Leptodactylus pentadactylus (Laurenti 1768)

Leptodactylus pentadactylus (Laurenti)

MATERIALS.—USNM uncatalogued (single specimen dissected, stage 34, SVL 18.4 mm). Collected from 6 km west of Turrialba, Cartago, Costa Rica. Specimen subsequently destroyed in SEM preparation.

REFERENCES.—External descriptions are in Heyer (1970, 1979). Discussions of ecology, habitat occurrence, and facultative carnivory are found in Heyer, McDiarmid, and Weigmann (1975) and Muedeking and Heyer (1976).

GENERAL

Leptodactylus pentadactylus (Laurenti), 1768

Rana pentadactyla Laurenti, 1768:32. [Type-locality: “Indiis.” Type: based on pl. 75: fig. 1 in Seba, 1734.]

Rana gigas Spix, 1824:25. [Type-locality: Amazon River, Brazil. Type: lost.]

Rana coriacea Spix, 1824:29. [Type-locality: Amazon River, Brazil. Type: lost.]

Leptodactylus goliath Jiménez de la Espada, 1875:57. [Type-locality: Archidona and Chinitambo, Sierra de Guacamayos, Oriente, Ecuador. Lectotype: Museo Nacional, Madrid, 328, adult female.]

Leptodactylus macroblepharus Miranda-Ribeiro, 1926:144. [Type-locality: Manaos, Amazonas, Brazil. Holotype: MZUSP 377, adult male.]

Leptodactylus pentadactylus dengleri Melin, 1941:51. [Type-locality: Roque, San Martin, Peru. Holotype: Naturhistoriska Muséet, Göteborg.]

Leptodactylus pentadactylus rubidoides Andersson, 1945:47. [Type-locality: Río Pastaza, Santiago-Zamora and Napo-Pastaza border, Ecuador. Holotype: Stockholm Royal Museum.]

DIAGNOSIS.—Individuals of L. pentadactylus have a distinctive pair of dorsolateral folds. Dorsolateral folds are also found in some or all individuals of fallax, flavopictus, labyrinthicus, kundseni, rhodomystax, rhodonotus, rugosus, and stenodema. Leptodactylus pentadactylus lack distinct light lip stripes; all flavopictus and rhodomystax individuals have distinct light upper lip stripes. Most rugosus lack dorsolateral folds; when present, the folds are interrupted, contrasting with the continuous folds found in pentadactylus. Leptodactylus pentadactylus is a large species (minimum adult SVL 106 mm) with a single thumb spine in males; rhodonotus is a moderate-sized species (maximum adult SVL 90 mm) with two thumb spines per hand in males; stenodema is a moderate-sized species (maximum adult SVL 100 mm) in which the males lack thumb spines. Leptodactylus pentadactylus most closely resembles fallax, labyrinthicus, and knudseni in the pentadactylus group. A fold from above the tympanum to the side of the body, usually dark outlined for the extent, is found in pentadactylus; this fold is absent in fallax, labyrinthicus, and knudseni.

ADULT CHARACTERISTICS (N=173).—Dorsum uniform, spotted, or barred (Figure 1, A–D, G, J); lip uniform or usually with dark triangular bars (Figure 2, A–C, E, J); posterior surface of thigh uniform or variously spotted and mottled (Figure 3, A–G, I-K, P, R, S); upper tibia distinctly or indistinctly broadly barred (Figure 4, A, C, E); 2 distinct, dark outlined dorsolateral folds from eye to sacrum; dark outlined fold from eye to over tympanum, dark outline continuing or not on continuation of fold to shoulder; dark outlined fold from above tympanum to side of body usually continuous, rarely interrupted in Middle American specimens; males with one thumb spine per hand, rarely a weakly developed second; upper tibia and distal tarsal surfaces smooth or with scattered to many white- or black-tipped tubercles; sole of foot smooth, rarely with scattered white tubercles; female SVL 148.1±14.7 mm, maximum 176.2 mm, male SVL 137.2±14.0 mm, 185.0 mm maximum; female interorbital distance/head length ratio 0.19±0.02, male 0.19±0.02; female eye-nostril distance/head length ratio 0.25±0.01, male 0.25±0.01; female head length/SVL ratio 0.37±0.01, male 0.38±0.01; female head width/SVL ratio 0.39±0.02, male 0.39±0.02; female femur/SVL ratio 0.42±0.03, male 0.42±0.02; female tibia/SVL ratio 0.45±0.02, male 0.44±0.02; female foot/SVL ratio 0.46±0.02, male 0.45±0.02.

DISTINCTIVE COLORS IN LIFE.—Posterior surfaces of thigh mottled black and white or black and gray, most distinctive in juveniles (Villa, 1972, from Nicaragua; R. W. McDiarmid color notes from Peruvian specimen).

LARVAL CHARACTERISTICS.—Eye diameter 7–13 percent head-body length; oral disk position anterior; oral disk entire; oral disk width 15–22 percent head-body length; anterior oral papilla gap 50–74 percent oral disk width; anterior split tooth row about ⅓ length of entire anterior tooth row; 45–67 denticles in split tooth row anterior to beak; tooth row formula head body length 28–39 percent total length; maximum total length, stage 40, 83 mm.

MATING CALL.—Dominant frequency modulated between 250–500 Hz (Figure 12); no harmonic structure in call; call barely pulsed (Costa Rica), partially pulsed (Panama, Barro Colorado Island), or pulsed (Ecuador), (Figure 13); call duration 0.22–0.33 s (Figure 13).

KARYOTYPE.—Diploid number 22, 2 pair median, 6 pair submedian, 3 pair subterminal (Heyer and Diment, 1974).

DISTRIBUTION (Figure 14).—Middle America from Honduras south into South America; in South America, the species occurs in coastal Colombia and Ecuador and the Amazon Basin.

BRAZIL. AMAZONAS: Benjamin Constant (MNRio 1743, 1751(2), 2559); Ducke Reserve (KU 129945); Fonte Boa, Alto Solimões (MNRio 796); Igarapé Belém, Rio Solimões (MZUSP 24897); São José (Jacaré), Rio Solimões (MZUSP 40303). MARANHÃO: Rio Aurá (CAS-SU 11830). PARÁ: Canindé, Rio Gurupi (MZUSP 25010–11); Furo do Panaquera (MZUSP 35677); IPEAN (KU 127400, 128336); Marambaia (KU 140094); Monte Cristo, Rio Tapajóz (MZUSP 38956); Óbidos (MZUSP 22126); Rio Mapuera, at Equator (AMNH 49482); Sudam Floral Reserve, 74 km SE Santarem (KU 129944).

COLOMBIA. AMAZONAS: Leticia (USNM 147053). ANTIOQUIA: Chigorodó, near Turbo (USNM 153914); Finca Chibiquí, 5 km W, Río Arquia (LACM 51211). CHOCÓ: Boca de Raspadura (AMNH 39792); Condoto (UMMZ 121418); Playa de Oro, 2 km above, upper Río San Juan (FMNH 54707). MAGDALENA: Parque Nacional Tayruna, trail between Canaveral and El Pueblito (USNM 200376). META: Río Guayabero, Angostura (USNM 150490); San Juan de Arama, Los Micos (FMNH 81329); about 30 mi WNW Vista Hermosa Caño Sardinata (UTA 5233–34). PUTUMAYO: About 15 km airline SW Mocoa, 1180 m (AMNH 84865). VALLE: Río Anchicaya, 8 km W Danubio, 300 m (KU 169090); Río Calima (USNM 150756); lower Río Calima (USNM 145094); Río Raposo, Virology Field Sta, (USNM 151460–61). VAUPÉS: Río Ariari and Río Guaviari (UTA 2719, 5232).

COSTA RICA. CARTAGO: La Suiza (KU 25713–14, 28188); Moravia de Turrialba (KU 30407, 65709, UTA 1395); 2.7 mi NE Río Reventazon Bridge, Peralta Rd, 3100’ (UMMZ 117274–76); Tunnel Camp, near Peralta (KU 33165, 33167); Turrialba (AMNH 54485, 62251, FMNH 57532, 101800, 101802, KU 30408–11, 65707–08, USNM 29953–54); 10 mi S Turrialba (USNM 192577). HEREDIA: Puerto Viejo and vicinity (KU 33135–39, 33144–52, 65710–12). LIMÓN: Guapiles (MCZ 7981–84); La Lola (KU 34967, UMMZ 117277); Los Diamantes (FMNH 101797, 101799, KU 25716–18, 30405–06(3), 65706); Puerto Viejo (KU 35924); Suretka (KU 35927, UMMZ 124019, 135387, 135392); Tortuguero (AMNH 75098–5100, MCZ 29134). PUNTARENAS: Agua Buena (KU 34965, 35928–29); Esterillos Oeste, 15 km SE Jacó (KU 33153, 34968–69, 65715–17, TCWC 19305–06); Gromaco (UMMZ 123320); Rincón de Osa and vicinity (LACM 116317, USNM field 1628, 1630–31, 1634, 1641, 1663, 1666–68, 1682); Villa Neily and vicinity, 75 m (KU 65713–14, 100338–39, 100354). SAN JOSÉ: El General (KU 25715); San Isidro del General 2½ mi E (FMNH 101801); 13 mi WSW San Isidro del General on Dominical Rd, 710 m (KU 34966, 35925–26, LACM 114334).

ECUADOR. COTOPAXI: Region of Sigchos (USNM 196743). ESMERALDAS: Hacienda Equinox, 30 km NNW Santo Domingo de los Colorados, 1000’; (USNM 196739). MORONA-SANTIAGO: Cusuime, 320 m (AMNH 93705–06). NAPO: Dureno, 320 m (KU 104714); Lago Agrio, 340 m (KU 126258–59, UMMZ 129062); Limoncocha (KU 99040, LACM 92128–31); Puerto Libre, Río Aguarico, 570 m (KU 122579); Santa Cecilia (KU 104713, 104715, 104717–18, 109152–53, 111405, 111408–09, 122574–75, 122577, 122580, 126257, 146176–80, 149354–58; MCZ 57944–46, 57948–52, 57954–56, 57959–60, UMMZ 129280–81). PASTAZA: Andoas (AMNH 71025); Canelos (USNM 196742); 2 km E Puyo (USNM 196741(2)); headwaters of Río Arajuno, tributary of Río Napo (USNM 196744); Río Pindo (USNM 196747); Río Rutuno, tributary of Río Bobonaza (USNM 196746). PICHINCHA: Espinosa, 9 km W Santo Domingo de los Colorados, road to Chone (CAS-SU 10453–54); 18 km W Santo Domingo de los Colorados, Ramsey Farm, km 19 Chone Rd. (USNM 196740(2), 196745).

FRENCH GUIANA. Sophie (MCZ 44562).

GUYANA. Rupununi, N of Acarahy Mts (KU 69685); Shudi-kar-wau (AMNH 70118–20).

HONDURAS. COLON: Belfate (AMNH 45704, 45722).

NICARAGUA. BLUEFIELDS: 6 mi W Rama, 50’ (TCWC 19307). GRANADA: N Granada (LACM 37870). MATAGALPA: 10.5 km N Finca Tepeyac, 9 km E Matagalpa, 960 m (KU 85146). RÍO SAN JUAN: San Juan del Norte (Greytown) (USNM 19765). ZELAYA: Bonanza (KU 85147–48, 101168); Camp Corozo, Río Huahuashan (AMNH 54980–81); Camp Santa Ana, Río Huahuashan (AMNH 54999); Cara de Mono, 50 m (KU 112673–75); Masawas, Waspuk R (AMNH 58435); El Recreo, 25 km W Rama (KU 112666–72, LACM 13945, 20475); 10 mi above Recreo, Río Mico (UMMZ 79751–52).

PANAMA. BOCAS DEL TORO: Almirante (KU 79992, USNM 142334); Boca del Drago (USNM 142318); Punta de Peña (USNM 38714). CANAL ZONE: Barro Colorado Island (AMNH 52878–79, 69728, FMNH 175986, KU 76568, MCZ 15266, UMMZ 64594, 124019, 135387, 135392, USNM 161151); Candelaria and Peluca Stas, near Boqueron (AMNH 53747); Cave A, Chili Brillo (AMNH 62338); Frijoles (USNM 196303); Las Cruces trail near Corozal (AMNH 38977); near Madden Dam (AMNH 87143, KU 155306, UMMZ 78481). CHIRIQUÍ: Progreso (UMMZ 58221). COCLÉ: El Valle, 560 m (AMNH 59590, KU 76575, 107229, 116825.) DARIÉN: Camp Creek, Camp Townsend (AMNH 40785–88, 40793, 41061, 41765); Casita (USNM 141783); Jaqué (USNM 161215); Laguna, 820 m (KU 76570–72); along Río Canglón, near mouth (UMMZ 123160–61); across from mouth of Río Canglón, along Río Chucunaque (UMMZ 123158–59); Río Chucunaque, about 7 km above Río Mortí, 150 m (KU 107230); Río Silugandí (UMMZ 124018); Río Tuira at Río Mono (KU 115308); Río Ucurgantí, about 7 km above mouth, 30 m (KU 97014–17), 107231–32); Tacarcuna, 550 m (KU 76573–74). LOS SANTOS: Guánico Arriba, 60 m (KU 107228). PANAMÁ: Altos de Maje (AMNH 88743); Cerro Campana (KU 76569); Ciricito (CAS 71514–16); Juan Mina, Chagres R (USNM 129908); Tapia (AMNH 18920, 18922–23). SAN BLAS: Armila (USNM 150090); Camp Sasardí, 12 m (KU 108688–89); Quebrada Venado (USNM 150089). VERAGUAS: Mojara (UMMZ 123320); mouth of Río Concepción, 1 m (KU 11537).

PERU. AYACUCHO: La Mar, Candalosa (FMNH 39720). HUANUCO: Ganzo Azul (FMNH 45145); Río Llullapichis, 4–5 km upstream from Río Pachitea (KU 171901). LORETO: Cashiboya (AMNM 42404); Iquitos (AMNH 42002, 42560); Peru-Brazil border, upper Utoquinea (AMNH 42751, 43371); 61 mi SE Pucallpa, 500’ (TCWC 4967); Río Pisqui (AMNH 43539); Río Ucayali, Yarinacocha (FMNH 56293, 56300). SAN MARTÍN: Upper Biabo Valley (AMNH 42656); Pachisa (AMNH 42001); Tocache Nuevo, Río Huallaga (AMNH 42562).

SURINAM. Godo Drai (TCWC 23561–63); King Frederick William IV Falls (FMNH 30912–13).