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 calcaratus http://species-id.net/wiki/Hypsiboas_calcaratus (Troschel, 1848)Hyla calcarata Troschel, 1848: 660. Type material not designated and likely lost. Type locality “Britisch-Guiana” (= Guyana; Frost 2013).Diagnosis.

Hypsiboas calcaratus (Figs 4C, 8C, and 9) is characterized by: (1) mean SVL 36.82 mm in males (range 27.61–42.50; n = 36), 50.92 mm in females (range 45.94–56.29; n = 4); (2) basal webbing on the fingers; (3) calcar large and triangular; (4) dorsal background color ranging from reddish brown to cream, pinkish white or grayish brown, in most cases dark marks are present (e.g., broad transversal marks, large black stains); (5) often middorsal dark brown line 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 yellow to orange band; (12) prepollical spine present in males.

Hypsiboas calcaratus is most similar to Hypsiboas fasciatus and Hypsiboas almendarizae sp. n. It differs from both species by the shape of the calcar (large and triangular in Hypsiboas calcaratus, small and conical in Hypsiboas fasciatus, and large and conical in Hypsiboas almendarizae sp. n.; Fig. 14C–D) and by the number of notes in the advisement call (Fig. 12). Hypsiboas calcaratus can be further distinguished from Hypsiboas fasciatus by the color of the upper band in the iris: red to reddish brown in Hypsiboas fasciatus, yellow to orange in Hypsiboas calcaratus. Hypsiboas almendarizae sp. n. differs from Hypsiboas calcaratus 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. 7.89%, SD = 1.2, in Hypsiboas calcaratus; differences are significant: t = -6.72, df = 18, P < 0.001) and smaller calcars.

Hypsiboas calcaratus differs from Hypsiboas maculateralis sp. n. in advertisement call (lower dominant frequency, higher fundamental frequency; Figs 12E–F and 13A–B) and by 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 calcaratus 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) 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 calcaratus from other species are shown in Table 6.

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
Variation.

Variation in dorsal and ventral coloration of preserved specimens is shown in Figure 9. Background dorsal coloration varies from cream (e.g., QCAZ 40085) to pinkish white (e.g., QCAZ 44530), reddish brown (e.g., QCAZ 14957, 43256, 44422), pale reddish brown (e.g., QCAZ 43259) or pale grayish brown (e.g., QCAZ 48718). Irregular dorsal marks may be present in diverse patterns. A dark middorsal line extends from the tip of the snout to the mid-sacrum (e.g., QCAZ 43256), but in some specimens it only extends along the head (e.g., QCAZ 25514) or on the anterior half of the body (e.g., QCAZ 43131). There is variation in the number, size, and shape of dorsal marks. Some individuals (e.g., QCAZ 43256) have five to seven brown diffuse transversal bands (sometimes interconnected). 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, the dorsum and dorsal surfaces of the forearms and shanks have large black stains (e.g., QCAZ 14957) or scattered brown or white dots (e.g., QCAZ 40085, 44178, 14971). The coloration of flanks and hidden surfaces of thighs vary from pale cream to creamy white or light gray, with dark brown transversal bars. The number of bars on the flank varies from 4 to 13; the number of bars on the thigh varies from 4 to 9. The extent of the area with bars varies from the groin to the mid flank (e.g., QCAZ 43259) to from the groin to the axilla (e.g., QCAZ 43256). In some individuals, the bars 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 43256).

Ventral surfaces of preserved specimens vary from creamy white (e.g., QCAZ 44530) to yellowish white (e.g., QCAZ 43256). In some individuals, scattered minute pale brown blotches are present on the lips (e.g., QCAZ 31446, 44178). Coloration of webbing and discs vary from yellowish white to brown or gray. Coloration of bones is white or green.

Coloration in life.

(based on photographs; Figs 4C and 8C). Dorsal surfaces vary from light brown (e.g., QCAZ 40056) to reddish brown (e.g., QCAZ 36869) or brown (e.g., QCAZ 24282) with a middorsal dark brown line (e.g., QCAZ 40985); some individuals have brown diffuse transversal bands (e.g., QCAZ 43256); the dorsal surfaces of the limbs have pale brown transversal bars (e.g., QCAZ 43256); scattered minute white and black dots can be present on the dorsum (e.g., QCAZ 40056); in some individuals there are large dark brown blotches on the dorsum, dorsal surfaces of the forearms and shanks (e.g., QCAZ 43245); flanks are white, light blue or blue with dark brown vertical bars (e.g., QCAZ 40083); hidden surfaces of thighs and shanks are white, light blue or blue with dark brown transversal bars (e.g., QCAZ 43034); in some specimens there are dark brown transversal bars on the hidden surfaces of the shanks, ventral surfaces of the upper arms, and dorsal surfaces of the feet (e.g., QCAZ 43034); a faint creamy white stripe usually is evident on the outer edge of the feet, tarsus, forearms, and hands (e.g., QCAZ 26062); venter creamy white with belly yellowish white; ventral surfaces of hindlimbs and forelimbs translucent white (e.g., QCAZ 43824) or yellowish (e.g., QCAZ 40085); in some individuals, ventral surfaces of the thighs are creamy white (e.g., QCAZ 43047); discs and webbing yellowish (e.g., QCAZ 40085) or brown (e.g., QCAZ 40985); iris creamy silver (e.g., QCAZ 40056) or bronze (e.g., QCAZ 40085) with an upper yellow to orange band (e.g., QCAZ 43047); bones are white (e.g., QCAZ 40083) or green (e.g., QCAZ 43824).

In the examined adult series, the largest male has a SVL of 42.50 mm, and the largest female 56.29 mm; mean male SVL = 37.08 mm (n = 35; SD = 2.09), mean female SVL = 50.92 mm (n = 4; SD = 4.80). Females are significantly larger than males (t = -5.71, df = 3, P = 0.009). Inter-population variation in size and other morphometric variables is shown in Tables 1 and 2.

Advertisement call.

Two males were recorded at Tena (Provincia Napo) on 1 March 2009 and five males at Estación Científica Yasuní PUCE (Provincia Orellana) on 20 June 2009, in vegetation next to streams or ponds. Acoustic parameters of the advertisement call are shown in Table 7. The call (Fig. 12E–F) consists of a single quack note with a mean duration of 0.05 s (SD = 0.00) and mean rise time of 0.04 s (SD = 0.01). The mean dominant frequency is 1780.50 Hz (SD = 112.73) and the mean fundamental frequency is 557.13 Hz (SD = 46.21).

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
Distribution and ecology.

Hypsiboas calcaratus has confirmed records (based on DNA sequences and specimens listed in Appendix) from French Guiana, Guyana and the Amazon basin of Brazil, Ecuador, and Peru (Fig. 17). A photograph published by De la Riva et al. (2000) confirms its presence in Bolivia. Records from Colombia and Venezuela need confirmation. Known localities range in elevation from sea level (Kaw) to 650 m (Canelos).

Hypsiboas calcaratus occurs in Terra Firme forest, flooded forests (Várzea and Igapó), and swamps. It is generally found next to streams, ponds, and lakes. Individuals have been recorded at night perching on vegetation 15 to 200 cm above the ground. Their occurrence in secondary forests and artificial open areas suggest at least some tolerance of anthropogenic habitat disturbance.

Vegetation types at known localities include Southwest Amazon Moist Forest and Napo Moist Forest for the Peruvian and Ecuadorian localities, Guianan Moist Forest for the Guyana and French Guiana localities, and Madeira-Tapajós Moist Forest for the Brazilian locality (according to the World Wildlife Fund, 2012).

Conservation status.

Its distribution polygon has 3’586,597 km2 and overlaps with protected areas and large regions of pristine forest. Hypsiboas calcaratus is relatively frequent in scientific collections suggesting that, at least in part of its range, it is not a rare species. For these reasons we propose assigning Hypsiboas calcaratus to the Red List category of Least Concern.

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.

Adult Hypsiboas calcaratus showing variation in dorsal and ventral coloration of preserved specimens. From left to right, first and second rows: QCAZ 44422, 44530 (females), 14957 (male); third and fourth rows: QCAZ 43259, 40085, 43256 (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).

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 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.

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

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 Hypsiboas calcaratus and Hypsiboas maculateralis. 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.

FrostDR (2013) Amphibian Species of the World: an Online Reference. Version 5.6 (9 January 2013).Electronic database accessible athttp://research.amnh.org/vz/herpetology/amphibia/American Museum of Natural History, New York, USA.De la RivaIKöhlerJLöttersSReichleS (2000) Ten years of research on Bolivian amphibians: updated checklist, distribution, taxonomic problems, literature and iconography.Revista Española de Herpetología14: 19-164.