Bioacoustics of Limnonectes lauhachindai

Description of holotype . Habitus moderately stocky; body tapering to groin. Head broad and depressed; head length and width subequal. Snout obtusely pointed in dorsal view, round in profile, projecting well beyond lower jaw in profile; nostril dorsolateral, much closer to tip of snout than to eye, below canthus, internarial distance subequal to interorbital distance; canthus rostralis indistinct, rounded, slightly constricted behind nostrils; lores concave, oblique; eye diameter 80% snout length, interorbital distance greater than upper eyelid width; pineal ocellus visible; tympanum round, not elevated from side of head, annulus weakly visible, tympanum diameter about 82% eye diameter and greater than distance between tympanum and eye; small, slit-like vocal sac openings on floor of mouth near lateral margin of tongue; vomerine teeth on two oblique ridges, equal in distance to each other as to choanae; two large odontoid processes at front of mandible; median triangular protuberance at mandibular symphisis. Forelimb moderately robust. Fingers moderately slender, without webbing; tip of fingers rounded, weakly expanded into discs; relative finger lengths II <IV <I <III; moveable flap of skin on preaxial sides of Fingers II– IV; distinct subarticular tubercles, one on Fingers I–II, two on Fingers III–IV; distinct palmar tubercles, one at base of Finger I, two in contact as base of Fingers II–IV; nuptial pad absent. Hindlimb moderately robust. Toes moderately slender; tips of toes rounded, expanded into small discs; relative toe lengths I<II<V<III<IV; webbing on Toe I to base of disc, on preaxial side of Toe II to midway between subarticular tubercle and tip and continuing as a fringe to base of tip, on postaxial side of Toe II to base of tip, on preaxial side of Toe III to level of distal subarticular tubercle continuing as a fringe to base of tip, on postaxial side of Toe III to base of tip, on preaxial and postaxial sides of Toe IV to level of distal subarticular tubercle and continuing as a fringe to base of tip, and on Toe V to base of tip; moveable flap of skin on outer margins of Toes I and V; distinct fold on distal half of tarsus; distinct, elongate, oval, inner metatarsal tubercle, length about 44% distance between tip of toe I and tubercle; no outer metatarsal tubercle. Skin above shagreened with irregular rows of large oval warts, most concentrated near flank and lower back; low-profiled, U-shaped caruncle with free posterior margin that fully occupies, but does not extend beyond, the interorbital region; distinct supratympanic ridge from posterior corner of eye to axilla; large rictal gland; no dorsolateral fold; skin on venter smooth. Measurements of holotype given in Table 2 . Measurement Holotype male n =1 All males n =20 Range; Mean ± SD Paratype females n =4 Range; Mean ± SD SVL 35.9 30.5–42.0; 33.8 ± 2.8 32.9–37.9; 35.4 ± 2.6 HDL 16.8 13.7–20.6; 15.8 ± 1.5 13.8–15.8; 14.8 ± 0.9 HDW 16.6 13.6–20.0; 15.3 ± 1.4 13.1–15.3; 14.2 ± 1.0 SNT 6.3 4.7–6.8; 5.6 ± 0.6 5.2–5.8; 5.5 ± 0.3 EYE 5.0 4.0–5.7; 4.5 ± 0.4 4.7–5.1; 4.9 ± 0.2 IOD 3.6 2.9–4.2; 3.6 ± 0.3 2.1–3.6; 2.9 ± 0.6 TMP 4.2 3.0–5.0; 3.7 ± 0.4 2.9–3.4; 3.1 ± 0.2 IND 3.5 3.0–4.0; 3.4 ± 0.2 2.8–3.7; 3.3 ± 0.4 SHK 17.0 13.9–19.1; 15.8 ± 1.2 17.1–18.6; 17.7 ± 0.8 TGH 17.8 14.2–20.4; 16.7 ± 1.4 15.3–19.0; 17.3 ± 1.8 LAL 7.5 6.3–8.7; 7.0 ± 0.6 6.9–8.2; 7.5 ± 0.6 HND 8.5 6.9–9.8; 7.9 ± 0.7 6.8–9.0; 8.1 ± 1.0 FTL 16.7 14.1–19.2; 15.9 ± 1.1 16.6–19.0; 17.9 ± 1.1 IML 2.0 1.6–2.5; 2.0 ± 0.2 1.7–2.7; 2.2 ± 0.4 IMW 0.9 0.9–1.2; 1.0 ± 0.1 0.8–1.1; 1.0 ± 0.1 Color of holotype in preservative. Dorsum dark brown; tips of dorsal warts white, warts encircled with black. Venter cream, chin dark brown, throat with large brown spots, belly and ventral surfaces of limbs with very small dark spots. Color of paratype ZMKU AM 0 0 586 in life. Dorsum brown, with irregular black spots, becoming brassy on dorsal surfaces of limbs and upper flank; continuous black streak under canthus and supratympanic fold, extending from nostril to upper half of tympanum; lips brown with broad black bars; iris bronze; broad cream-yellow vertebral stripe from anterior margin of upper jaw to vent; upper surfaces of hindlimb with broad black bands; lower flank beige and gray ( Fig. 2 ). Variation. Females lack caruncles and have narrower heads in dorsal view than males. The largest male paratype (ZMKU AM 01111; SVL 42.0 mm) is noticeably larger than the second largest (NCSM 81269; SVL 37.7 mm ). Seven (NCSM 81269, ZMKU AM 0 1106, ZMKU AM 00552–53, ZMKU AM 00586–87, ZMKU AM 00593) of 25 (28%) specimens have a pale vertebral stripe, a polymorphic character seen in other Limnonectes , including L. dabanus and L. kohchangae ( Stuart & Emmett 2006 ; Stuart et al. 2006b ). Measurements are summarized in Table 2 . Molecules. The aligned dataset contained 1,624 characters. The new species was recovered as sister to a clade containing L. dabanus and L. gyldenstolpei ( Fig. 3 ). The holotype and six paratypes (NCSM 81269, ZMKU AM 01104–07, ZMKU AM 01109) are identical in the 16S gene fragment, but have an uncorrected pairwise divergence of 5.98–6.72% from L. dabanus ( n =2) and 4.25–8.40% from L. gyldenstolpei ( n =3). Calls. Paratype male ZMKU AM 0 0 589 had two different advertisement call types , referred to here as Type 1 and Type 2 ( Fig. 4 ). Type 1 (n = 30) had 1–10, one-pulsed notes lasting 27.44 to 4342.91 ms; note duration of 10.29–34.50 ms; interval between notes 182.50–972.41 ms; note repetition rate of 0.98–4.00 notes per second; and dominant frequency of 1.12–2.33 kHz. Type 2 (n = 3) was a single, multi-pulsed note lasting 1749.06–2148.03 ms with 31–37 pulses; pulse duration of 21.24–25.00 ms; interval between pulses 33.74–35.36 ms; pulse rate 16.98– 17.49 pulses per second; and dominant frequency of 2.15–2.24 kHz ( Table 3 ). The advertisement calls were repeated at a rate of approximately 0.12 calls per second and intercall interval varied from 0.36 to 30.26 s. These calls showed frequency modulation with or without weak harmonics.