Mannophryne herminae

Remarks: The name herminae has long been associated with the abundant populations found in the Cordillera de la Costa. Unpublished genetic data (by J. C. Santos) indicates some of these populations belong to an undescribed species. Mannophryne herminae sensu stricto apparently is found only in San Esteban (Carabobo state) and surrounding areas, near Puerto Cabello. The identity of other populations from Cordillera de la Costa traditionally determined as M. herminae need confirmation. Rojas-Runjaic et al. (2018) redefined M. herminae and restricts Mannophryne herminae sensu stricto to the northern slope of Cordillera de la Costa between Carabobo and Aragua states. In the Mannophryne collaris species group (Manzanilla et al. 2007; Grant et al. 2017). Selected references: Boettger (1893); Stejneger (1901); Lutz (1927); Schmidt (1932); Alemán (1952); Mertens (1957 b); Ginés (1959); Sexton (1960); Rivero (1961, 1964 a, 1988); Test (1962); Gremone et al. (1986); La Marca (1991 c “ 1994, ” 1992, 1994 a, 1995 b); Myers et al. (1991); Manzanilla et al. (1995); Barrio-Amorós (2006 c); Barrio-Amorós et al. (2010 b); Rojas-Runjaic et al. (2018).

Distribution: Region 2. Central Cordillera de la Costa (to be better determined).

Type locality: “ Puerto Cabello in Venezuela. ”

Call description. Advertisement calls of Mannophryne herminae are composed of long trills of short tonal notes arranged in duplets (Fig. 12 a, c). The notes show amplitude modulation and the number of amplitude peaks typically varies between 2 – 3 (Fig 12 c). The duration of calls between the two males recorded was 30.16 ± 15.27 (7.00 – 49.49) s. The rate of note emission was 11.02 ± 0.17 (10.60 – 11.29) notes / s, and each call contained 331.73 ± 166.98 (78 – 546) notes. Note duration was 0.03 ± 0.00 (0.02 – 0.04) s, and the duration of silent interval between notes of each duplet was 0.03 ± 0.00 (0.02 – 0.04) s; the duration of duplets was 0.08 ± 0.00 (0.08 – 0.10) s, and duration of silent intervals between duplets 0.10 ± 0.01 (0.09 – 0.14) s. Notes have a very slight ascending frequency modulation (Figs. 12 b, d). Fundamental frequency was 2.05 ± 0.04 (1.98 – 2.15) kHz. Peak frequency was 4.04 ± 0.09 (3.79 – 4.24) kHz, and its lower and upper frequencies were 3.89 ± 0.10 (3.65 – 4.11) kHz and 4.17 ± 0.10 (3.93 – 4.39) kHz, respectively; the bandwidth was 0.28 ± 0.04 (0.21 – 0.42) kHz. Three additional harmonics (3 rd – 5 th; Fig. 12 d) were observed above peak frequency, at 5.68 – 6.29, 7.60 – 8.44, and 9.52 – 10.51 kHz. Some small differences were observed in the spectral parameter values between the first and second note of each duplet, being the second note the one that shows the higher values. Discriminated temporal and spectral values for the first and second note of each duplet are listed in Table 2.

Remarks: Previously considered a species widely distributed in northern Venezuela (Rivero 1984 b; La Marca 1992, 1994). In the last two decades it has been demonstrated that in fact as formerly defined geographically and morphologically, Mannophryne herminae constitutes a conglomerate of different species-level taxa, and until today five species (M. caquetio, M. lamarcai, M. leonardoi, M. venezuelensis and M. vulcano) have been described from populations originally considered as corresponding to M. herminae (Mijares-Urrutia & Arends 1999 a, b; Manzanilla et al. 2007 a, b; Barrio-Amorós et al. 2010 a). However, after the above mentioned species delimitations, the geographic distribution of M. herminae is yet wide and probably there are still several different species under this name. In order to facilitate the future recognition of undescribed taxa related to M. herminae, we only consider as pertaining to this species the populations from the northern slope of Cordillera de la Costa that fit with the amended definition provided below, which is based on the type series (Figs. 9 – 10) and on additional specimens (Fig. 11) from the lower basins of San Esteban (near Puerto Cabello, the type locality), Patanemo, Cata, and Cuyagua rivers, and from Rancho Grande. Regarding the type locality, as previously commented by La Marca (1994) Puerto Cabello probably only was the port of shipping of the type series and not the place where it was collected. Puerto Cabello is located within a Tropical dry forest area, an unlikely habitat for a collared frog. The type series most likely comes from the vicinities of Puerto Cabello (e. g. San Esteban valley), where small rivers surrounded by Premontane very humid forests and inhabited by Mannophryne herminae populations still occur. Amended definition: (1) Small body size, with adult males smaller than females (males: 20.5 – 23.1 mm of SVL vs. females: 24.6 – 29.0 mm); (2) dorsal skin of body and hind limbs shagreen, moderately granular on flanks; small tubercles present on posterior third of dorsum, body flanks, and dorsal surface of thighs and shanks; ventrally smooth; (3) snout rounded to nearly truncate in dorsal view, rounded to protruding in profile; (4) nares visible ventrally; (5) tympanum small (~ 1 / 3 – 2 / 5 of ED), defined, about 1 / 4 – 1 / 3 concealed posterodorsally; (6) short teeth present on maxillary arch; (7) median lingual process absent; (8) vocal sac single subgular in males; (9) carpal pad absent; (10) metacarpal ridge low; (11) thenar tubercle conspicuous; (12) nuptial excrescences on thumb absent; (13) FIII not swollen; (14) tip of FIV surpassing distal subarticular tubercle of FIII; (15) FI and FII equal in size; (16) thin lateral fringes present on preaxial side of FII – FIII; (17) weak and poorly defined lateral keels on pre- and postaxial sides of FI and FIV, and postaxial side of FII – FIII; (18) tarsal keel well-defined in all its extension, nearly straight, extending from the base of TI where is continuous with the preaxial fringe, to the mid-tarsus, not merged with the inner metatarsal tubercle nor reduced at the level of this tubercle; (19) tarsal fringe absent; (20) middle metatarsal tubercle present, similar in size to the inner one, non-protuberant, weakly defined; (21) metatarsal fold present, strong, almost reaching the outer metatarsal tubercle; (22) wide lateral (pre- and postaxial) fringes in all toes; (23) toes basally webbed, webbing formula: I (1 + – 2 –) – (2 + – 2 ¾) II (1 ½ – 2 –) – (3 – – 3 ⅓) III (2 ⅓ – 3 –) – (3 ¾ – 4 +) IV (4 + – 4 ½) – (2 ½ – 3 –) V; (24) disc weakly expanded on FI, moderately expanded on FIII – FIV; moderately expanded TI – TIV, and weakly to moderately expanded on TV; (25) paired dorsal digital scutes present on fingers and toes; (26) cloacal sheath short; (27) supracloacal dermal flap present, conspicuous; (28) cloacal tubercles present; (29) iridescent golden to cream spot at dorsal forelimb and hind limb insertions present, diffuse; (30) pale paracloacal mark present, diffuse; (31) diffuse yellowish spots on hidden parts of hind limbs absent; (32) thigh dorsally pale brown with three to four transverse dark brown bands; (33) pale dorsolateral stripe solid to diffuse, straight, reaching the level of the arm insertion or up to the level of the groin; (34) lateral dark band solid, wider at the middle of the flank; (35) oblique lateral stripe partial (not reaching the posterior border of the eye), solid, cream or subtly tinged with yellow at the groin; (36) ventrolateral stripe cream or yellowish, poorly defined, formed by a wavy series of spots; (37) dark dermal collar wide, diffuse to solid, non-speckled with small white dots and complete in males; broad (10.9 – 15.9 % SVL), solid, with or without some small whitish dots, and complete in females; (38) dark lower labial stripe present, diffuse; throat color in life: extensively colored with yellow in females, gray in males; (39) abdomen color white to spotted with yellow (in life), and almost free to irregularly stippled with melanophores in females; pale gray, evenly stippled with melanophores in males; (40) iris bronze, finely reticulated of black, with a dark horizontal band, pupil ring bronze, complete; (41) tongue ocher yellow; (42) large intestine unpigmented; (43) adult testis unpigmented; (44) mature oocytes with the animal pole pigmented with dark brown; (45) skin blackening in males during call activity; (46) mercaptanlike odor present; (47) diurnal activity; (48) tadpole transport by males; (49) riparian habitat; (50) advertisement call composed of long trills of tonal notes arranged in duplets (51) peak frequency of calls: 3.79 – 4.24 (4.04 ± 0.09) kHz; (52) fundamental frequency: 1.98 – 2.15 (2.05 ± 0.04); (53) rate of note emission: 10.60 – 11.29 notes / s (11.02 ± 0.17); (54) note duration: 0.02 – 0.04 (0.03 ± 0.00) s; (55) duration of silent intervals between notes of a duplet: 0.14 – 0.57 (0.31 ± 0.10) s; (56) duration of silent intervals between duplets: 0.09 – 0.14 (0.10 ± 0.01) s.