6.17 (12.04 – 34.41) – – – D. aff. diastema , EPL 3.1 ƚ 0.68 (2.2 – 3.9) 19.7 ƚ 4.43 (14.8 – 25.8) 8.5 28.4 40.26 D. aff. diastema , MM 3.4 ƚ 1.08 (2.5 – 4.8) 17.9 ƚ 5.02 (12 – 22.8) D. aff. diastema , CP 1.65 34.41 10.2 17.33 30.25 D. igneus 6.40 – 9.67 8.18 – – – D. majeensis sp. nov. 2.67 – 6.02 12.32 – – – D. peque ~ no 3.51 – 6.85 11.61 – – – D. aff. quidditus 4.71 ƚ 0.78 (3.27 – 5.44) 12.19 ƚ 2.24 (10.35 – 16.71) – – – D. tinker 2.96 ƚ 2.05 (1.35 – 7.55) 25.38 ƚ 11.84 (7.76 – 39.16) 11 – 13 15.97 – 27.85 19.71 – 30.58 D. ventrimaculatus * 5.15 11.45 D. vocator 1.91 ƚ 0.57 (1.4 – 2.52) 32.98 ƚ 9.35 (23.7 – 42.39) 13.23 ƚ 10.51 (5.8 – 20.67) 19.25 ƚ 13.31 311.02 ƚ 304.7 (9.84 – 28.66) (95.56 – 526.47) Mean ƚ SD (range). Number in parentheses next to the species names represents the number of individuals analysed. DF, dominant frequency. *Information obtained from literature. palmar and plantar accessory tubercles indistinguishable; no nuptial pads; no fringes on fingers; hindlimbs of moderate length, TL 43% of SVL; relative lengths of adpressed toes I <II <III <V <IV; when adpressed, tip of toe I reaches to tubercle of toe II; disc of toe IV slightly expanded, 1.3 times wider than distal end of adjacent phalanx; no fringes on toes; between one and three nonprotuberant subarticular tubercles present (one each on toes I and II, two on toes III and V, and three on toe IV); inner metatarsal tubercle ovoid; outer metatarsal tubercles slightly pointed and smaller than inner; tarsal ridge absent; hands and feet without webbing; finger and toe discs even, broadened; ungual flap expanded, almost 278 A. BATISTA ET AL. flanks brick red (36); groin, axilla, and ventral areas mottled with brick red (36). Coloration in preservative Dorsal ground color raw amber (23), with a couple of dorsolateral lines light buff (2); groin and ventral areas buff (5), with small points sepia (279); ungual flaps dark drab (45). rounded; pads broadened and globular in profile ( Figs 7A , 10 ). Coloration of holotype in life Holotype (SMF 97304; Figs 10 , 14A, B ) recorded as follows: iris light Pratt’s rufous (71) with a couple of lateral and irregular lines tawny (60), lumbar region tawny (60), bordered by two lines light buff (2); Measurements of holotype (mm) SVL 17.40; HL 6.70; HW 6.30; IOD 3.24; EL 2.72; TD 0.92; FL 6.42; TL 7.50; HAL 3.47; 3FW 0.40; 3FD 0.64; 3TW 0.36; 3TD 0.49; 4TW 0.31; 4TD 0.67; BW 5.22 (for variation of the species, see Table 1 ). Vocalization The calls produced by two specimens from Cerro Sapo ( Fig. 6 ; Table 2 ), one paratype ( SMF 97313, environmental temperature 21.5 °C; humidity 84%; 22:06 h) and an uncollected specimen (environmental temperature 21.7 °C; humidity 80%; 21:00 h) were analysed. The calls consist of single, short, monophasic notes that are reminiscent of a ‘whistle’ ( Fig. 6 ). Note duration is 0.04 – 0.05 s , with an interval between calls of 16.91 – 16.25 s , and with a call rate of four calls per minute. The peak frequency band ranges from 2.79 to 4.30 kHz; the first harmonic contains the dominant frequency at 3.34 – 3.81 kHz . Natural history This species is found in the eastern Panamanian montane forest ( Fund & Hogan, 2012 ) along the PM and JSM ( Fig. 1 ). The vegetation consists predominantly of trees covered with moss, bromeliads ( Werauhia spp. and Guzmania spp. ), giant ferns ( Cyathea spp. ), and orchids. Diasporus darienensis sp. nov. is an inhabitant of the cloud forest ( 869 – 1169 m a.s.l.), usually found 1 – 5 m above ground. During the day, specimens seek retreats between bromeliad leaves. At night they actively move across tree bark and bromeliads. Only two males were encountered calling, both during the end of the rainy season (December) at the top of Cerro Sapo. One male (SMF 97313) was observed calling from the underside of a leaf in a tree about 5 m above ground, the other was calling from a branch on a ridge 3 m above the ground on the same day. Diet is not known, but as with other Diasporus it may eat small arthropods ( Batista, 2009 ). Etymology The species name is derived from the province name Darien where the holotype was found, with the Latin suffix - ensis donating a place or locality. The species is known to occur only in this province, and it is present in the main mountain ranges of the region.