Hypsiboas maculateralis, Caminer, Marcel A. & Ron, Santiago R., 2014

Caminer, Marcel A. & Ron, Santiago R., 2014, Systematics of treefrogs of the Hypsiboas calcaratus and Hypsiboas fasciatus species complex (Anura, Hylidae) with the description of four new species, ZooKeys 370, pp. 1-68 : 26-32

Hypsiboas maculateralis sp. n.

Common name.

English: Stained treefrog; Spanish: Rana arbórea manchada

Holotype.

(Fig. 14F) QCAZ 40082 (field no. SC-PUCE 23221), adult male from Ecuador, Provincia Napo, Comunidad Santa Rosa (1.02337°S, 77.48359°W), 354 m above sea level, collected by Marcel A. Caminer and Edwin Carrillo Ponce on 28 February 2009.

Paratypes.

ECUADOR: PROVINCIA ORELLANA: Río Napo, Santa Teresita, 4 km NW from Nuevo Rocafuerte (0.9008°S, 75.4135°W), 186 m, QCAZ 44651, adult female, 44673, adult male, collected by S. R. Ron, E. Toral, and I. G. Tapia on 9 July 2009; Río Napo, Huiririma (0.7116°S, 75.6239°W), 194 m, QCAZ 44636, adult male, collected by S. R. Ron, E. Toral and I. G. Tapia on 8 July 2009; Río Napo, San Vicente (0.6790°S, 75.6511°W), 203 m, QCAZ 44531, adult female, 44532, adult male, collected by S. R. Ron, E. Toral and I. G. Tapia on 7 July 2009; Río Napo, Chiroisla (0.5756°S, 75.8998°W), 203 m, QCAZ 44452, adult male, collected by S. R. Ron, E. Toral and I. G. Tapia on 5 July 2009; Río Napo, Edén (0.4983°S, 76.0711°W), 216 m, QCAZ 44182-83, 44186, 44188-90, adult males, collected by S. R. Ron, E. Toral and I. G. Tapia on 4 July 2009; Río Napo, sector La Primavera (0.4442°S, 76.7868°W), 244 m, QCAZ 43825, adult female, 43827, adult male, QCAZ 43800, juvenile, collected by S. R. Ron, E. Toral and I. G. Tapia on 29 June 2009; Río Napo, El Descanso (0.4310°S, 76.7864°W), 244 m, QCAZ 43897, adult male, collected by S. R. Ron, E. Toral and I. G. Tapia on 30 June 2009; El Coca (0.4778°S, 76.9898°W), 267 m, QCAZ 43712, adult male, 43709-10, 43715, ju veniles, collected by S. R. Ron, E. Toral and I. G. Tapia on 27 June 2009; Guiyero community (0.6248°S, 76.4944°W), 227 m, EPN 10319, adult male, collected by A. Almendáriz and J. Hernández; Guiyero community, Nambai km 32 (0.6248°S, 76.4944°W), 240 m, EPN 10887, adult male, collected by A. Almendáriz, J. Awa and P. Ima; Parque Nacional Yasuní, km 74 on the road Pompeya-Iro (0.8331°S, 76.3416°W), 257 m, QCAZ 5238, adult female, collected by S. de la Torre and S. R. Ron on 20 February 1994; Nuevo Rocafuerte, Tambococha (0.9783°S, 75.4256°W), 177 m, QCAZ 55359, adult male, 55395, 55374, adult females, collected by F. Ayala-Varela, E. Carillo, J. Brito, A. Varela and D. Quirola on 13 March 2013; PROVINCIA Sucumbíos: Zábalo, Familia Criollo (0.3181°S, 75.7662°W), 220 m, QCAZ 28004, 28024, adult females, collected by M. R. Bustamante, N. Acosta-Buenaño, M. Guerra and C. Proaño on 30 September 2003 and 2 October 2003; Playas de Cuyabeno (0.2654°S, 75.8917°W), 230 m, QCAZ 28401, adult female, collected by M. R. Bustamante, N. Acosta-Buenaño, M. Guerra and C. Proaño on 8 October 2003; Playas de Cuyabeno, Agurico Pañacoha (0.3106°S, 76.0335°W), 200 m, EPN 13452, adult male, collected by P. Meza Ramos; Putumayo, Güepi camp 20 km N of Zábalo (0.1779°S, 75.6767°W), 220 m, EPN 7189, adult female, collected by A. Almendáriz; Reserva Faunística Cuyabeno (0.0849°N, 76.1344°W), 273 m, QCAZ 2148, 29454, adult males, 2156, adult female, collected by S. de la Torre and F. Campos on 13 May 1990 and 9 May 2004; Río Napo, La Selva Lodge (0.5086°S, 76.3649°W), 229 m, QCAZ 44020-21, adult males, collected by S. R. Ron, E. Toral and I. G. Tapia on 2 July 2009; Indillana, (0.43370°S, 76.5419°W), 265 m, QCAZ 24672, subadult, collected by F. Ayala-Valera on 11 January 2002; Campo Vinita, on the road Palma Roja-Pto El Carmen de Putumayo (0.1220°N, 75.8611°W), 217 m, QCAZ 29280-82, adult males, collected by S. Aldás-Alarcón on 7 May 2004; PROVINCIA PASTAZA: Bataburo Lodge, S of the road from Cononaco (1.2083°S, 76.7166°W), 250 m, QCAZ 39430, 39452, adult males, collected by S. D. Padilla and C. Meyer on 10 October 2008 and 13 October 2008.

Referred specimens.

ECUADOR: PROVINCIA SUCUMBÍOS: Reserva de Producción Faunística Cuyabeno, Estación Científica Universidad Católica del Ecuador, Laguna Grande (0.0195°S, 76.1712°W), 220 m (QCAZ 6141-43, 6146-51); PERU: REGION MADRE DE DIOS: Manu National Park, Cocha Cashu Biological Station, Río Manu (11.9166°S, 71.3°W), 340 m (USNM 299774).

Diagnosis.

A member of the genus Hypsiboas characterized by: (1) mean SVL 36.00 mm in males (range 31.86-39.17; n = 22), 45.18 mm in females (range 32.04-55.31; n = 6); (2) basal webbing on fingers; (3) calcar large, triangular; (4) dorsal coloration varying from creamy white to reddish brown, pinkish white or brown, sometimes with dark markings (e.g., narrow longitudinal lines, broad transversal marks, or large black stains); (5) dark brown middorsal line restricted to the head or the anterior half of the body often present; (6) flanks pale cream or creamy white (in life, blue in large females and light blue or white in males) with dark brown blotches; (7) hidden surfaces of thighs pale cream or gray (in life, blue in large females and light blue or white in males) with dark brown blotches; (8) ventral surfaces of thighs yellowish white or brown; (9) ventral coloration varying from creamy white to yellowish white with or without brown flecks on the neck and chest; (10) webbing on feet; (11) in life, iris cream silver, yellowish or cream with faint yellow to orange color on its upper quarter; (12) prepollical spine present in males.

Hypsiboas maculateralis (Figs 4D, 8D and 16) differs from Hypsiboas fasciatus , Hypsiboas almendarizae , and Hypsiboas calcaratus in advertisement call (Figs 12 and 13 A–B) and by having dark blotches on the flanks and thighs (dark lines in Hypsiboas almendarizae , Hypsiboas calcaratus , and Hypsiboas fasciatus ). Hypsiboas maculateralis can be distinguished from Hypsiboas alfaroi sp. n. and Hypsiboas tetete sp. n. by the presence of a calcar (instead of a small tubercle on the heel). Morphological characters useful to differentiate Hypsiboas maculateralis from other species are shown in Table 6.

Description of holotype.

Adult male, 36.16 mm SVL, foot length 14.33 mm, head length 8.98 mm, head width 11.71 mm, eye diameter 4.07 mm, tympanum diameter 2.44 mm, tibia length 20.29 mm, femur length 18.44 mm, calcar length 1.44 mm, arm length 6.36 mm, eye-nostril distance 2.86 mm, head wider than long and wider than body; snout rounded in lateral view, truncate in dorsal view; distance from nostril to eye shorter than diameter of eye; canthus rostralis indistinct, rounded; loreal region concave; internarial area convex; nostrils not protuberant, directed laterally; interorbital area slightly convex; eye large, strongly protuberant; diameter of eye 1.7 times diameter of tympanic annulus; tympanum concealed beneath skin;tympanic annulus evident, ovoid, longer dorsoventrally and concealed dorsally by supratympanic fold, separated from eye by ca. 1.1 times its diameter; posterior end of supratympanic fold reaches anterior border of arm insertion. Arm slender, axillary membrane absent; indistinct low tubercles along ventrolateral edge of forearm; relative length of fingers I<II<IV<III; fingers bearing large, oval discs, that of third finger about three fourths of tympanum diameter; subarticular tubercles prominent, round to ovoid, single; supernumerary tubercles present; palmar tubercle small, elongated; prepollical tubercle large, flat, elliptical; prepollex enlarged, claw shaped; nuptial excrescences absent; webbing absent between fingers. Large triangular calcar on tibiotarsal articulation; ill defined, scattered tubercles on tarsus and along ventrolateral edge of foot; toes bearing discs slightly wider than long, smaller than those of fingers; relative length of toes I<II<V<III<IV; outer metatarsal tubercle ill defined, small, round; inner metatarsal tubercle large, elliptical; subarticular tubercles single, round, flat; supernumerary tubercles restricted to the soles; webbing formula of toes I2-2+II 1¾-3-III2--3IV3-1½ V. Skin on dorsum, head, and dorsal surfaces of limbs smooth; skin on flanks smooth with weak longitudinal wrinkles posterior to the arm; skin on venter coarsely granular; skin on ventral surfaces of head and thighs granular, those of shanks smooth. Cloacal opening directed posteriorly at upper level of thighs; cloacal sheath short and simple, covering cloacal opening; round tubercles below and on sides of vent, larger proximally. Tongue ovoid, widely attached to mouth floor; vomerine odontophores triangular with arched base, narrowly separated, posteromedial to choanae, each bearing 8-12 teeth; choanae ovoid.

Color of holotype in preservative. Dorsum pale reddish brown with two faint brown narrow longitudinal lines interconnected in the scapular region (Fig. 16); scattered minute black dots distributed on dorsum; dark brown middorsal line along snout; dorsal surfaces of limbs pale reddish brown with five pale brown narrow transversal bars on each thigh; flanks and hidden surfaces of thighs pale cream with dark brown blotches; narrow white stripe above vent; venter creamy white; ventral surfaces of limbs and webbing yellowish white; narrow to wide pale brown stripe on the outer edge of the hands, forearms, thighs, tarsal folds, and feet; minute brown blotches on lips; bones white.

Color of holotype in life. (Fig. 14F). Dorsum creamy white with six narrow pale brown longitudinal lines; a dark brown middorsal line; five pale brown faint transversal bars on dorsal surface of each thigh and two narrow pale brown longitudinal lines on dorsal surface of each shank; scattered minute black dots on dorsum; flanks and hidden surfaces of the thighs light-bluish white with dark brown blotches; discs and webbing pale yellowish tan; iris cream with faint yellow coloration on its upper margin.

Etymology.

The specific name is derived from the Latin words macula = stain, and lateralis = lateral, in reference to the brown dark blotches on the flanks of these frogs.

Variation.

Variation in dorsal and ventral coloration of preserved specimens is shown in Figure 16. Background dorsal coloration varies from creamy white (e.g., QCAZ 44020) to reddish brown (e.g., QCAZ 28401), pale reddish brown (e.g., QCAZ 28004), pale pinkish white (e.g., QCAZ 44021, 43897), pinkish white (e.g., QCAZ 44531, 44651, 44184) or brown (e.g., QCAZ 43825). A dark middorsal line extends from the tip of the snout to the middle of dorsum (e.g., QCAZ 2156), but in some specimens is restricted to the snout (e.g., QCAZ 43825, 44651, 28004) or is absent (e.g., QCAZ 44531). There is variation in the number, size, and shape of the dorsal marks. Five to seven broad transversal bands on the dorsum (sometimes interconnected) and narrower brown transversal bars on the dorsal surfaces of the limbs (one or two each on upper arm and forearm and three to five each on thigh, shank and foot) may be present (e.g., QCAZ 28401). In some individuals, faint, narrow longitudinal dark lines may be present (e.g., QCAZ 43897). In a few specimens, the dorsum, dorsal surfaces of forearms, and shanks may present large black stains (e.g., QCAZ 44021) and/or few scattered brown dots (e.g., QCAZ 43897, 44020). The coloration of the flanks and hidden surfaces of thighs vary from pale cream to creamy white or gray, with dark brown blotches. There is variation in the distribution of blotches on flanks; the extent covered by the blotches ranges from groin to the mid-flank (e.g., QCAZ 28004) or to the axilla (e.g., QCAZ 55374). In some individuals, similar blotches can also be present on the hidden surfaces of the shanks, ventral surfaces of the forelimbs, and dorsal surfaces of the feet (e.g., QCAZ 43825). In few individuals, the dark blotches of the thighs and flanks are faint (e.g., QCAZ 43897).

Ventral surfaces of preserved specimens vary from creamy white (e.g., QCAZ 44020, 44184; Fig. 16) to yellowish white (e.g., QCAZ 28004). Some individuals have brown flecks on the gular region, chest, and lips (e.g., QCAZ 43825). A narrow to wide brown stripe is present in some individuals on the outer edge of the hands, forearms, thighs, tarsal folds and feet (e.g., QCAZ 28004, 28401, 44021). Coloration of webbing and disc varies from yellowish white to brown or gray. Vomerine odontophores are triangular (with arched base in some specimens). Coloration of bones is white.

In the adult type series, the largest male has a SVL of 39.17 mm, and the largest female 55.31 mm; mean male SVL = 36 mm (n = 22; SD = 1.92), mean female SVL = 45.18 mm (n = 6; SD = 9.39). Females are significantly larger than males (t = -3.43, df = 4, P = 0.025). Inter-population variation in size and other morphometric variables are shown in Tables 1 and 2.

Coloration in life.

(based on photographs; Figs 4D and 8D). Dorsal surfaces vary from pale yellowish tan (e.g., QCAZ 43827) to yellowish cream (e.g., QCAZ 44636) with a middorsal dark brown line (e.g., QCAZ 43827); sometimes, ill-defined broad pale brown transversal bands (e.g., QCAZ 43827) or narrow pale brown longitudinal lines are present on the dorsum (e.g., QCAZ 44636); in some individuals, there are pale brown transversal bars on the dorsal surfaces of the limbs (e.g., QCAZ 44636); scattered minute black dots may be present on the dorsum (e.g., QCAZ 44636); flanks white, light blue or blue with dark brown blotches (e.g., QCAZ 43825, 43827, 44636); hidden surfaces of thighs white, gray, orange, light blue or blue with dark brown blotches (e.g., QCAZ 43825, 43825, 44636); in some specimens, hidden surfaces of the shanks, ventral surfaces of the upper arms and dorsal surfaces of the feet are blue with dark brown blotches (e.g., QCAZ 43825); venter creamy, sometimes with yellowish on its anterior half and with brown flecks on the neck, chest, and lips (e.g., QCAZ 43827); ventral surfaces of hindlimbs and forelimbs translucent white (e.g., QCAZ 43827) or translucent orange (e.g., QCAZ 44636); in some specimens, ventral surfaces of the thighs creamy white (e.g., QCAZ 43825); discs and webbing brown (e.g., QCAZ 43825), orange (e.g., QCAZ 44636) or pale yellowish tan (e.g., QCAZ 43827); a narrow to wide brown stripe may be present on the outer edge of the hands, forearms, thighs, feet, and tarsal folds (e.g., QCAZ 43825, 43827); iris yellowish (e.g., QCAZ 43715) or cream with faint yellow to orange color on its upper quarter (e.g., QCAZ 43825); bones vary from white to green (e.g., QCAZ 43827, 44636). Juveniles have similar coloration to adults, except that the blotches on the flanks and hidden surfaces of thighs are absent (e.g., QCAZ 43709-10, 43715, 43800).

Advertisement call.

A call from a male (QCAZ 40082) at Comunidad Santa Rosa (Napo Provincia) recorded on 28 February 2009 (Table 7) consists of three to four notes that sound like a high-pitched chuckle. The mean call duration is 0.35 s (SD = 0.04) and the mean rise time is 0.19s (SD = 0.1). The dominant frequency is higher (mean = 2217.93 Hz, SD = 56.94) and the fundamental frequency lower (mean = 488.10 Hz, SD = 12.47) compared to Hypsiboas fasciatus , Hypsiboas almendarizae , Hypsiboas calcaratus , Hypsiboas alfaroi sp. n. and Hypsiboas tetete sp. n.

Distribution and ecology.

Hypsiboas maculateralis inhabits the Amazon basin of Ecuador (Napo, Orellana, Pastaza, and Sucumbíos provinces) and Peru ( Región de Madre de Dios) (Fig. 17). Localities with known elevation range vary between 186 and 354 m of elevation. The elevation of Comunidad Santa Rosa (354 m) is the highest known and Santa Teresita (186 m) is the lowest.

All the specimens in Zábalo, Reserva Cuyabeno, Santa Teresita, Bataburo Lodge, and km 74 Pompeya-Iro road were found at night, in primary or secondary forests, perching on vegetation between 40 and 200 cm above the ground, in areas of pasture and flooded forest.

Vegetation types for Ecuadorian localities are: (1) Amazonian Lowland Evergreen Forest, characterized by high plant alpha-diversity and a canopy of 30 m with emergent trees that reach 40 m, (2) Floodplain Lowland Forest of White-Waters, characterized by periodical flooding with white-waters from large rivers, the vegetation reaches 35 m of height, and there are several horizontal strata of vegetation, and (3) Lowland Forest of Palms and black-waters, swamps characterized by a canopy of 30 m with dens e understory and a dominance of the palm Mauritia flexuosa .

Vegetation type of the Peruvian locality is Southwest Amazon Moist Forest.

Conservation status.

The distribution polygon of Hypsiboas maculateralis has an area of 209,304 km2. Because its distribution range is large and includes extensive areas of undisturbed forest, we propose that Hypsiboas maculateralis be assigned to the category Least Concern.