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 fasciatus http://species-id.net/wiki/Hypsiboas_fasciatus (Günther, 1858)Hyla fasciata Gunther, 1858: 327. Holotype: BMNH 58.4.25.22, a female from “Anden von Ecuador” (Andes of Ecuador).Diagnosis.

A member of the genus Hypsiboas characterized by: (1) mean SVL 35.40 mm in males (range 32.65–37.74; n = 19), 51.89 mm in females (range 47.16–54.84; n = 5); (2) basal webbing on fingers; (3) calcar small and conical; (4) dorsal coloration varying from cream to grayish brown, pinkish white or brown, with dark marks in some individuals (e.g., broad transversal bands); (5) middorsal brown stripe often present; (6) flanks pale cream or gray (in life, blue in large females and light blue or white in males) with dark brown vertical bars; (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 transversal bars; (8) ventral surfaces of thighs creamy white, yellowish white or brown; (9) venter creamy white or yellowish white; (10) webbing on feet; (11) in life, iris creamy silver or bronze with upper red to reddish brown band; (12) prepollical spine present in males.

Hypsiboas fasciatus is most similar to Hypsiboas almendarizae sp. n. and Hypsiboas calcaratus. Hypsiboas calcaratus can be distinguished from Hypsiboas fasciatus by the shape of the calcar (large and triangular in Hypsiboas calcaratus vs. small and conical in Hypsiboas fasciatus), structure of the advertisement calls (Fig. 12A–B and 12E–F), and by the yellow to orange color of the upper band in the iris (red to reddish brown in Hypsiboas fasciatus). Hypsiboas almendarizae sp. n. differs from Hypsiboas fasciatus in having narrower transversal dark bars on the flanks and thighs (mean width of bars on thighs = 5.05% of femur length, SD = 0.61, in Hypsiboas almendarizae sp. n.vs. 8.58%, SD = 1.56, in Hypsiboas fasciatus). The color of the upper band in the iris also distinguishes both species (red to reddish brown in Hypsiboas fasciatus vs. yellow to orange in Hypsiboas almendarizae sp. n.; Fig. 14A–B).

Hypsiboas fasciatus differs from Hypsiboas maculateralis sp. n. in advertisement call (Figs 12A–B and 13A–B) and in the presence of transversal bars on the flanks and hidden surfaces of the thighs (dark blotches instead of bars in Hypsiboas maculateralis sp. n.) Hypsiboas fasciatus can easily 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) and by the absence of dark flecks on the gular region and chest (present in Hypsiboas alfaroi sp. n. and Hypsiboas tetete sp. n.) Morphological characters useful to differentiate Hypsiboas fasciatus from other species are shown in Table 6.

Variation.

Variation in dorsal and ventral coloration of preserved specimens is shown in Figure 5. Background dorsal coloration varies from cream (e.g., QCAZ 41488, 48584) to pale grayish brown (e.g., QCAZ 18271), grayish brown (e.g., QCAZ 27258), pinkish white (e.g., QCAZ 41575), pale brown (e.g., QCAZ 23148) or brown (e.g., QCAZ 26497). Irregular dorsal marks can be present in diverse patterns. A dark middorsal stripe extends from the tip of the snout to the mid-sacrum (e.g., QCAZ 26497) or to the vent (QCAZ 23144). In a few individuals, the middorsal line is absent (QCAZ 17123, 31040). In some individuals (e.g., QCAZ 41575) broad transversal bands are present on the dorsum; narrower brown transversal bars are present on the dorsal surfaces of the limbs (one or two on the upper arm and forearm and three to five on the thigh, shank, and foot). In some individuals, scattered minute brown dots may be present on dorsum (e.g., QCAZ 48584). The coloration of flanks and hidden surfaces of thighs vary from pale cream to creamy white, brown or gray, with dark brown vertical bars (4 to 12 on each flank and 6 to 12 on each thigh). The area with vertical bars extends from the groin to the axilla (e.g., QCAZ 41575) or to the mid flank (e.g., QCAZ 41576, 18271). In some individuals, similar transversal bars can be present on the hidden surfaces of the shanks, ventral surfaces of the forelimbs, and dorsal surfaces of the feet (e.g., QCAZ 41575). The shape of the calcar is small and conical and shows little variation among individuals.

Adult Hypsiboas fasciatus showing variation in dorsal and ventral coloration of preserved specimens. From left to right, first and second rows: QCAZ 24866, 26497, 41575 (females); third and fourth rows: QCAZ 18271, 41488, 41576, 48584 (males). See Appendix for locality data. All specimens are shown at the same scale.

Ventral areas vary from creamy white (e.g., QCAZ 48584) to yellowish white (e.g., QCAZ 41576, 41488). A narrow to wide brown stripe can be present on the outer edge of the hands, forearms, thighs, feet, and tarsal folds (e.g., QCAZ 48584, 41576, 41575, 26497). In some specimens, scattered minute pale brown blotches can be present on the lips (e.g., QCAZ 26497, 24866). Coloration of webbing and discs vary from yellowish white to brown or gray. Bones are white.

In the examined series of adults, the largest male has a SVL of 37.74 mm, and the largest female 54.84 mm; mean male SVL = 35.40 mm (n = 19; SD = 1.65), mean female SVL = 51.89 mm (n = 5; SD = 3.18). Females are significantly larger than males (t = -16.24, df = 22, P < 0.001). Inter-population variation in size and other morphometric variables is shown in Tables 1 and 2.

Coloration in life.

(based on digital photographs; Figs 4A and 8A). Dorsal surfaces vary from pale yellowish tan (e.g., QCAZ 47070) to brown (e.g., QCAZ 48611) with a middorsal dark brown line (e.g., QCAZ 48585) and scattered minute black dots (e.g., QCAZ 47051); some individuals have diffuse broad pale brown transversal bands on the dorsum (e.g., QCAZ 48583); pale brown transversal bars are present on the dorsal surfaces of the limbs; flanks and hidden surfaces of thighs are white, light blue or blue with dark brown vertical bars (e.g., QCAZ 47070, 47051); in some specimens, dark brown vertical bars are present on the hidden surfaces of the shanks, ventral surfaces of the upper arms and dorsal surfaces of the feet (e.g., QCAZ 48671); venter creamy white, sometimes with yellowish white on its posterior half (e.g., QCAZ 48670); ventral surfaces of hindlimbs and forelimbs translucent pinkish white (e.g., QCAZ 47051) or brown (e.g., QCAZ 48611); in some specimens, ventral surfaces of the thighs creamy white (e.g., QCAZ 47070); discs and webbing pale yellowish tan (e.g., QCAZ 48584) or brown (e.g., QCAZ 48611); a narrow to wide brown stripe is present on the outer edge of the hands, forearms, thighs, feet, and tarsal folds (e.g., QCAZ 48611); iris creamy silver (e.g., QCAZ 48584) or bronze (e.g., QCAZ 48611) with upper red to reddish brown band (e.g., QCAZ 48628); bones vary from green (e.g., QCAZ 48671) to white (e.g., QCAZ 48628).

Advertisement call.

We recorded calls of one male at Bombuscaro (Provincia Zamora Chinchipe) on 9 June 2010, three males at La Pradera (Provincia Morona Santiago) on 10 June 2010, and one male at Comunidad San Luis (Provincia Morona Santiago) on 13 June 2010. The advertisement call consists of three to five quack notes (Fig. 12A–B). Mean call duration is 0.52 s (SD = 0.24) and mean rise time is 0.24 s (SD = 0.06). Sound frequency increases from the beginning to the end of the note; mean dominant frequency is 1855.81 Hz (SD = 148.08) and mean fundamental frequency is 884.89 Hz (SD = 105.52). Other call parameters are listed in Table 7.

Distribution and ecology.

Hypsiboas fasciatus has been recorded in the Ecuadorian and Peruvian Amazon basin (Morona Santiago and Zamora-Chinchipe provinces in Ecuador and Región Amazonas in Peru) (Fig. 15). Localities with known elevation range from 730 to 1593 m above sea level. The elevation at Romerillos Alto (1593 m) is the highest known locality for Hypsiboas fasciatus, while Tink (730 m) is the lowest.

Specimens from Comunidad San Luis, La Pradera, La Pituca, Limón, Miazi Alto, Nueva Principal, and Tiink were found in primary and secondary forest, perching on vegetation 30 to 200 cm above the ground, in flooded areas, ponds, swamps, and near streams. Individuals in Zamora were found in grassy swamps roosting in shrubs between 30 and 110 cm above ground.

Vegetation types for Ecuadorian localities are: (1) Amazonian Evergreen Foothill Forest, characterized by a mixture of Amazonian and Andean vegetation with a canopy of 30 m, (2) Evergreen Lower Montane Forest of the Amazonian Range, characterized by trees reaching 20–30 m of height and abundant epiphytes and hemiepiphytes, with Dictyocaryum lamarckianum (Arecaceae) as dominant species, and (3) Evergreen Lower Montane Forest of the East of the Southern Andes, characterized by abundant epiphytes, trees reaching 30 m of height with Podocarpus as dominant species.

Conservation Status.

Its distribution polygon has 8,572 km2 of which 2,198 km2 (25.6%) have been degraded by human activities, especially agriculture and cattle raising (estimated from Ministerio de Ambiente del Ecuador 2013). Because habitat degradation is increasing, we assign Hypsiboas fasciatus to the Red List category Near Threatened. Its distribution polygon overlaps with three protected areas: Parque Nacional Podocarpus, Reserva Biológica el Quimi and Refugio de Vida Silvestre el Zarza. Its distribution polygon does not overlap with the polygon reported in the Red List assessment for this species by Icochea et al. (2004). The new range represents < 1% of the range reported in the IUCN Red List.

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.

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.

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
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).

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

Dorsolateral views of adult females of A Hypsiboas fasciatus, QCAZ 48611, SVL = 51.79 mm B Hypsiboas almendarizae, QCAZ 32638, SVL = 51.26 mm C Hypsiboas calcaratus, QCAZ 24282, SVL = 51.26 mm D Hypsiboas maculateralis, QCAZ 43825, SVL = 55.31 mm E Hypsiboas alfaroi, QCAZ 43252, SVL = 45.37 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.

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.

Ministerio de Ambiente del Ecuador (2013) Sistema de Clasificación de los Ecosistemas del Ecuador Continental.Subsecretaría de Patrimonio Natural, Quito, Ecuador.IcocheaJColomaLARonSR (2004) Hypsiboas fasciatus. In: IUCN 2013. IUCN Red List of Threatened Species. Version 2013.1. (15 August 2013). http://www.iucnredlist.org