Bioacoustics of Pristimantis relictus

Description of the holotype . Adult male (URCA-H 13344; Figure 2 ). Head longer than wide; head width 37% of SVL; head length 40% of SVL; snout subovoid in dorsal view, rounded in lateral view; nostrils not protuberant, directed anterolaterally; internarial distance less than eye-nostril distance; canthus rostralis angular, weakly concave, loreal region slightly concave; eye large, with horizontally elliptical pupil; eye diameter slightly larger than interorbital distance; cranial crests absent; upper eyelid smooth, 74% of eye diameter; tympanum large, with distinct rounded tympanic annulus; tympanic membrane present, visible; supratympanic fold present; labial bars absent; vocal sac subgular, small; tongue ovoid covering the entire floor of mouth, free and notched behind; vocal slits present, lateral to tongue; choana small, rounded; odontophores oblique and widely separated posterior to the choanae, each one bearing six teeth; skin on dorsum shagreen with scattered low tubercles; dorsolateral fold absent; skin on venter areolate, weakly spotted, discoidal fold present; hands large, 29% of SVL; relative lengths of fingers II <IV <I <III; finger fringes absent; finger discs expanded, truncate; disc of Finger III wider than the others, discs of fingers II and IV wider than disc of Finger I; nuptial pad present, single; outer metacarpal tubercle divided inner metacarpal tubercle large, in contact with nuptial pad; subarticular tubercles large, rounded; one subarticular tubercle on fingers I and II, and two on fingers II and IV; supernumerary tubercles present; three horizontal bars present on thigh and tibia; tibia length greater than thigh length, tibia length 51% of SVL; tarsal fold present; inner metatarsal tubercle present, larger than outer metatarsal tubercle, both rounded; toes bearing narrow lateral fringes; webbing absent; relative lengths of toes I <II <III <V <IV; toe discs expanded, smaller on Toe I; subarticular tubercles rounded, one on toes I and II; two on toes III and V, and three on Toe IV; supernumerary tubercles present, barely distinct. Measurements of the holotype (mm). SVL 24.4, HL 9.7, HW 8.9, ED 3.4, SL 4.4, IND 2.0, END 2.8, IOD 3.3, UEW 2.5, TD 2.0, FAL 5.4, HAL 7.0, FIII 1.4, FIV 1.3, THL 12.4, TL 13.1, TAL 7.1, FL 10.7, and TIV 1.4. Color in preservative . Dorsal and lateral surfaces pale cream, snout light brown, brown interocular band in dorsal view, distinct dark brown stripe from tip of snout to posterior region of supratympanic fold. Irregular brown marks in lower orbital region. Two brown chevrons on dorsum with irregular brown dots in inguinal region, light brown stripes on dorsal surfaces of limbs. Ventral surface background pale cream in color. Advertisement call. The call of Pristimantis relictus sp. nov. is described based on three individuals (MNRJ 55582 and two unvouchered individuals) (see Table 2 ;). The vocalization is composed of 1–8 multi pulsed notes (2–5 pulses per note), with an ascendant amplitude modulation. The call duration ranges 0.02– 0.66 s (0.06 ± 0.08; n = 181), with intercall interval of 0.8– 9.6 s (2.12 ± 1.5; n = 74), note duration ranges 0.02– 0.05 s (0.03 ± 0.01; n = 150), with a note rate of 0.52–2.20 notes per second ( Figure 3 . The call has three visible bands. The fundamental frequency is in the first band, ranging 1894.9–2153.3 Hz, followed by a second band located in the dominant frequency, ranging 3617.6–4220.5 Hz, and a third weak band with a higher frequency, ranging 4565–6287.7 Hz. Acoustic parameters Pristimantis relictus sp. nov. Peak Frequency (Hz) 3922.8 ± 125.6 (3617.6–4220.5, n =149) Fund. Frequency (Hz) 2032.5 ± 54.7 (1894.9–2153.3, n =149) Third band Frequency (Hz) 5692.3 ± 341.7 (4565–6287.7, n =149) Note duration (s) 0.03 ± 0.01 (0.02–0.05, n =150) Call duration (s) 0.06 ± 0.08 (0.02–0.66, n =181) Notes/call 1.94 ± 1.24 (1–8, n = 151) Note rate(s) 1.39 ± 0.8 (0.52–2.20, n =3) Pulses/call 2.6 ± 1.0 (2–5, 386) Intercall interval (s) 2.12 ± 1.5 (0.8–9.6, n =74) Males ( n = 18) Females ( n = 33) SVL 25.4 ± 1.7 (23.1–29.7) 30.5 ± 1.5 (26.8–32.8) HL 9.7 ± 0.6 (9.0–11.3) 12.4 ± 1.1 (10.8–15.9) HW 8.9 ± 0.7 (7.9–10.9) 11.6 ± 0.6 (9.9–12.3) ED 3.2 ± 0.4 (2.6–4.3) 3.7 ± 4.2 (3.1–4.2) SL 4.5 ± 0.3 (3.8–5.4) 5.5 ± 0.4 (4.6–6.4) IND 2.1 ± 0.2 (1.7–2.6) 2.6 ± 0.3 (1.9–3.3) END 3.2 ± 0.4 (2.7–4.2) 3.9 ± 0.4 (3.1–4.7) IOD 3.1 ± 0.4 (2.4–3.7) 4.0 ± 0.4 (3.3–4.8) UEW 3.3 ± 0.8 (2.2–4.3) 4.0 ± 0.8 (2.6–5.3) TD 1.9 ± 0.2 (1.6–2.5) 2.2 ± 0.2 (1.8–2.5) FAL 5.8 ± 0.5 (5.1–6.8) 7.3 ± 0.5 (6.4–8.4) HAL 6.8 ± 0.4 (6.1–7.7) 8.4 ± 0.6 (6.8–9.9) FIII* 1.5 ± 0.1 (1.4–1.6) 1.5 ± 0.1 (1.3–1.6) FIV* 1.4 ± 0.1 (1.3–1.5) 1.4 ± 0.1 (1.2–1.7) THL 12.3 ± 0.9 (10.2–13.8) 15.2 ± 1.3 (12.8–18.4) TL 13.6 ± 0.6 (12.5–15.5) 17.0 ± 1.2 (14.9–19.8) TAL 7.3 ± 0.5 (6.4–8.3) 9.2 ± 1.1 (7.4–12.3) FL 10.7 ± 0.6 (9.9–12.6) 13.4 ± 0.8 (11.8–14.9) TIV* 1.3 ± 0.1 (1.2–1.4) 1.4 ± 0.1 (1.2–1.6) Variation among paratypes . Adult females (SVL 26.8–32.8 mm ) are slightly larger than adult males (SVL 23.1–29.7 mm ) ( Table 3 ). Variation in coloration of fixed specimens is mostly related to the presence of chevrons on the dorsum (URCA-H 6980, 2373, 1544, 13342, 6988, and 9828). In life, the dorsal coloration is highly polymorphic: dark brown, reddish to yellowish-brown, or grey ( Figure 4 ), with the presence of dark brown chevrons and black spots in some individuals. Other variations in coloration among paratypes are: brown bars on the leg and arm; dark brown stripe from the tip of the nose to the eye, including the internarial area; dark brown bar in the interorbital region; labial region yellowish to white; supratympanic fold bold black; gular region yellowish; and iris cooper-colored with brown reticulations. Geographic distribution. Pristimantis relictus is endemic to the “Brejos de Altitude” of the state of Ceará , Northeast Brazil . This species occurs in the Planalto da Ibiapaba (municipalities of Ipu, Tianguá, Ubajara, Granja, Viçosa do Ceará ), Serra de Maranguape (municipality of Maranguape), Serra da Aratanha (municipality of Pacatuba), Serra de Baturité (municipalities of Guaramiranga, Pacoti, Aratuba, Mulungu), Serra da Uruburetama ( Roberto & Loebmann 2016 ), and Serra da Meruoca (municipality of Meruoca) ( Figure 5 ). Natural history. Pristimantis relictus occurs in dry forests and humid forests ( Loebmann & Haddad 2010 ; Roberto & Loebmann 2016 ; Castro et al . 2019 ), from 100 to 900 m .a.s.l. The species has a prolonged breeding period throughout the raining season from December to April. Vocalization occurs from 5 pm throughout the night, with a peak of individuals vocalizing during 6–9 pm. Individuals were found vocalizing on leaves, tree trunks and twigs, mostly facing downwards. Amplexus is axillary, and the eggs are deposited on humid surfaces of rock and wooden debris. Mature females range 28–33 mm and 1.45–3.27 g (n = 13), and clutches have 21– 53 eggs (n = 13). We observed territorial behavior involving agonistic combat and the emission of territorial and release calls; unfortunately, these calls were not recorded. At Serra de Baturité mountain range we recorded the following species occurring in sympatry with P. relictus : Boana raniceps , Dendropsophus minusculus , D. minutus , D . tapacuarensis , Pithecopus gonzagai , Scinax tropicalia , Trachycephalus typhonius , Adelophryne baturitensis , Odontophrynus carvalhoi , Rhinella casconi , R. dapsilis , R. diptycha , R. granulosa , Adenomera juikitam , Leptodactylus macrosternum , L. natalensis , L. pustulatus , L. mystaceus and Physalaemus cuvieri . See Loebmann & Haddad (2010) , and Castro et al . (2019) for the anuran species that occur in sympatry with Pristimantis relictus at the Planalto da Ibiapaba mountain range.