CaminerMarcel A.RonSantiago R.Systematics of treefrogs of the Hypsiboas calcaratus and Hypsiboas fasciatus species complex (Anura, Hylidae) with the description of four new speciesZookeys1512014201437016810.3897/zookeys.370.6291 Hypsiboas tetete http://zoobank.org/D9448B7C-4778-4912-A21A-3E32E4266EF1 http://species-id.net/wiki/Hypsiboas_tetete sp. n.Common name.

English: Tetete’s treefrog; Spanish: Rana arbórea de los Tetetes

Holotype.

(Fig. 14G) QCAZ 40081 (field no. SC-PUCE 23220), adult male from Ecuador, Provincia Napo, Comunidad Santa Rosa, road to Tena (1.0214°S, 77.4782°W), 344 m above sea level, collected by Marcel A. Caminer and Edwin Carrillo Ponce on 28 February 2009.

Paratopotypes.

QCAZ 40060–61, 40079–80, adult males, collected with the holotype.

Paratypes.

ECUADOR: PROVINCIA NAPO: Jatun Sacha (1.0649°S, 77.6142°W), 420 m, QCAZ 48094, adult male, 48095–96, adult females, collected by S. R. Ron and Morley Read on 31 March 2010.

Referred specimens.

PERU: REGIÓN LORETO: San Jacinto (2.3125°S, 75.8628°W), 180 m (KU 221864).

Diagnosis.

Hypsiboas tetete (Figs 8F and 19) is characterized by: (1) mean SVL 31.72 mm in males (range 31.15–32.24; n = 5), 45.59 mm in females (range 45.33–45.85; n = 2); (2) basal webbing on the fingers; (3) tubercle on the heel; (4) dorsal background coloration varying from grayish brown to pale brown, sometimes with dark markings (e.g., diffuse broad transversal bands); (5) faint brown middorsal line often present; (6) flanks creamy white or gray (in life, light blue in large females and white in males) with dark brown irregular spots; (7) hidden surfaces of thighs creamy white or brown with dark brown irregular spots or blotches; (8) ventral surfaces of thighs creamy white or yellowish white; (9) ventral areas creamy white or yellowish white with brown flecks on the head, neck, and chest; (10) webbing on the feet; (11) in life, iris yellow or cream with yellow on its anterior half; (12) prepollical spine present in males.

Hypsiboas tetete differs from Hypsiboas fasciatus, Hypsiboas almendarizae, Hypsiboas calcaratus, and Hypsiboas maculateralis in advertisement call (Figs 12A–F and 13A–H) and by having a tubercle on the heel instead of a calcar. Hypsiboas tetete is most similar to Hypsiboas alfaroi from which differs in advertisement call (Fig. 13C–H). Although with overlapping values, Hypsiboas tetete has a statistically significant larger tympanum than Hypsiboas alfaroi (see Hypsiboas alfaroi diagnosis). Morphological characters useful to differentiate Hypsiboas tetete from other species are shown in Table 6.

Description of holotype.

Adult male, SVL 31.15 mm, foot length 12.01 mm, head length 7.48 mm, head width 10.31 mm, eye diameter 3.51 mm, tympanum diameter 2.25 mm, tibia length 17.76 mm, femur length 16.09 mm, arm length 5.50 mm, eye-nostril distance 1.97 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 1.6 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. 1.03 times its diameter; posterior end of supratympanic fold reaches anterior border of arm insertion. Arm slender, axillary membrane absent; indistinct low tubercles present 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. Small tubercle 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, elongated and elliptical; subarticular tubercles single, round, flat; supernumerary tubercles restricted to the soles; webbing formula of toes I2-—2½II2—3+III2—3+IV3+—2-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; short simple cloacal sheath covering cloacal opening; round tubercles below and on the sides of vent. Tongue ovoid, widely attached to mouth floor; vomerine odontophores triangular with arched base, barely separated, posteromedial to choanae, bearing eight vomerine teeth on each side; choanae ovoid.

Color of holotype in preservative. Dorsum grayish brown with scattered minute black dots; faint brown narrow middorsal line extends from the tip of the snout to the mid-sacrum; dorsal surfaces of hindlimbs and forelimbs grayish brown with narrow transversal brown bars (one or two on each forearm and three or four on each thigh, shank, and tarsus); flanks creamy white with dark brown irregular spots; hidden surfaces of thighs grayish brown with dark brown irregular spots; venter creamy white with brown flecks on the neck, chest, and lips; ventral surfaces of hindlimbs and forelimbs yellowish white with a narrow to wide brown stripe on the outer edge of the hands, forearms, thighs, tarsal folds, and feet; bones white.

Color of holotype in life. (Fig. 14G). Dorsum pale yellowish tan with four narrow pale brown longitudinal lines; a dark brown middorsal line extends from the tip of snout to mid-sacrum; dorsal surfaces of hindlimbs yellowish tan with pale brown transversal bars; scattered minute dark brown dots on the dorsal surfaces of limbs and dorsum; flanks white with well-defined dark brown irregular spots; hidden surfaces of thighs pale yellowish tan with dark brown spots; venter creamy white with yellowish white belly; ventrally, scattered brown flecks on the chest, gular region, and jaw margin; ventral surfaces of hindlimbs and forelimbs creamy white; discs and webbing pale yellow tan; iris cream with faint yellow coloration on its upper half; bones white.

Etymology.

The specific name is a noun and refers to the Tetete, a Western Tucanoan indigenous group that inhabited the Colombian and Ecuadorian Amazon. It was decimated by the rubber exploitation and became extinct during the 1970s (Wasserstrom et al. 2011). Its recent disappearance parallels the destruction of increasingly large areas of forest in the Ecuadorian Amazon with the ensuing decline of biodiversity.

Variation.

Variation in dorsal and ventral coloration of preserved specimens is shown in Figure 19. Background dorsal coloration varies from grayish brown (e.g., QCAZ 48094) to pale grayish brown (e.g., QCAZ 48096) or pale brown (e.g., QCAZ 48094). Irregular dorsal marks may be present in diverse patterns. A faint middorsal line extends from the tip of snout to mid-dorsum (e.g., QCAZ 40079), mid-sacrum (e.g., QCAZ 40060) or to the cloaca (e.g., QCAZ 48095–96). There is variation in the number, size and shape of the dorsum marks. Five to seven broad transversal bands (sometimes interconnected) may be present on the dorsum; the dorsal surfaces of the limbs have brown transversal bars (one or two each on upper arm and forearm and three to five each on thigh, shank, and foot) (e.g., QCAZ 40080). Some individuals have scattered black or white dots on the dorsum (e.g., QCAZ 40060). The coloration of flanks varies from creamy white to gray with conspicuous dark brown irregular spots distributed from the groin to mid-flank. In some individuals, similar spots can also be present on the hidden surfaces of shanks and dorsal surfaces of feet (e.g., QCAZ 48096). The coloration of the hidden surfaces of thighs varies from creamy white to brown, with dark brown spots (e.g., QCAZ 40060) or blotches (QCAZ 48096).

Ventral areas of preserved specimens vary from creamy white (e.g., QCAZ 48094, 48096) to yellowish white (e.g., QCAZ 40079) with scattered flecks on the head and chest. In some individuals, the flecks are also present on hindlimbs, forelimbs, and belly (e.g., QCAZ 48095). Some individuals (e.g., QCAZ 48096) have a narrow to wide brown stripe on the outer edge of the hands, forearms, thighs and tarsal folds. Coloration of webbing and discs vary from brown to gray. Vomerine odontophores are triangular (with arched base in some specimens). Bones white.

In the adult type series, the largest male has a SVL of 32.24 mm, and the largest female 45.85 mm; mean male SVL = 31.72 mm (n = 5; SD = 0.42), female SVL range is 45.33 to 45.85 mm (n = 2). Inter-population variation in size and other morphometric variables are shown in Tables 1 and 2.

Coloration in life.

(based on photographs; Fig. 8F). Dorsal surfaces vary from pale yellowish tan (e.g., QCAZ 40060) to reddish brown (e.g., QCAZ 40079), with a brown middorsal line (e.g., QCAZ 48096); narrow pale brown longitudinal lines (e.g., QCAZ 40060, 48094) may be present. Some individuals have pale brown transversal bars on the dorsal surfaces of hindlimbs (e.g., QCAZ 40080); scattered minute black dots can be present on the dorsal surfaces of limbs and dorsum (e.g., QCAZ 40060); flanks are white (light blue in large females) with dark brown irregular marks with rounded (e.g., QCAZ 40060) or elongated shapes (e.g., QCAZ 48096); hidden surfaces of thighs are white or gray with dark brown spots (e.g., QCAZ 40060); in some specimens, the hidden surfaces of the shanks and dorsal surfaces of feet also have dark brown irregular spots (e.g., QCAZ 48096); venter creamy (e.g., QCAZ 48096) or yellowish white (e.g., QCAZ 40080) with scattered brown flecks on the ventral surfaces of the head and chest (e.g., QCAZ 48094); ventral surfaces of hindlimbs and forelimbs creamy white (e.g., QCAZ 40060) or yellowish white (e.g., QCAZ 40061); discs and webbing pale yellow tan; iris bronze (e.g., QCAZ 48096) or cream with faint yellow coloration on its upper half (e.g., QCAZ 40060); bones are white (e.g., QCAZ 40080).

Calls.

We recorded the calls of four malesat Comunidad Santa Rosa (Provincia Napo) on 28 February 2009, in flooded areas of secondary forest. Call parameters are shown in Table 7. Two call types were recorded. Type one (Fig. 13E–F) was the most common and consisted of a beep-like note with a mean duration of 0.10 s (SD = 0.02), average rise time 0.03 s (SD = 0.02) and average dominant frequency 1938.47 Hz (SD = 26.24). Type two (Fig. 13G–H) consisted of a single pulsed note with a mean duration of 0.11 s (SD = 0.02), mean rise time 0.05 s (SD = 0.02), and mean dominant frequency 1829.12 Hz (SD = 12.61). Call type two was alternated between calls of type one. Only males QCAZ 40060, 40080–81 produced this type of call.

Distribution and ecology.

Hypsiboas tetete is distributed in the Ecuadorian (Provincia Napo) and Peruvian Amazon basin (Región Loreto) (Fig. 15). Known localities range in elevation from 180 m (San Jacinto) to 420 m (Jatun Sacha). It is likely to have a larger distribution. Unfortunately, the lack of distinctive morphological characters relative to Hypsiboas alfaroi, preclude the unequivocal identification of museum specimens not associated with advertisement calls or genetic data. All specimens from Comunidad Santa Rosa and Jatun Sacha were found in flooded areas, in secondary forest, roosting on vegetation, 50 to 80 cm above ground.

Vegetation type for the Ecuadorian localities is Amazonian Lowland Evergreen Forest characterized by high plant alpha-diversity and a canopy of 30 m with emergent trees that reach 40 m.

Vegetation type at the Peruvian locality is Napo Moist Forest.

Conservation status.

The distribution polygon has 2,950 km2 of which 106 (3.5%) have been degraded by human activities (estimated from Ministerio de Ambiente del Ecuador 2013). Because its known distribution range is small with less than five localities and habitat degradation is increasing, Hypsiboas tetete is assigned to the Endangered category under criteria B1ab(iii).

Variation in color of the iris of A Hypsiboas fasciatus (QCAZ 47051), and B Hypsiboas almendarizae (QCAZ 42055). Calcar in C Hypsiboas almendarizae (QCAZ 39639) and D Hypsiboas calcaratus (QCAZ 40055). Note that the calcar in Hypsiboas almendarizae is large and conical while in Hypsiboas calcaratus is large and triangular. Dorsolateral views of the holotypes of E Hypsiboas almendarizae, adult male (QCAZ 39650), SVL = 36.72 mm F Hypsiboas maculateralis, adult male (QCAZ 40082), SVL = 36.16 mm; and G Hypsiboas tetete, adult male (QCAZ 40081), SVL = 31.15 mm.

Dorsolateral views of adult males of A Hypsiboas fasciatus, QCAZ 48584, SVL = 33.77 mm B Hypsiboas almendarizae, QCAZ 39649, SVL = 36.54 mm C Hypsiboas calcaratus, QCAZ 43256, SVL = 40.07 mm D Hypsiboas maculateralis, QCAZ 40082, SVL = 36.16 mm E Hypsiboas alfaroi, QCAZ 43260, SVL = 30.35 mm F Hypsiboas tetete, QCAZ 40081, SVL = 31.15 mm.

Adult Hypsiboas tetete showing variation in dorsal and ventral coloration of preserved specimens. From left to right, first and second rows: QCAZ 48095–96 (females), 40060 (male); third and fourth rows: QCAZ 40079, 40081 (holotype), 48094 (males). See Appendix for locality data. All specimens are shown at the same scale.

AppendixExamined specimens

Hyla leptoscelis. BRAZIL: AMAZONAS: Lago do Tachy, above São Paulo Olivença, Rio Solimöes (BMNH 1947.2.23.10, holotype).

Hyla steinbachi. BOLIVIA: PROVINCIA SARA: Departamento Santa Cruz de la Sierra (BMNH 1947.2.13.61–63, syntypes).

Hypsiboas calcaratus. ECUADOR: PROVINCIA ORELLANA: Río Napo, San Vicente (0.6790°S, 75.6511°W), 203 m (QCAZ 44529–30); Río Napo, Chiroisla (0.5756°S, 75.8998°W), 203 m (QCAZ 44422); Río Napo, Añangu (0.5249°S, 76.3844°W), 255 m (QCAZ 43933–36, 43979); Río Napo, Edén (0.4983°S, 76.0711°W), 216 m (QCAZ 44176–79, 44225, 44246–47, 44286); Estación Científica Tiputini, Universidad San Francisco de Quito (0.6387°S, 76.1492°W), 230 m (QCAZ 12343–44); Río Rumiyacu (0.89566°S, 75.94783°W), 250 m (QCAZ 20547–51); Parque Nacional Yasuní, between km 80 and km 75, on the road Pompeya-Iro (0.8401°S, 76.3024°W), 243 m (QCAZ 43057, 43060, 43062–63); Pompeya-Iro road, 38.8 km from Pompeya (0.6535°S, 76.4535°W), 237 m (QCAZ 8201); Parque Nacional Yasuní, km 9 Pompeya (0.4598°S, 76.5931°W), 253–271 m (QCAZ 43034–35, 43038, 43046–47); Estación Científica Yasuní, Universidad Católica del Ecuador (0.6713°S, 76.4005°W), 230–250 m (QCAZ 8210, 8812, 11912–13, 12386, 14815–17, 16792–97, 16798–99, 17825, 18264, 19202, 19204–05, 20290, 20305, 20837, 22488–89, 22561–66, 22866, 22988, 23038–39, 23065–69, 23848–54, 24214, 24282, 31445–46, 36869–78, 40985–86, 41005–06, 41015, 43242–48, 43256–59, 49205); Río Napo, sector La Primavera (0.4442°S, 76.7868°W), 244 m (QCAZ 43824); El Coca (0.4778°S, 76.9898°W), 257 m (QCAZ 43713, 43789); La Belleza, Comunidad Bocana del Suno (0.6922°S, 77.1353°W), 309 m (QCAZ 33522, 33524); Aguarico, confluence of the Yasuní and Jatuncocha rivers (0.9836°S, 75.4183°W), 200 m (EPN 5085–86); Aguarico, Ishpingo II oil well (1.0947°S, 75.6494°W), 178 m (EPN 3865); Loreto, San José Daguano (0.8255°S, 77.4347°W), 450 m (EPN 5674); San Luis de Armenia (0.4822°S, 77.0683°W), 300 m (EPN 11739); PROVINCIA SUCUMBÍOS: Cuyabeno, Caña de Canangüeno (0.3990°S, 75.8753°W), 222 m (QCAZ 11924); Reserva Faunística Cuyabeno (0.08498°N, 76.13444°W), 273 m (QCAZ 2034, 2046–49, 2242, 4612); Laguna de Mateococha (0.01846°N, 76.22155°W), 220 m (QCAZ 26062); Puerto Bolívar (0.0886°S, 76.1420°W), 240 m (QCAZ 28181, 28185, 28197); La Selva Lodge (0.4981°S, 76.3738°W), 245 m (QCAZ 4333, 25419, 25434); La Selva Lodge, Mandi Cocha (0.41666°S, 76.1333°W), 250 m (QCAZ 11540–41, 11545, 12005–07, 12009); Reserva Limoncocha (0.4062°S, 76.6194°W), 261 m (QCAZ 43100, 43131,43268); Shushufindi (0.0331°S, 76.6535°W), 270 m (QCAZ 15188); Zancudococha (QCAZ 4535); Tarapoa (QCAZ 23095); Cofán Duvuno community (0.0333°S, 77.1166°W), 340 m (EPN 4917); PROVINCIA NAPO: Reserva Yachana (0.8333°S, 77.1666°W), 300 m (QCAZ 48833); Comunidad Santa Rosa, on the road to Tena (0.9895°S, 77.4412°W), 341–439 m (QCAZ 40055–56, 40083); Tena, on the road to Jatun Sacha (1.0449°S, 77.76951°W), 428 m (QCAZ 18173, 40084–85); on the road Hollín-Loreto (QCAZ 649); La Selva (QCAZ 7436–40, 7443); PROVINCIA PASTAZA: Canelos Sacha (1.5822°S, 77.7155°W), 500–650 m (QCAZ 14956–75, 48712, 48718–20); Bataburo Lodge (1.2083°S, 76.7166°W), 220 m (QCAZ 39442); Kapawi Lodge (2.5386°S, 76.8583°W), 239 m (QCAZ 9018, 25513–14); Misión (2.2143°S, 76.5142°W), 240 m (EPN 885); Montalvo (2.0666°S, 76.9666°W), 305 m (EPN 884); Shionayacu (2.1001°S, 76.6334°W), 360 m (EPN 886); FRENCH GUIANA: Kaw (4.7166°N, 52.1333°W), 3 m (105PG); Cayenne, Crique Arataye (4°N, 52.66°W), 70 m (USNM 247780); Lac Toponow (3.0527°N, 52.7102°W), 152 m (123PG); Trinite (4.5833°N, 53.35°W), 290 m (192BM); Crique Margot (5.4666°N, 53.95°W), 44 m (131MC); GUYANA: Rupununi, Iwokrama Forest Reserve, 5 hrs S (downstream) of Kurupukari (4.285°N, 58.50944°W), 99 m (USNM 531370, 531372); BRAZIL: RONDONIA: Nova Brasilia (11.15°S, 61.566 °W), 431 m (USNM 304048); PERU: REGIÓN MADRE DE DIOS: Cuzco Amazónoico, 15 km E Puerto Maldonado (12.583°S, 69.083°W), 200 m (MUSM 14456, 14547, KU 215205); Río Tambopata, W bank of Zona Reservada Tambopata-Candamo (13.1416°S, 69.6066°W), 211 m (USNM 332415); Pakiza, Manu National Park (11.866°S, 71.3°W), 360 m (USNM 345216–17); REGIÓN LORETO: Río Lagarto Cocha, Aguas Negras (0.5230°S, 75.2569°W), 183 m (USNM 520924).

Hypsiboas fasciatus. ECUADOR: Andes of Ecuador (BMNH 58.4.25.22; holotype); PROVINCIA MORONA SANTIAGO: 7.9 km N Limón, on the road Limón-Macas (2.8858°S, 78.3970°W), 1571 m (QCAZ 26496–97); Comunidad Nueva Principal, 3 km near to the town (3.1686°S, 78.3870°W), 1059 m (QCAZ 48611, 48628); Tiink, Río Zamora (3.3330°S, 78.4518°W), 730 m (QCAZ 17016, 17123, 17128, 17030–31, 17040–41, 17148); Comunidad San Luis (3.3420°S, 78.4677°W), 792 m (QCAZ 48633, 48636, 48639, 48670–71); La Pradera, on the road to Gualaquiza (3.3365°S, 78.6504°W), 1015–1036 m (QCAZ 18270–71, 48584–93); road to Río Abanico Jimbitano, 1000 m (QCAZ 21188–89); road between Gualaquiza and Limón, 3 km from Limón (3.3990°S, 78.5786°W), 1140 m (QCAZ 27610); San Juan Bosco, Cóndor Mirador (3.5192°S, 78.4314°W), 1100 m (EPN 14060–65); 3 km Gualaquiza-Limón (3.3700°S, 78.5680°W), 1140 m (QCAZ 21707, 21957, 33976); PROVINCIA ZAMORA-CHINCHIPE: Gravel road East to Sarsa (3.8078°S, 78.6059°W), 1500 (QCAZ 47051); Piuntza, Finca Don Mesías (3.8564°S, 78.8646°W), 1154–1192 m (QCAZ 40797, 40802); near to Zamora (4.0491°S, 78.9256°W), 927 m (QCAZ 27258); Zamora (4.0669°S, 78.9508°W), 1060 m (QCAZ 23144, 23147–49, 48583); Bombuscaro (4.0893°S, 78.9609°W), 1091 m (QCAZ 12444, 12446, 12448, 12452, 12494–97); La Pituca (4.1461°S, 78.9446°W), 1343 m (QCAZ 24650); Romerillos Alto, 26 km of Zamora (4.1850°S, 78.9352°W), 1300–1593 m (QCAZ 24866, 47070); Las Orquidias, around the town (4.2290°S, 78.6577°W), 874 m (QCAZ 41488); Miazi Alto, “Tepuy” camp (4.2562°S, 78.6222°W), 1250–1300 m (QCAZ 41575–76, 41659, 41518); Destacamento Militar Miasi (4.2833°S, 78.6333°W), 800 m (EPN 4112–13, 4115–16, 4118); Paquisha, Blanco river (3.8945°S, 78.5161°W), 1630 m (EPN 14218); Yantzaza, Colibrí Goldmarca (3.7665°S, 78.5055°W), 1377 m (EPN 12672, 12674, 13288, 13290–92); Yantzaza, Concesión Colibrí (3.7588°S, 78.5009°W), 1424 m (EPN 11387–91, 12316); Yantzaza, Sachavaca sector (3.7042°S, 78.4827°W), 1280 m (EPN 14216–17); Yantzaza, road to Pindal (3.7522°S, 78.5528°W), 1224 m (EPN 13677); Centro Shuar Yawi (4.4301°S, 78.6489°W), 945 m (QCAZ 31037–40); near to Tundaime (QCAZ 16466); PERU: REGIÓN AMAZONAS: Cordillera del Cóndor, Upper Río Comainas, Puesto Vigilancia (4.1°S, 78.3833°W), 1100 m (USNM 525495, 525499).

Calls of the Hypsiboas calcaratus species complex. A–B Hypsiboas fasciatus (QCAZ 48585) from La Pradera, Provincia Morona Santiago C–D Hypsiboas almendarizae (QCAZ 39650) from Limón, Provincia Morona Santiago E–F Hypsiboas calcaratus (QCAZ 40085) from Tena, Provincia Napo. A, C, E are oscilograms and B, D, F spectrograms.

Calls of the Hypsiboas calcaratus species complex. A–B Hypsiboas maculateralis (QCAZ 40082) from Comunidad Santa Rosa, Provincia Napo C–D Hypsiboas alfaroi (QCAZ 43260) from Estación Científica Yasuní PUCE, Provincia Orellana E–H Hypsiboas tetete (QCAZ 40081) from Comunidad Santa Rosa, Provincia Napo. A, C, E, G are oscilograms and B, D, F, H spectrograms.

Diagnostic characters of male specimens of the Hypsiboas calcaratus species complex. Coloration corresponds to preserved specimens unless otherwise noted.

CharactersHypsiboas alfaroiHypsiboas almendarizaeHypsiboas calcaratusHypsiboas fasciatusHypsiboas maculateralisHypsiboas tetete
Flank ColorationCreamy white or gray with dark brown irregular spotsPale cream or creamy white with thin dark brown vertical barsPale cream, creamy white or light gray with dark brown vertical barsPale cream, creamy white, brown or gray with dark brown vertical barsPale cream, creamy white or gray with dark brown blotchesCreamy white or gray with dark brown irregular spots
Coloration of the hidden surfaces of thighsCreamy white, gray or brown with dark brown irregular spotsPale cream or creamy white with thin dark brown transversal barsPale cream, creamy white or light gray with dark brown transversal barsPale cream, creamy white, brown or gray with dark brown transversal barsPale cream, creamy white or gray with dark brown blotchesCreamy white or brown with dark brown irregular spots
Size and shape of the calcarCalcar absent. Only small tubercle presentLarge and conicalLarge and triangularSmall and conicalLarge and triangularCalcar absent. Only small tubercle present
Brown flecks on the neck and chestPresentAbsentAbsentAbsentSometimes presentPresent
Iris upper band, in lifeIll-defined, yellowWell-defined, yellow to orangeWell-defined, yellow to orangeWell-defined, red to reddish brownIll-defined, yellowIll-defined, yellow

Descriptive statistics for morphometric measurements of male Hypsiboas alfaroi, Hypsiboas almendarizae, Hypsiboas calcaratus, Hypsiboas fasciatus, Hypsiboas maculateralis, and Hypsiboas tetete used for Principal Component Analysis. Mean ± SD is given with range below. Bold figures represent combined values for males of all populations. Abbreviations are: SVL = Snout-vent length; FOOT = Foot length; HL = Head length; HW = Head width; ED = Eye diameter; TD = Tympanum diameter; TL = Tibia length; FL = Femur length; CL = Calcar length. All measurements are in mm.

SpeciesSVLFOOTHLHWEDTDTLFLCL
Hypsiboas alfaroi(n = 32)32.80 ± 1.9727.91–36.2712.54 ± 0.6511.33–13.808.95 ± 0.987.13–11.8410.07 ± 0.618.72–10.953.25 ± 0.362.52–4.032.28 ± 0.281.70–3.0118.50 ± 0.9516.54–20.5016.25 ± 0.9614.71–17.91Calcar absent
Yasuní PUCE(n = 7)32.20 ± 1.9827.91–35.1912.73 ± 0.6111.33–13.468.35 ± 0.607.40–8.9910.10 ± 0.538.79–10.803.29 ± 0.362.90–4.032.24 ± 0.112.08–2.5018.66 ± 1.1016.54–20.5016.25 ± 0.9814.81–17.89Calcar absent
Huiririma(n = 2)33.61–34.2212.95–13.809.69–9.7510.37–10.773.25–3.322.22–2.5118.10–19.3215.50–17.17Calcar absent
Nuevo Rocafuerte(n = 5)33.99 ± 2.1330.34–35.8812.51 ± 0.8111.41–13.719.91 ± 1.168.98–11.8410.23 ± 0.639.26–10.953.11 ± 0.412.52–3.612.05 ± 0.231.70–2.2818.82 ± 1.2316.95–20.0516.96 ± 1.2214.92–17.91Calcar absent
Pañacocha(n = 3)33.04 ± 0.5232.64–33.6312.10 ± 0.1911.95–12.319.58 ± 0.109.46–9.649.65 ± 0.688.87–10.143.34 ± 0.332.97–3.622.29 ± 0.052.25–2.3417.97 ± 0.2917.66–18.2216.27 ± 0.2016.04–16.41Calcar absent
Selva Lodge(n = 2)32.11–33.8512.72–13.148.58–9.069.18–10.532.73–3.612.11–2.9118.47–18.9014.71–16.21Calcar absent
Hypsiboas almendarizae(n = 23)37.64 ± 2.0134.31–44.5615.04 ± 0.8212.99–16.009.74 ± 0.868.33–11.3511.8 1 ± 0.6010.80–13.074.02 ± 0.343.36–4.562.27 ± 0.241.63–2.7421.27 ± 0.9419.53–23.0419.24 ± 1.0615.84–21.521.51 ± 0.241.09–1.88
Baños(n = 2)37.39–37.9813.43–15.169.51–9.6911.88–12.004.22–4.322.31–2.3019.53–21.2118.48–19.381.32–1-39
El Rosario(n = 3)38.90 ± 1.7937.56–40.9215.29 ± 0.4014.89–15.7011.06 ± 0.4610.52–11.3412.40 ± 0.2712.15–12.684.42 ± 0.124.35–4.562.30 ± 0.202.13–2.5322.45 ± 0.6621.74–22.5819.37 ± 0.2319.10–19.501.64 ± 0.071.56–1.68
Limón Indanza(n = 12)36.72 ± 1.2334.31–38.6014.97 ± 0.8812.99–16.009.19 ± 0.398.33–9.7411.73 ± 0.5610.80–12.683.94 ± 0.343.36–4.542.25 ± 0.232.02–2.7421.01 ± 0.7319.64–22.2419.13 ± 1.1915.84–20.231.51 ± 0.281.09–1.88
Río Hollín(n = 2)38.46–44.5615.73–15.7510.20–11.3511.72–13.073.60–4.381.63–2.2521.80–22.6218.83–21.521.33–1.57
Hypsiboas calcaratus(n = 35)36.82 ± 2.5927.61–42.5015.00 ± 1.3110.68–17.4410.14 ± 1.067.71–12.6311.93 ± 1.019.16–13.723.63 ± 0.372.47–4.522.48 ± 0.211.92–2.9422.10 ± 1.5716.76–24.6119.03 ± 1.4813.82–21.442.03 ± 0.331.32–2.65
Canelos(n = 3)36.59 ± 1.4934.89–37.6914.64 ± 0.5914.08–15.2610.48 ± 0.5510.05–11.1011.68 ± 0.5911.01–12.133.87 ± 0.303.69–4.222.43 ± 0.242.18–2.6621.78 ± 0.3421.39–22.0318.28 ± 0.5317.67–18.662.06 ± 0.251.89–2.34
Tena(n = 5)35.23 ± 2.8032.10–39.1513.87 ± 1.1012.65–15.209.12 ± 1.158.18–10.6911.95 ± 1.2410.69–13.723.71 ± 0.313.33–4.042.57 ± 0.232.39–2.9420.69 ± 1.2918.77–21.9818.28 ± 1.1316.51–19.432.05 ± 0.391.56–2.54
Edén(n = 2)36.88–37.0914.39–14.579.75–10.7211.30–12.243.29–3.552.41–2.7120.70–21.8517.42–17.631.84–2.13
El Coca(n = 2)34.83–34.9614.21–14.429.05–9.1611.19–11.312.47–2.942.10–2.3720.04–22.5216.46–19.221.68–1.98
Estación Científica Yasuní PUCE(n = 18)37.78 ± 1.5335.21–40.1015.73 ± 0.8914.54–17.4410.43 ± 0.898.99–12.6312.20 ± 0.909.59–13.333.71 ± 0.303.36–4.522.48 ± 0.172.23–2.8722.99 ± 0.9821.12–24.6119.91 ± 0.8518.40–21.442.06 ± 0.361.32–2.47
Parque Nacional Yasuní, Pompeya(n = 2)36.11–36.3113.50–15.5210.42–11.4311.02–13.133.60–4.122.43–2.6321.14–23.4719.38–19.551.90–2.04
Hypsiboas fasciatus(n = 19)35.40 ± 1.6532.65–37.7414.00 ± 0.5912.85–15.249.651 ± 0.668.42–11.5311.39 ± 0.5910.41–12.333.71 ± 0.343.15–4.422.42 ± 0.252.03–2.7820.07 ± 0.8918.63–21.7118.18 ± 0.8716.12–19.941.35 ± 0.151.08–1.63
Centro Shuar Yawi(n = 4)35.79 ± 2.1436.22–37.7213.81 ± 1.0212.85–15.249.69 ± 0.579.03–10.4311.60 ± 0.7110.66–12.333.91 ± 0.383.50–4.422.40 ± 0.252.20–2.7320.55 ± 1.3418.63–21.7118.14 ± 1.4216.12–19.421.35 ± 0.071.25–1.41
La Pradera(n = 4)34.69 ± 0.8233.74–35.7414.18 ± 0.1414.00–14.229.61 ± 0.319.36–10.0610.98 ±0.4310.41–11.443.66 ± 0.163.56–3.822.31 ± 0.332.03–2.7819.27 ± 0.4319.06–19.8318.00 ± 0.5317.30–18.441.50 ± 0.101.39–1.63
Tiink(n = 6)34.79 ± 1.8932.65–37.5013.96 ± 0.5213.46–14.779.72 ± 1.068.42–11.5311.17 ± 0.6310.60–12.103.55 ± 0.423.15–4.092.50 ± 0.192.26–2.7220.30 ± 0.8019.25–21.1218.44 ± 1.0017.29–19.941.23 ± 0.161.08–1.47
Zamora(n = 2)34.55–36.9513.52–13.588.94–9.5511.54–11.853.85–3.952.51–2.7819.21–19.6117.71–18.041.25–1.40
Hypsiboas maculateralis(n = 22)36.00 ± 1.9231.86–39.1713.55 ± 1.0311.10–16.229.59 ± 1.186.62–11.1911.08 ± 0.709.21–12.293.55 ± 0.492.63–4.392.17 ± 0.281.26–2.6320.67 ± 1.4816.94–23.2318.18 ± 1.4015.31–20.571.67 ± 0.321.15–2.52
Bataburo Lodge(n = 2)34.20–38.1813.02–13.909.58–11.1010.27–11.813.36–4.242.09–2.1418.60–22.2117.77–19.191.24–1.47
Edén(n = 3)34.74 ± 2.6831.86–37.1712.65 ± 1.3711.10–13.687.93 ± 1.346.62–9.3010.38 ± 1.029.21–11.093.29 ± 0.632.63–3.882.12 ± 0.161.94–2.2418.83 ± 1.6416.94–19.8216.85± 1.3515.31–17.841.81 ± 0.221.56–1.96
Reserva de Producción Faunística Cuyabeno PUCE(n = 7)36.40 ± 1.5333.71–38.6913.59 ± 0.6012.57–14.1910.23 ± 0.759.15–11.1911.32 ± 0.4510.69–12.143.97 ± 0.263.67–4.392.29 ± 0.152.10–2.5721.65 ± 0.6920.63–22.4319.56 ± 0.8318.36–20.571.89 ± 0.351.55–2.52
La Primavera(n = 2)35.26–38.4913.41–14.589.36–11.0110.90–11.143.02–3.671.26–2.2820.84–20.9117.20–17.691.27–1.86
Selva Lodge(n = 2)37.48–39.1712.98–16.228.77–9.9411.72–12.293.24–3.332.43–2.6320.86–23.2316.12–19.901.61–1.90
Hypsiboas teteteComunidad Santa Rosa(n = 5)31.72 ± 0.4231.15–32.2412.22 ± 0.1512.01–12.408.13 ± 0.537.48–8.7510.32 ± 0.249.97–10.643.60 ± 0.253.38–4.022.62 ± 0.352.25–3.0117.62 ± 0.2417.30–17.9316.50 ± 0.3616.09–17.00Calcar absent

Descriptive statistics for morphometric measurements of female Hypsiboas alfaroi, Hypsiboas almendarizae, Hypsiboas calcaratus, Hypsiboas fasciatus, Hypsiboas maculateralis, and Hypsiboas tetete used for Principal Component Analysis. Mean ± SD is given with range below. Bold figures represent combined values for females from all populations. Abbreviations are: SVL = Snout-vent length; FOOT = Foot length; HL = Head length; HW = Head width; ED = Eye diameter; TD= Tympanum diameter; TL = Tibia length; FL = Femur length; CL = Calcar length. All measurements are in mm.

SpeciesSVLFOOTHLHWEDTDTLFLCL
Hypsiboas alfaroi(n = 12)44.51 ± 3.0939.68–49.2116.72 ± 1.9611.39–18.9411.59 ± 1.178.90–13.0914.21 ± 1.331.55–15.764.12 ± 0.563.23–5.002.96 ± 0.322.22–3.4425.47 ± 1.9322.88–9.4422.77 ± 1.6419.88–25.66Calcar absent
Estación Científica Yasuní PUCE(n = 4)42.81 ± 2.0840.27–45.3715.82 ± 2.9911.39–17.7111.71 ± 0.6611.03–12.2813.93 ± 1.0512.41–14.704.12 ± 0.363.59–4.342.87 ± 0.232.60–3.1424.83 ± 1.5223.18–26.8022.91 ± 1.3221.17–24.35Calcar absent
Playas de Cuyabeno(n = 2)44.51–48.0116.60–17.4112.15–13.0914.99–15.714.91–5.003.18–3.4424.61–25.1121.16–23.38Calcar absent
Hypsiboas almendarizae(n = 4)48.11 ± 6.8837.80–51.9419.36 ± 2.9215.11–21.6411.59 ± 1.968.70–12.9914.77 ± 2.4511.15–16.594.08 ± 0.663.45–4.682.52 ± 0.412.07–3.0025.81 ± 6.8815.51–29.7424.46 ± 3.4119.40–26.621.89 ± 0.551.20–2.53
Nueve de Octubre(n = 3)51.54 ± 0.3551.26–51.9420.78 ± 0.8819.87–21.6412.55 ± 0.4312.14–12.9915.98 ± 0.5415.61–16.594.28 ± 0.623.57–4.682.68 ± 0.342.32–3.0029.25 ± 0.4428.87–29.7426.15 ± 0.6025.47–26.622.12 ± 0.361.87–2.53
Hypsiboas calcaratus(n = 4)50.92 ± 4.8045.94–56.2920.56 ± 2.0518.44–23.1713.08 ± 2.0311.04–15.8316.46 ± 1.8514.59–18.424.14 ± 0.863.57–5.423.12 ± 0.502.67–3.7731.00 ± 3.1328.10–35.2926.05 ± 2.1524.47–29.092.62 ± 0.162.42–2.78
Hypsiboas fasciatus(n = 5)51.89 ± 3.1847.16–54.8420.44 ± 1.2318.79–21.9813.91 ± 0.7912.59–14.5316.57 ± 0.7215.80–17.384.83 ± 0.434.28–5.323.25 ± 0.382.70–3.7729.58 ± 1.4628.55–32.0927.02 ± 1.3425.74–29.201.95 ± 0.151.73–2.09
Hypsiboas maculateralis(n = 6)45.18 ± 9.3932.04–55.3116.27 ± 3.2911.25–19.7711.76 ± 2.408.28–14.1014.21 ± 2.9510.01–17.373.78 ± 0.263.34–4.062.43 ± 0.521.70–3.0025.23 ± 4.4218.04–29.6621.90 ± 4.2215.48–26.541.61 ± 0.401.09–2.22
Zábalo(n = 2)32.04–38.2611.25–14.588.28–9.2810.01–12.383.34–3.611.70–2.0318.04–22.8215.48–19.151.09–1.39
Hypsiboas teteteJatun Sacha(n = 2)45.33–45.8516.81–18.1711.15–12.9613.66–14.114.09–4.963.56–3.8525.45–25.7821.18–21.81Calcar absent

Descriptive statistics for call parameters of Hypsiboas alfaroi (QCAZ 43260–63), Hypsiboas almendarizae (QCAZ 39645, 39647–50), Hypsiboas calcaratus (QCAZ 40084–85, 43247, 43256–59), Hypsiboas fasciatus (QCAZ 48583–86, 48633), Hypsiboas maculateralis (QCAZ 40082), and Hypsiboas tetete (QCAZ 40060, 40080–81, 48095). The n values indicate the number of males analyzed. Mean ± SD is given with range in parentheses. Values for Hypsiboas maculateralis were obtained from three calls from a single male. See Table 3 for a description of each parameter.

Hypsiboas alfaroi(n = 4)Hypsiboas almendarizi(n = 5)Hypsiboas calcaratus(n = 7)Hypsiboas fasciatus(n = 5)Hypsiboas maculateralis(n = 1)Hypsiboas tetete(n = 4)
Type 1Type 2
Call Duration (s)0.20 ± 0.05(0.16–0.19)0.48 ± 0.07(0.39–0.55)0.05 ± 0.00(0.05–0.06)0.52 ± 0.24(0.27–0.90)0.35 ± 0.04(0.317–0.398)0.10 ± 0.02(0.08–0.12)0.11 ± 0.02(0.09–0.13)
Number of notes4–53–413–53–411
Rise time (s)0.07 ± 0.03(0.05–0.08)0.30 ± 0.10(0.20–0.45)0.04 ± 0.01(0.03–0.05)0.24 ± 0.06(0.17–0.30)0.19 ± 0.10(0.084–0.289)0.03 ± 0.02(0.02–0.06)0.05 ± 0.02(0.03–0.07)
Call dominant frequency (Hz)2079.53 ± 83.43(1924.50–2032.74)1954.43 ± 128.43(1828.18–2115.65)1780.50 ± 112.73(1619.32–1927.25)1855.81 ± 148.08(1712.00–2088.70)2217.93 ± 56.94(2174.92–2282.51)1938.47 ± 26.24(1830.31–1959.23)1829.12 ± 12.61(1815.97–1841.10)
Call fundamental frequency (Hz)2036.31 ± 130.30(1951.05–2146.50)951.76 ± 61.38(882.86–1029.95)557.13 ± 46.21(512.60–651.38)884.89 ± 105.52(722.60–1001.30)488.10 ± 12.47(473.71–495.33)1940.14 ± 28.29(1830.31–1964.23)1821.96 ± 16.96(1808.80–1841.10)
Number of pulses14.85 ± 3.03(11.90–19.17)14.12 ± 0.74(12.60–14.75)11.80 ± 1.69(9.60–13.56)10.22 ± 1.68(8.67–12.00)
Pulse repetition rate (pulses/s)200.41 ± 31.89(168.95–249.22)268.46 ± 11.08(250.71–280.11)179.53 ± 18.08(150.31–195.95)92.97 ± 2.26(90.72–95.24)
Dominant frequency at the beginning to the third note (Hz)1869.09 ± 52.61(1808.80–1905.70)1827.51 ± 175.70(1722.70–2140.10)
Fundamental frequency at the beginning to the third note (Hz)826.16 ± 58.21(766.58–882.90)841.64 ± 91.67(728.30–969.00)
Dominant frequency at the end to the third note (Hz)2058.93 ± 33.98(2026.28–2094.10)1962.32 ± 190.41(1798.80–2282.50)
Fundamental frequency at the end to the third note (Hz)937.43 ± 83.53(882.90–1033.60)950.18 ± 144.22(709.50–1055.10
Interval between calls (s)7.75 ± 0.95(7–9)16.2 ± 6.37(10–15)12 ± 2.64(6–23)51 ± 24.72(13–78)152.5 ± 3.53(150–155)9 ± 4.24(6–15)
Recording temperatures (°C)25.921.523.9–25.620.1–23.621.522.2–2522.2–25

Call traits of Hypsiboas spp. analyzed in this study. See text for details.

CharacterDescription
Call durationTime from the beginning of the first note to the end of the last note of the call
Number of notesNumber of notes in the call
Rise timeTime from the beginning of the call to the point of its maximum amplitude
Call dominant frequencyThe frequency with the greatest amount of sound energy along all the call
Call fundamental frequencyThe frequency with the greatest amount of sound energy in the first harmonic, measured along all the call
Dominant frequency at the beginning to the third noteThe frequency with the greatest amount of sound energy measured at the beginning of the third note
Fundamental frequency at the beginning to the third noteThe frequency of the first harmonic measured at the beginning of the third note
Dominant frequency at the end to the third noteThe frequency with the greatest amount of sound energy measured at the end of the third note
Fundamental frequency at the end to the third noteThe frequency of the first harmonic measured at the end of the third note
Number of pulsesNumber of pulses per note
Pulse repetition rateNumber of pulses per second
Interval between callsTime from end of call to the beginning of next call

Distribution of species of the Hypsiboas calcaratus complex. Localities are based on museum specimens from Museo de Historia Natural Gustavo Orcés of Escuela Politécnica Nacional, Museo de Zoología of Pontificia Universidad Católica del Ecuador, Natural History Museum University of Kansas, and Smithsonian Institution National Museum of Natural History.

WasserstromRReiderSLaraR (2011) Nobody knew their names: the black legend of Tetete extermination.Ethnohistory58: 421-444. doi: 10.1215/00141801-1263848Ministerio de Ambiente del Ecuador (2013) Sistema de Clasificación de los Ecosistemas del Ecuador Continental.Subsecretaría de Patrimonio Natural, Quito, Ecuador.