GENERIC RELATIONSHIPSAmong the 33 genera currently recognized in the family Hylidae, there are two basic types of vocal sac structure (Duellman, 1970b),namely the subgular type and the lateral type. Only four hylid genera, all Neotropical lowland groups, are known to possess paired lateral vocal sacs; these areOsteocephalus,Argenteohyla,Phrynohyas, andTrachycephalus. The geographical distributions and morphological characteristics of these four genera suggest that they are more closely related to one another than with any other hylid genera.Of the four genera,Osteocephalusis the most generalized in morphology, and, likePhrynohyas, has no specialized habits.OsteocephalusandArgenteohylaare similarly distinguished fromPhrynohyasandTrachycephaluson the basis of vocal sac structure. The vocal sacs ofOsteocephalusandArgenteohylaare posterior and protrude posterolateral to the angles of the jaws when they are inflated, whereas those ofPhrynohyasandTrachycephalusare more lateral and protrude posterior to the angles of the jaws when inflated.AlthoughOsteocephalusandArgenteohylahave similar vocal sac structure, they are obviously distinct. The monotypicArgenteohylais a rather specialized, semifossorial frog (Trueb, 1970b), characterized by smooth skin, moderate-sized digital discs, and a large inner metatarsal tubercle. The general architecture of the skull is not unlike that ofOsteocephalus; the skulls of both are well roofed, broader than long, and characterized by posterolaterally oriented parasphenoid alae.Argenteohylabears small, slightly curved prevomerine dentigerous processes in contrast to the large, angular processes ofOsteocephalus. The skull ofArgenteohylashows specializations, apparently adaptations to its semifossorial mode of existence, which further distinguish the genus fromOsteocephalus. In comparison withOsteocephalus, the cranium ofArgenteohylais slightly depressed anteriorly, the roofing bones extensively casqued, and the palatines robust.Osteologically,Osteocephalusmore closely resemblesPhrynohyasthan either of the other two genera, butOsteocephalusandPhrynohyasare clearly distinct on the basis of their respective vocal sac structure. LikeOsteocephalus, skulls of the members of the genusPhrynohyasare broader than long, have extensive dermal roofing bones, and have posterolaterally oriented parasphenoid alae. In contrast toOsteocephalus, the dentigerous processes of the prevomers are curved, rather than angular inPhrynohyas. Furthermore, the latter genus is singularly distinguished fromOsteocephalus,Argenteohyla, andTrachycephalusby having extensively developedparotoid glands that produce a viscous, milky volatile secretion.Trachycephalusis the most readily identifiable of the four genera under discussion. Members of this genus are large frogs with heavily casqued and co-ossified skulls (Trueb, 1970a). The dermal roofing bones bear ornate and characteristic patterns of sculpturing. The medial ramus of the pterygoid does not articulate with the otic capsule, and the parasphenoid alae are laterally, rather than posterolaterally, oriented. A dermal sphenethmoid is present, and the parasphenoid bears odontoids. The basic structure of the skull has many characters in common with bothOsteocephalusandPhrynohyas. The obvious modifications of dermal roofing bones and of palatal and suspensory elements seem to be specializations adapting members of the genusTrachycephalusto their peculiar phragmotic habits. The vocal sac structure ofTrachycephalusis like that ofPhrynohyasand therefore further distinguishes it fromOsteocephalus.Morphologically,Osteocephalusseems to be sufficiently diverse and generalized so as to represent a modern derivative of an ancestral type which might have given rise toPhrynohyas,Trachycephalus, andArgenteohyla. The specialized vocal sac structure inPhrynohyasandTrachycephalussuggests that these two genera may be rather closely allied and represent a single phyletic line from an ancestral stock similar toOsteocephalus.Argenteohylais quite distinct fromPhrynohyasandTrachycephalusand apparently represents a distinct phyletic line from the ancestral stock.OCCURRENCE OFOSTEOCEPHALUSIN AMAZONIAN ECUADORAll of our observations on members of this genus have been made at four localities: 1) Santa Cecilia at an elevation of 340 meters on the Río Aguarico, a tributary of the Río Napo, 2) Lago Agrio, 330 meters, about 14 kilometers east of Santa Cecilia, 3) Puerto Libre, 570 meters, on the Río Aguarico just east of its formation by the confluence of the Río Cofanes and Río Chingua, and 4) south slope of the Cordillera del Dué, above the Río Coca, 1150 meters.Osteocephalus leprieuriiwas found at all four localities, andbuckleyiwas found at all but the last;taurinuswas found at Santa Cecilia and Lago Agrio, andverrucigeruswas found only in the Cordillera del Dué. Our data are based on collections of 113 frogs and three lots of tadpoles, as well as observations on calling sites and young. The observations are summarized by species, as follows:Osteocephalus buckleyi.—No breeding activity was observed. Males were found only at night in March, June, and July. One was perched on aHeliconialeaf in a swamp at Puerto Libre, and two were on bushes in the forest at Santa Cecilia. A gravid female was found on a recently felled tree at Lago Agrio on the night of 12 May 1969.Osteocephalus leprieurii.—Males were heard calling sporadically at Puerto Libre in July 1968, and at Santa Cecilia in May 1969. A small chorus was found on the night of 12 May 1969 at Lago Agrio, where the frogs were perched on branches of fallen trees over a temporary pool. The call is a soft rattling chuckle. In late April and May many gravid females and males with well-developed nuptial excrescences were obtained from trees as they were felled at Lago Agrio. The reproductive condition of the frogs indicates that they probably breed in May. One individual called nearly every night from a large tree at Puerto Libre between 4-17 July 1968. The tree was felled on the latter date, but no frog was found. Two nights later apparently the same individual called from abromeliadat a height of about 10 m on a large bamboo adjacent to the felled tree; the frog was collected when the bamboo was cut down.Throughout the rainy months that we have worked in Ecuador (April-August) we have found occasional individuals perched on bushes or low trees at night. Large numbers of adults were observed only during a clearing operation which resulted in the felling of many large trees. Thus, it seems likely thatleprieuriiis a tree-top inhabitant. A partially digested adult male was removed from the stomach of aHemiphractus proboscideus.At Santa Cecilia many recently metamorphosed young and juveniles were found in June and July 1968. Most of these were on low bushes or herbs in swamp forest at night; some were found in unfoldedHeliconialeaves by day, and one was observed on the forest floor by day. Snout-vent lengths of 18 specimens are 12.3-17.0 (mean 15.1) mm. The smaller frogs were recently metamorphosed as evidenced by the melanophore deposits above the vent. The coloration of the young is strikingly different from that of the adults (see account ofO. leprieurii), so the association of the young and adults was not made until individuals with intermediate patterns were obtained at Lago Agrio in May 1969. Probably juveniles obtained in June and July are the offspring of an April or May breeding. We have been unable to associate tadpoles with this species.Osteocephalus taurinus.—A small chorus occurred at Lago Agrioon 12 May 1969. Males were calling from the ground adjacent to a small pool amidst recently felled trees. The males were very wary and, when approached, jumped onto limbs and ran up branches; this behavior was noted by Bokermann (1964). The call consists of a series of low-pitched, short notes—like a slow trill—four to six notes per call group. Call groups are repeated two, three, or four times followed by a lapse of several minutes. Although no amplectant pairs were found, several gravid females were collected at Lago Agrio in May, so it can be safely assumed that the species breeds in May. From April through July occasional individuals were observed on bushes and trees at night. During clearing operations at Lago Agrio several individuals were obtained from the tops of trees as they were felled.Osteocephalus verrucigerus.—Observations were made in a broad, shallow ravine, in which there was a small stream. On 2-4 August 1968, males were observed calling from low bushes and rocks at the edge of a quiet pool in the stream. The call consists of a series of well-pulsed, low-pitched, guttural notes produced at the rate of 5-10 per minute. One amplectant pair was found at the base of a bush adjacent to the pool on 3 August. Another female was found on a branch of a tree 2 m above the ground and 10 m from the stream. Tadpoles of this species were found in the quiet silt-bottomed pool.SPECIMENS EXAMINEDThe localities for each of the specimens examined are given in the following paragraphs. The arrangement of the data is as follows: alphabetically by country, state (department or province), and locality; alphabetically by the first letter in the abbreviations for the museums, and numerically after each museum abbreviation. Specimens lacking precise locality data are listed first in the most restricted political unit possible; localities which have not been found on maps or the positions of which are not known to us are given in quotation marks. Where more than one specimen is included under one museum number, the number of specimens is given in parentheses after the museum number. Unless noted otherwise, all specimens are alcoholics.Osteocephalus buckleyiBOLIVIA:El Beni: Ivón, BMNH 1967.2070-1.Santa Cruz: Buenavista, CM 4333, 4339, UMMZ 66563-5.BRASIL:Amapá: No specific locality, WCAB 13284.COLOMBIA:Amazonas: Río Guacaya, USNM 152759.Huila: Acevedo,Río Suaza, FMNH 69702.Nariño: Rumiyacu, FMNH 54756.Meta: Río Guejar, Campamento La Macarena, USNM 152199.ECUADOR: No specific locality, NHMW 6209, WCAB 35499.Chimborazo: Pallatanga, BMNH 1947.2.13.46; Santiago, FMNH 42529.Morona-Santiago: "Río Santiago" (= Río Zamora), MIZS 2950.Napo: Lago Agrio, KU 126646; Puerto Libre, Río Aguarico, KU 123172; Santa Cecilia, AUM 8138, KU 105208-9, 109506, 123171.Pastaza: Alpayacu, BMNH 1912.11.1.64; Canelos, BMNH 1947.2.13.40-1, 1947.2.13.43-5; Colonia Mena, Río Conambo, ZSM 33/1962; Don Tomás, USNM 166014; Guaché, Río Pastaza, AMNH 79986; Río Bobonaza, USNM 166005; Río Capahuari, USNM 166554; Río Conambo at Río Shiona-yacu, USNM 166018; Río Copataza, upper Río Pastaza, USNM 166007-13; Río Pastaza, NHRM 1946; Río Pucyacu, USNM 165997 (skeleton), 165998-6001; Río Rutuno, USNM 166006; Río Villano, USNM 166002-4; Sarayacu, BMNH 1947.2.13.36-9, MCZ 26090, ZMB 10166.GUYANA:Mazaruni-Potaro: Kartabo, AMNH 70971; Membaru River, upper Mazaruni River, UMMZ 85168; Oko Mountains, FMNH 26722-3.North West: Amakura River, Haulover, UMMZ 83558-9.Rupununi: Marudi River, AMNH 46233; Shudi-kar-wau, AMNH 49252.West Demerara: Dunoon, UMMZ 52449, 52508.PERÚ:Junín: Chanchamayo, BMNH 1911.12.13.79-80.Loreto: Andoas, AMNH 79984-5; Cashiboya, AMNH 43454; San Antonio, Río Itaya, AMNH 43218.Puno: Yahuaramayo, BMNH 1913.2.25.7.SURINAM:Suriname: Powakka, CM 44217.SOUTH AMERICA: No specific locality, NHMW 6208.Osteocephalus leprieuriiBRASIL:Acre: Tarauacá, FMNH 83247.Amazonas: Rio Javarí, Benjamin Constant, CAS-SU 12620; Río Uaupés, north of Rio Japú, NHMG 489.COLOMBIA:Amazonas: Gino-goje, lower Río Apoporis, MCZ 28038, 28040-2, 28044, USNM 152136-8.ECUADOR: No specific locality, WCAB 35452-3; "Napo-Pastaza," USNM 166571.Napo: Avila, UMMZ 92093; south slope Cordillera del Dué, KU 123170; Lago Agrio, KU 125961-2 (skeletons), 126611-44, UMMZ 129326 (2); Limón Cocha, Río Napo, KU 99210-6, UIMNH 63087-9, 63098, 63106-9, 63118-9, 64802-4, 64858, 87998-9, 88001-30, 88437-8, 88580, 88604-5, 89852-97, 89999-90000; Loreto, CAS-SU 11439, WCAB 36526; Puerto Libre, Río Aguarico, KU 123190-1; Puerto Napo, UIMNH 55818-20; Río Cotapino, UMMZ 92094; Río Napo, UMMZ 92078; Santa Cecilia, AUM 8099, 8102, 8113-5, 8127-9, 8131, 8137, 8139-46, 8148, KU 105210-20, 109509-11, 111971, 122964-87, 123169, 126645.Pastaza: Canelos, BMNH 1947.2.13.42, KU 120915; Río Alpayacu, UMMZ 92079; Río Arajuno, USNM 166560-2, WCAB 40176; Río Oglán, USNM 16655203, 166558; Río Rutuno, USNM 166559; Río Shilcayacu, below Puyo, USNM 166557; Río Villano, USNM 166551.FRENCH GUIANA: No specific locality, MNHN 4629.Inini: Lunier River, MNHN 98/217.GUYANA:Mazaruni-Potaro: Kartabo, AMNH 70967-8, 70972, 70976.Rupununi: Shudi-kar-wau, AMNH 49255.West Demerara: Demerara Falls, BMNH 72.10.16.23, 72.10.16.37-8.PERÚ:Loreto: Estirón, Río Ampiyacu, MZUSP 31033-4; Pebas, CAS-SU 3158, 3160; Roaboya, AMNH 43064.SURINAM: No specific locality, MCZ 2036, RMNH 11468.Marowijne: Camp 3, RMNH 13045-6; Wane Creek North, RMNH 11469-70.Saramacca: Right Coppename River, RMNH 11467.Osteocephalus pearsoniBOLIVIA:El Beni: upper Río Beni, below mouth of Río Mapiri, MCZ 15565, UMMZ 57548, 67464-5; Rurrenbaque, UMMZ 57533.PERÚ:Pasco: Yaupi, KU 136312.Osteocephalus taurinusBOLIVIA:El Beni: Ivón, BMNH 1967.2040; Reyes, UMMZ 57532.La Paz: San Ernesto, Mapiri District, BMNH 1901.8.2.54.Santa Cruz: Buenavista, AMNH 33951-2, 33958, BMNH 1927.8.1.19, 1927.8.1.118, FMNH 27091, UMMZ 63319-21, 63959(2), 63961(2), 66566(2), 66567, 66568(2), 66569 (2), 66570, 66571(2), 66575-6, 68196; Río Mamore, 2 km N Boca Chaparé, AMNH 79324; Sara, CM 3840-1; Surutu, CM 3814-5.BRASIL: No specific locality: "Interior," BMNH 74.7.16.8-9.Acre: Plácido de Castro, MZUSP 6518; Tarauacá, WCAB 2496.Amazonas: Cucuí, NHMW 16495; Manacapurú, ZMB 28492, ZSM 278/1925; Manáus, MCZ 56281, NHMW 16492; Maués, AMNH 69623, 76177; Taracuá, NHMG 488, WCAB 18463-4.Mato Grosso: Mabuca, MZUSP 4272; Posto Coluene, Rio Xingú, WCAB 812; "Puerto Cabello," AMNH 3154; Tapirapé, AMNH 73647-62, CAS-SU 12351-6, MNHN 46/324.Pará: No specific locality, MPEG 623-6; Belém, KU 129866; Cachimbo, FMNH 175876, UIMNH 42149, WCAB 813; Cametá, NHMW 15892; Gurupá, BMNH 96.6.29.13; Ilha de Marajó, BMNH 1923.11.9.20-4; Ilha Mexicana, ZSM 111/1911, 112/1912; "Ponto Dois Indios," BMNH 1939.1.5.5; Santarém, BMNH 75.10.22.1-4, MCZ 354.Rondonia: Abuná, CAS 49773-4, FMNH 64239; Forte Principe da Beira, WCAB 10230; Igarapé Marmelo, FMNH 140254, KU 84725 (skeleton), 92243-6, 92247-8 (skeletons), WCAB 9997, 10001, 10003-4; Porto Velho, MZUSP 16343.COLOMBIA:Amazonas: Gino-goje, lower Río Apoporis, USNM 152139; Leticia, USNM 152010-1; Raudal de la Playa, lower Río Apoporis, MCZ 28050; Río Apoporis, MCZ 28060.Boyacá: Sutatenza, USNM 152054-6.Cundinamarca: Medina, MCZ 16269-71, USNM 152089-90, 152092-7, 152757.Meta: El Mico, Río Guejar, USNM 152203; Río Duda, Sierra de Macarena, AMNH 79914; Río Guapaya, Sierra de Macarena, FMNH 81332; Río Guaviari, Casa de Piedra, UTA No number.Putumayo: Río Mecaya, FMNH 69711-4, 69716.Vaupés: Gomogoje, lower Río Apoporis, MCZ 28048.ECUADOR: No specific locality, WCAB 35451, 35785; "Oriente," UMMZ 90418.Carchí: below Salinas, USNM 166059.Esmeraldas: Carondelet, UIMNH 53560-9; Lagartera, Río Caoni, UIMNH 53441, 53458-79.Morona-Santiago: Macuma, UIMNH 63142-3, 63145, 63147, 63151, 63154, 63157, USNM 166060.Napo: Avila, UMMZ 92077; Cuyabeno, UIMNH 63158, 90111; Lago Agrio, KU 126647-55; Limón Cocha, Río Napo, AUM 8132-4, KU 99207-8, 99421-3, 99424 (skeleton), 99425, UIMNH 64801, 87798, 87800, 88032-5, 88576, 90066, 90082, 90102, 90104, 90314, 90984; Loreto, WCAB 35352; Río Cotapino, UMMZ 92080; Río Napo, UMMZ 84120; San José Abajo, AMNH 1295, 1449, 22180, 79990; Santa Cecilia, AUM 8117, 8150, KU 105230-3; south slope Volcán Sumaco, USNM 166570.Pastaza: No specific locality, ZSM 31/1956; Arajuno, USNM 165995; Bufeo, lower Río Bobonaza, USNM 166046-8; Canelos, BMNH 80.12.5.179, 1947.2.13.48, UMMZ 89066;Don Tomás, Río Bobonaza, USNM 166049-50; Montalvo, CAS-SU 10320, USNM 165987-9, 166058, 166566; 2.5 km SE Puyo, USNM 166051; Río Arajuno, USNM 166043-5; Río Arajuno (headwaters), USNM 166053; Río Bobonaza, WCAB 3613-4, 35504; Río Capahuari, USNM 165990, 166555-6; Río Capahuari (headwaters), USNM 166057; Río Conambo, USNM 166569, ZSM 28/1962, 35/1962; Río Conambo at Río Ollaguanga, USNM 166568; Río Conambo at Río Shiona-yacu, USNM 166019, 166563-5; Río Corrientes, USNM 195994, 166020-38, WCAB 3841-2; Río Huiyo-yacu, Pico de Conambo, USNM 166052; Río Pastaza, MCZ 19697; Río Pastaza (drainage), NHRM 1966, USNM 165996; Río Pindo, USNM 166039-41; Río Pindo at Río Tigre (village), USNM 165992-3, 166042; Río Pucayacu, USNM 166054, 166056; Río Rutuno, USNM 166055; Río Solis, upper Río Bobonaza, WCAB 39914; Río Villano, USNM 165991, 166567; Sarayacu, BMNH 80.12.5.213, 80.12.5.239-40, MZUSP 323; Shell Mera, KU 99420.Zamora-Chinchipe: "Yani-Inzari," AMNH 43259, 43394; Zamora, AMNH 78928.FRENCH GUIANA:Cayenne: Crique Grégoire, UP 40; Maripa, Oyapok River, UP 72; Oyapok River, UZM 1473.Inini: Crique Gabrielle, UP 118-20.GUYANA: No specific locality: RMNH 1873(3), ZMB 3102(2).East Demerara: Atkinson Field, ASU 11622.Mazaruni-Potaro: Chinapora River, upper Potaro River, BMNH 1905.11.1.20-1; Kamakusa, AMNH 21416, 21418-9, 21422; Kartabo, AMNH 11689, 11691, 11697-9, 11703, 11706-8, 23107, 39730, 70966, 70969-70, 70973-5, USNM 118057; Moraballi Creek, Essequibo River, BMNH 1930.10.10.47-51; Oko Mountains, FMNH 26692-705; upper Potaro River, Tung District, BMNH 1905.11.1.40; Rockstone, FMNH 26591.North West: Amakura River, Haulover, UMMZ 83735.Rupununi: north of Acaray River, west of New River, KU 69747-8; Kuyuwini Landing, AMNH 46283; Pakaraima Mountains, BMNH 1933.6.19.49; Shudi-kar-wau, AMNH 10665, 39637, 49256(2).West Demerara: Demerara, CAS 54773-4; Demerara Falls, BMNH 72.10.16.16-22, 72.10.16.25-32; Dunoon, MCZ 4834, UMMZ 46736, 52493-4, 52502, 52504-5, 57271; Vryheid, BMNH 78.12.13.18.PERÚ:Amazonas: Río Cenepa, AMNH 43400.Huanuco: Monte Alegre, Río Pachitea, AMNH 43014, 43019.Loreto: Achinamisa, Río Huallaga, AMNH 42178, 42502; Andoas, Río Pastaza, AMNH 79991; Cashiboya, AMNH 43388, 43453; Estirón, Río Ampiyacu, CAS 93264-74, 93276, 93278-9, 93281, 93283-6, 93289, 93311, 93327; Igarapé Champuia, upper Río Curiuja, MZUSP 10339; Iquitos, AMNH 42204, 42442, 43468, NHMW 6118; Lago de Miraño, mouth of Río Napo, AMNH 42712, 43186; Nauta, ANSP 11399; Ollanta, AMNH 42865; Pampa Hermosa, Río Cushabatay, AMNH 43124, 43146; Pebas, CAS-SU 6375; Pucallpa, MJP 101(2), 140(3); Punga, Río Tapiche, AMNH 43194; "Rancho de Indiana, Iquitos District," MVZ 16890; upper Río Abujao, AMNH 42908; Río Itaya, AMNH 42755; upper Río Pisqui, AMNH 43536; Río Tapiche at Río Contaya, AMNH 42983; Río Utoquinia at Brasilian frontier, AMNH 43137; Sobral, Río Tamaya, AMNH 43242; Yurimaguas, BMNH 84.2.18.50.San Martín: Cainarachi, AMNH 42763; Moyobamba, ZSM 19/1914.SURINAM: No specific locality, BMNH 70.3.10.67, NHMW 18433.3.Brokopondo: Afobaka, RMNH 16536; Brownsweg, RMNH 16537; Railway km. 121, RMNH 16534.Marowijne: Djai Creek, RMNH 16513-4; Maroni River, ZMB 8240, 8531; Nassaugebergte, RMNH 16517-33; Paloemeu, USNM 159025; Swamp Camp, RMNH 16515.Nickerie: Sipaliwini, RMNH 16538.Saramacca: Left Coppename River, RMNH 16535; Tibiti, RMNH 16516.Suriname: Berlijn, RMNH 15064; Powakka, CM 44226; Zanderij, CM 50568.VENEZUELA:Amazonas: Cerro Duida, UPR-M 2875; Cerro Marahuaca, UPR-M 114-5; Esmeralda, AMNH 23174; Iniridi, SMF 2640; La Culebra, MCZ 28572, UPR-M 117; Laguna, between Tama Tama and Esmeralda, UPR-M 2760; Río Pescado, AMNH 23177; Tapara, UPR-M 113.Distrito Federal: Caracas, BMNH 51.7.17.182.Osteocephalus verrucigerusCOLOMBIA:Huila: Acevedo, Río Suaza, FMNH 69709-10.ECUADOR: No specific locality, ZMB 16589.Napo: Avila, UMMZ 90413; south slope Cordillera del Dué, KU 123176-88, 123189 (skeleton), 124208 (eggs), 124209-11 (tadpoles); L'Alegria, USNM 167472-3; Río Pacayacu, tributary of Río Cotapino, CAS-SU 13150; southeast slope Volcán Sumaco, CAS-SU 11442.Pastaza: Abitagua, CAS-SU 5067, FMNH 25791, 27619, UMMZ 90414, 92092; Alpayaca, Río Pastaza, BMNH 1912.11.1.64; Mera, UMMZ 90412(4).Tungurahua: Baños, NHRM 1960.PERÚ:Ayacucho: La Mar, Sivia, Río Apurimac, FMNH 39853.Huanuco: Río Pachitea, midway between Puerto Victoria and Puerto Inca, CAS-SU 17745.Junín: Satipo, MJP 38.LITERATURE CITEDAndersson, L. G.1945. Batrachians from east Ecuador collected 1937, 1938 by Win. Clarke-MacIntyre and Rolf Blomberg. Arkiv Zool., 37A(2):1-88.Bokermann, W. C. A.1964. Field observations on the hylid frogOsteocephalus taurinusFitz. Herpetologica, 20:252-255.1966. Lista anotada das localidades tipo de anfíbios Brasileiros. São Paulo, 183 pp.Boulenger, G. A.1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum, ed. 2, London, xvi+503 pp.Cochran, D. M.andC. J. Goin1970. Frogs of Colombia. Bull. U.S. Natl. Mus., 288:xii+655 pp.Cope, E. D.1867. On the families of the raniform Anura. Jour. Acad. Nat. Sci. Philadelphia, 2:189-206.1874. On some Batrachia and Nematognathi brought from the upperAmazon by Prof. Orton. Proc. Acad. Nat. Sci. Philadelphia, 25:120-137.Duellman, W. E.1970a. Identity of the South American hylid frogGarbeana garbei. Copeia, (3):534-538.1970b. The hylid frogs of Middle America. Monog. Mus. Nat. Hist., Univ. Kansas, 1:xi+753 pp.Duméril, A. M. C.andG. Bibron1841. Erpétologie générale ou histoire naturelle compléte des reptiles, vol. 8. Paris, 792 pp.Fitzinger, L.1843. Systema reptilium. Vienna, ix+106 pp.Funkhouser, J.1956. New frogs from Ecuador and southwestern Colombia. Zoologica, 91:73-80.Gaige, H. T.1929. Three new tree-frogs from Panama and Bolivia. Occas. Papers Mus. Zool. Univ. Michigan, 207:1-6.Goin, C. J.1961. Synopsis of the genera of hylid frogs. Ann. Carnegie Mus., 36:5-18.Melin, D.1941. Contribution to the knowledge of Amphibia of South America. Göteborgs Kungl. Vetensk.-och Vitterh.-Sam. Handl., Ser. B, 1(4):1-71.Peracca, M. G.1904. Viaggio del Dr. Enrico Festa nell' Ecuador e regioni vicine. Reptile ed amfibii. Boll. Mus. Zool. Anat. Comp., Univ. Torino, 19:1-41.Steindachner, F.1862. Über zwei noch unbeschriebene Batrachier. Arch. Zool. Anat. Fisiol., 2:77-82.1867. Amphibien. Novara Expedition. Zool. Theil, I, Vienna, 70 pp.Trueb, L.1970a. The evolutionary relationships of casque-headed treefrogs with co-ossified skulls (family Hylidae). Univ. Kansas Publ. Mus. Nat. Hist., 18:547-716.1970b. The generic status ofHyla siemersiMertens. Herpetologica, 26:254-267.Trueb, L.andW. E. Duellman1970. The systematic status and life history ofHyla verrucigeraWerner. Copeia (4):601-610.Tyler, M.1971. The phylogenetic significance of vocal sac structure in hylid frogs. Univ. Kansas Publ. Mus. Nat. Hist., 19:319-360.Werner, F.1901. Ueber Reptilien und Batrachier aus Ecuador und Neu-Guinea. Verh. Zool.-Bot. Gesell. Wien, 50:593-614.University of Kansas PublicationsMUSEUM OF NATURAL HISTORYThe University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly published in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous Publications, Museum of Natural History, began with number 1 in 1946. Longer research papers are published in that series. Monographs of the Museum of Natural History were initiated in 1970.Institutional libraries interested in exchanging publications may obtain the Occasional Papers and Miscellaneous Publications by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas 66044. Individuals may purchase separate numbers of all series. Prices may be obtained upon request addressed to Publications Secretary, Museum of Natural History, University of Kansas, Lawrence, Kansas 66044.Transcriber's NotesExcept for the list of corrections below and minor corrections not listed, the text presented here is that of the original printed version.Typographical CorrectionsPageCorrection5is ⇒ in5buckley ⇒ buckleyi17scaral ⇒ sacral19Provicia ⇒ Provincia25in ⇒ is25metalic ⇒ metallic41bromeiad ⇒ bromeliad
GENERIC RELATIONSHIPS
Among the 33 genera currently recognized in the family Hylidae, there are two basic types of vocal sac structure (Duellman, 1970b),namely the subgular type and the lateral type. Only four hylid genera, all Neotropical lowland groups, are known to possess paired lateral vocal sacs; these areOsteocephalus,Argenteohyla,Phrynohyas, andTrachycephalus. The geographical distributions and morphological characteristics of these four genera suggest that they are more closely related to one another than with any other hylid genera.
Of the four genera,Osteocephalusis the most generalized in morphology, and, likePhrynohyas, has no specialized habits.OsteocephalusandArgenteohylaare similarly distinguished fromPhrynohyasandTrachycephaluson the basis of vocal sac structure. The vocal sacs ofOsteocephalusandArgenteohylaare posterior and protrude posterolateral to the angles of the jaws when they are inflated, whereas those ofPhrynohyasandTrachycephalusare more lateral and protrude posterior to the angles of the jaws when inflated.
AlthoughOsteocephalusandArgenteohylahave similar vocal sac structure, they are obviously distinct. The monotypicArgenteohylais a rather specialized, semifossorial frog (Trueb, 1970b), characterized by smooth skin, moderate-sized digital discs, and a large inner metatarsal tubercle. The general architecture of the skull is not unlike that ofOsteocephalus; the skulls of both are well roofed, broader than long, and characterized by posterolaterally oriented parasphenoid alae.Argenteohylabears small, slightly curved prevomerine dentigerous processes in contrast to the large, angular processes ofOsteocephalus. The skull ofArgenteohylashows specializations, apparently adaptations to its semifossorial mode of existence, which further distinguish the genus fromOsteocephalus. In comparison withOsteocephalus, the cranium ofArgenteohylais slightly depressed anteriorly, the roofing bones extensively casqued, and the palatines robust.
Osteologically,Osteocephalusmore closely resemblesPhrynohyasthan either of the other two genera, butOsteocephalusandPhrynohyasare clearly distinct on the basis of their respective vocal sac structure. LikeOsteocephalus, skulls of the members of the genusPhrynohyasare broader than long, have extensive dermal roofing bones, and have posterolaterally oriented parasphenoid alae. In contrast toOsteocephalus, the dentigerous processes of the prevomers are curved, rather than angular inPhrynohyas. Furthermore, the latter genus is singularly distinguished fromOsteocephalus,Argenteohyla, andTrachycephalusby having extensively developedparotoid glands that produce a viscous, milky volatile secretion.
Trachycephalusis the most readily identifiable of the four genera under discussion. Members of this genus are large frogs with heavily casqued and co-ossified skulls (Trueb, 1970a). The dermal roofing bones bear ornate and characteristic patterns of sculpturing. The medial ramus of the pterygoid does not articulate with the otic capsule, and the parasphenoid alae are laterally, rather than posterolaterally, oriented. A dermal sphenethmoid is present, and the parasphenoid bears odontoids. The basic structure of the skull has many characters in common with bothOsteocephalusandPhrynohyas. The obvious modifications of dermal roofing bones and of palatal and suspensory elements seem to be specializations adapting members of the genusTrachycephalusto their peculiar phragmotic habits. The vocal sac structure ofTrachycephalusis like that ofPhrynohyasand therefore further distinguishes it fromOsteocephalus.
Morphologically,Osteocephalusseems to be sufficiently diverse and generalized so as to represent a modern derivative of an ancestral type which might have given rise toPhrynohyas,Trachycephalus, andArgenteohyla. The specialized vocal sac structure inPhrynohyasandTrachycephalussuggests that these two genera may be rather closely allied and represent a single phyletic line from an ancestral stock similar toOsteocephalus.Argenteohylais quite distinct fromPhrynohyasandTrachycephalusand apparently represents a distinct phyletic line from the ancestral stock.
OCCURRENCE OFOSTEOCEPHALUSIN AMAZONIAN ECUADOR
All of our observations on members of this genus have been made at four localities: 1) Santa Cecilia at an elevation of 340 meters on the Río Aguarico, a tributary of the Río Napo, 2) Lago Agrio, 330 meters, about 14 kilometers east of Santa Cecilia, 3) Puerto Libre, 570 meters, on the Río Aguarico just east of its formation by the confluence of the Río Cofanes and Río Chingua, and 4) south slope of the Cordillera del Dué, above the Río Coca, 1150 meters.Osteocephalus leprieuriiwas found at all four localities, andbuckleyiwas found at all but the last;taurinuswas found at Santa Cecilia and Lago Agrio, andverrucigeruswas found only in the Cordillera del Dué. Our data are based on collections of 113 frogs and three lots of tadpoles, as well as observations on calling sites and young. The observations are summarized by species, as follows:
Osteocephalus buckleyi.—No breeding activity was observed. Males were found only at night in March, June, and July. One was perched on aHeliconialeaf in a swamp at Puerto Libre, and two were on bushes in the forest at Santa Cecilia. A gravid female was found on a recently felled tree at Lago Agrio on the night of 12 May 1969.
Osteocephalus leprieurii.—Males were heard calling sporadically at Puerto Libre in July 1968, and at Santa Cecilia in May 1969. A small chorus was found on the night of 12 May 1969 at Lago Agrio, where the frogs were perched on branches of fallen trees over a temporary pool. The call is a soft rattling chuckle. In late April and May many gravid females and males with well-developed nuptial excrescences were obtained from trees as they were felled at Lago Agrio. The reproductive condition of the frogs indicates that they probably breed in May. One individual called nearly every night from a large tree at Puerto Libre between 4-17 July 1968. The tree was felled on the latter date, but no frog was found. Two nights later apparently the same individual called from abromeliadat a height of about 10 m on a large bamboo adjacent to the felled tree; the frog was collected when the bamboo was cut down.
Throughout the rainy months that we have worked in Ecuador (April-August) we have found occasional individuals perched on bushes or low trees at night. Large numbers of adults were observed only during a clearing operation which resulted in the felling of many large trees. Thus, it seems likely thatleprieuriiis a tree-top inhabitant. A partially digested adult male was removed from the stomach of aHemiphractus proboscideus.
At Santa Cecilia many recently metamorphosed young and juveniles were found in June and July 1968. Most of these were on low bushes or herbs in swamp forest at night; some were found in unfoldedHeliconialeaves by day, and one was observed on the forest floor by day. Snout-vent lengths of 18 specimens are 12.3-17.0 (mean 15.1) mm. The smaller frogs were recently metamorphosed as evidenced by the melanophore deposits above the vent. The coloration of the young is strikingly different from that of the adults (see account ofO. leprieurii), so the association of the young and adults was not made until individuals with intermediate patterns were obtained at Lago Agrio in May 1969. Probably juveniles obtained in June and July are the offspring of an April or May breeding. We have been unable to associate tadpoles with this species.
Osteocephalus taurinus.—A small chorus occurred at Lago Agrioon 12 May 1969. Males were calling from the ground adjacent to a small pool amidst recently felled trees. The males were very wary and, when approached, jumped onto limbs and ran up branches; this behavior was noted by Bokermann (1964). The call consists of a series of low-pitched, short notes—like a slow trill—four to six notes per call group. Call groups are repeated two, three, or four times followed by a lapse of several minutes. Although no amplectant pairs were found, several gravid females were collected at Lago Agrio in May, so it can be safely assumed that the species breeds in May. From April through July occasional individuals were observed on bushes and trees at night. During clearing operations at Lago Agrio several individuals were obtained from the tops of trees as they were felled.
Osteocephalus verrucigerus.—Observations were made in a broad, shallow ravine, in which there was a small stream. On 2-4 August 1968, males were observed calling from low bushes and rocks at the edge of a quiet pool in the stream. The call consists of a series of well-pulsed, low-pitched, guttural notes produced at the rate of 5-10 per minute. One amplectant pair was found at the base of a bush adjacent to the pool on 3 August. Another female was found on a branch of a tree 2 m above the ground and 10 m from the stream. Tadpoles of this species were found in the quiet silt-bottomed pool.
SPECIMENS EXAMINED
The localities for each of the specimens examined are given in the following paragraphs. The arrangement of the data is as follows: alphabetically by country, state (department or province), and locality; alphabetically by the first letter in the abbreviations for the museums, and numerically after each museum abbreviation. Specimens lacking precise locality data are listed first in the most restricted political unit possible; localities which have not been found on maps or the positions of which are not known to us are given in quotation marks. Where more than one specimen is included under one museum number, the number of specimens is given in parentheses after the museum number. Unless noted otherwise, all specimens are alcoholics.
Osteocephalus buckleyi
BOLIVIA:El Beni: Ivón, BMNH 1967.2070-1.Santa Cruz: Buenavista, CM 4333, 4339, UMMZ 66563-5.
BRASIL:Amapá: No specific locality, WCAB 13284.
COLOMBIA:Amazonas: Río Guacaya, USNM 152759.Huila: Acevedo,Río Suaza, FMNH 69702.Nariño: Rumiyacu, FMNH 54756.Meta: Río Guejar, Campamento La Macarena, USNM 152199.
ECUADOR: No specific locality, NHMW 6209, WCAB 35499.Chimborazo: Pallatanga, BMNH 1947.2.13.46; Santiago, FMNH 42529.Morona-Santiago: "Río Santiago" (= Río Zamora), MIZS 2950.Napo: Lago Agrio, KU 126646; Puerto Libre, Río Aguarico, KU 123172; Santa Cecilia, AUM 8138, KU 105208-9, 109506, 123171.Pastaza: Alpayacu, BMNH 1912.11.1.64; Canelos, BMNH 1947.2.13.40-1, 1947.2.13.43-5; Colonia Mena, Río Conambo, ZSM 33/1962; Don Tomás, USNM 166014; Guaché, Río Pastaza, AMNH 79986; Río Bobonaza, USNM 166005; Río Capahuari, USNM 166554; Río Conambo at Río Shiona-yacu, USNM 166018; Río Copataza, upper Río Pastaza, USNM 166007-13; Río Pastaza, NHRM 1946; Río Pucyacu, USNM 165997 (skeleton), 165998-6001; Río Rutuno, USNM 166006; Río Villano, USNM 166002-4; Sarayacu, BMNH 1947.2.13.36-9, MCZ 26090, ZMB 10166.
GUYANA:Mazaruni-Potaro: Kartabo, AMNH 70971; Membaru River, upper Mazaruni River, UMMZ 85168; Oko Mountains, FMNH 26722-3.North West: Amakura River, Haulover, UMMZ 83558-9.Rupununi: Marudi River, AMNH 46233; Shudi-kar-wau, AMNH 49252.West Demerara: Dunoon, UMMZ 52449, 52508.
PERÚ:Junín: Chanchamayo, BMNH 1911.12.13.79-80.Loreto: Andoas, AMNH 79984-5; Cashiboya, AMNH 43454; San Antonio, Río Itaya, AMNH 43218.Puno: Yahuaramayo, BMNH 1913.2.25.7.
SURINAM:Suriname: Powakka, CM 44217.
SOUTH AMERICA: No specific locality, NHMW 6208.
Osteocephalus leprieurii
BRASIL:Acre: Tarauacá, FMNH 83247.Amazonas: Rio Javarí, Benjamin Constant, CAS-SU 12620; Río Uaupés, north of Rio Japú, NHMG 489.
COLOMBIA:Amazonas: Gino-goje, lower Río Apoporis, MCZ 28038, 28040-2, 28044, USNM 152136-8.
ECUADOR: No specific locality, WCAB 35452-3; "Napo-Pastaza," USNM 166571.Napo: Avila, UMMZ 92093; south slope Cordillera del Dué, KU 123170; Lago Agrio, KU 125961-2 (skeletons), 126611-44, UMMZ 129326 (2); Limón Cocha, Río Napo, KU 99210-6, UIMNH 63087-9, 63098, 63106-9, 63118-9, 64802-4, 64858, 87998-9, 88001-30, 88437-8, 88580, 88604-5, 89852-97, 89999-90000; Loreto, CAS-SU 11439, WCAB 36526; Puerto Libre, Río Aguarico, KU 123190-1; Puerto Napo, UIMNH 55818-20; Río Cotapino, UMMZ 92094; Río Napo, UMMZ 92078; Santa Cecilia, AUM 8099, 8102, 8113-5, 8127-9, 8131, 8137, 8139-46, 8148, KU 105210-20, 109509-11, 111971, 122964-87, 123169, 126645.Pastaza: Canelos, BMNH 1947.2.13.42, KU 120915; Río Alpayacu, UMMZ 92079; Río Arajuno, USNM 166560-2, WCAB 40176; Río Oglán, USNM 16655203, 166558; Río Rutuno, USNM 166559; Río Shilcayacu, below Puyo, USNM 166557; Río Villano, USNM 166551.
FRENCH GUIANA: No specific locality, MNHN 4629.Inini: Lunier River, MNHN 98/217.
GUYANA:Mazaruni-Potaro: Kartabo, AMNH 70967-8, 70972, 70976.Rupununi: Shudi-kar-wau, AMNH 49255.West Demerara: Demerara Falls, BMNH 72.10.16.23, 72.10.16.37-8.
PERÚ:Loreto: Estirón, Río Ampiyacu, MZUSP 31033-4; Pebas, CAS-SU 3158, 3160; Roaboya, AMNH 43064.
SURINAM: No specific locality, MCZ 2036, RMNH 11468.Marowijne: Camp 3, RMNH 13045-6; Wane Creek North, RMNH 11469-70.Saramacca: Right Coppename River, RMNH 11467.
Osteocephalus pearsoni
BOLIVIA:El Beni: upper Río Beni, below mouth of Río Mapiri, MCZ 15565, UMMZ 57548, 67464-5; Rurrenbaque, UMMZ 57533.
PERÚ:Pasco: Yaupi, KU 136312.
Osteocephalus taurinus
BOLIVIA:El Beni: Ivón, BMNH 1967.2040; Reyes, UMMZ 57532.La Paz: San Ernesto, Mapiri District, BMNH 1901.8.2.54.Santa Cruz: Buenavista, AMNH 33951-2, 33958, BMNH 1927.8.1.19, 1927.8.1.118, FMNH 27091, UMMZ 63319-21, 63959(2), 63961(2), 66566(2), 66567, 66568(2), 66569 (2), 66570, 66571(2), 66575-6, 68196; Río Mamore, 2 km N Boca Chaparé, AMNH 79324; Sara, CM 3840-1; Surutu, CM 3814-5.
BRASIL: No specific locality: "Interior," BMNH 74.7.16.8-9.Acre: Plácido de Castro, MZUSP 6518; Tarauacá, WCAB 2496.Amazonas: Cucuí, NHMW 16495; Manacapurú, ZMB 28492, ZSM 278/1925; Manáus, MCZ 56281, NHMW 16492; Maués, AMNH 69623, 76177; Taracuá, NHMG 488, WCAB 18463-4.Mato Grosso: Mabuca, MZUSP 4272; Posto Coluene, Rio Xingú, WCAB 812; "Puerto Cabello," AMNH 3154; Tapirapé, AMNH 73647-62, CAS-SU 12351-6, MNHN 46/324.Pará: No specific locality, MPEG 623-6; Belém, KU 129866; Cachimbo, FMNH 175876, UIMNH 42149, WCAB 813; Cametá, NHMW 15892; Gurupá, BMNH 96.6.29.13; Ilha de Marajó, BMNH 1923.11.9.20-4; Ilha Mexicana, ZSM 111/1911, 112/1912; "Ponto Dois Indios," BMNH 1939.1.5.5; Santarém, BMNH 75.10.22.1-4, MCZ 354.Rondonia: Abuná, CAS 49773-4, FMNH 64239; Forte Principe da Beira, WCAB 10230; Igarapé Marmelo, FMNH 140254, KU 84725 (skeleton), 92243-6, 92247-8 (skeletons), WCAB 9997, 10001, 10003-4; Porto Velho, MZUSP 16343.
COLOMBIA:Amazonas: Gino-goje, lower Río Apoporis, USNM 152139; Leticia, USNM 152010-1; Raudal de la Playa, lower Río Apoporis, MCZ 28050; Río Apoporis, MCZ 28060.Boyacá: Sutatenza, USNM 152054-6.Cundinamarca: Medina, MCZ 16269-71, USNM 152089-90, 152092-7, 152757.Meta: El Mico, Río Guejar, USNM 152203; Río Duda, Sierra de Macarena, AMNH 79914; Río Guapaya, Sierra de Macarena, FMNH 81332; Río Guaviari, Casa de Piedra, UTA No number.Putumayo: Río Mecaya, FMNH 69711-4, 69716.Vaupés: Gomogoje, lower Río Apoporis, MCZ 28048.
ECUADOR: No specific locality, WCAB 35451, 35785; "Oriente," UMMZ 90418.Carchí: below Salinas, USNM 166059.Esmeraldas: Carondelet, UIMNH 53560-9; Lagartera, Río Caoni, UIMNH 53441, 53458-79.Morona-Santiago: Macuma, UIMNH 63142-3, 63145, 63147, 63151, 63154, 63157, USNM 166060.Napo: Avila, UMMZ 92077; Cuyabeno, UIMNH 63158, 90111; Lago Agrio, KU 126647-55; Limón Cocha, Río Napo, AUM 8132-4, KU 99207-8, 99421-3, 99424 (skeleton), 99425, UIMNH 64801, 87798, 87800, 88032-5, 88576, 90066, 90082, 90102, 90104, 90314, 90984; Loreto, WCAB 35352; Río Cotapino, UMMZ 92080; Río Napo, UMMZ 84120; San José Abajo, AMNH 1295, 1449, 22180, 79990; Santa Cecilia, AUM 8117, 8150, KU 105230-3; south slope Volcán Sumaco, USNM 166570.Pastaza: No specific locality, ZSM 31/1956; Arajuno, USNM 165995; Bufeo, lower Río Bobonaza, USNM 166046-8; Canelos, BMNH 80.12.5.179, 1947.2.13.48, UMMZ 89066;Don Tomás, Río Bobonaza, USNM 166049-50; Montalvo, CAS-SU 10320, USNM 165987-9, 166058, 166566; 2.5 km SE Puyo, USNM 166051; Río Arajuno, USNM 166043-5; Río Arajuno (headwaters), USNM 166053; Río Bobonaza, WCAB 3613-4, 35504; Río Capahuari, USNM 165990, 166555-6; Río Capahuari (headwaters), USNM 166057; Río Conambo, USNM 166569, ZSM 28/1962, 35/1962; Río Conambo at Río Ollaguanga, USNM 166568; Río Conambo at Río Shiona-yacu, USNM 166019, 166563-5; Río Corrientes, USNM 195994, 166020-38, WCAB 3841-2; Río Huiyo-yacu, Pico de Conambo, USNM 166052; Río Pastaza, MCZ 19697; Río Pastaza (drainage), NHRM 1966, USNM 165996; Río Pindo, USNM 166039-41; Río Pindo at Río Tigre (village), USNM 165992-3, 166042; Río Pucayacu, USNM 166054, 166056; Río Rutuno, USNM 166055; Río Solis, upper Río Bobonaza, WCAB 39914; Río Villano, USNM 165991, 166567; Sarayacu, BMNH 80.12.5.213, 80.12.5.239-40, MZUSP 323; Shell Mera, KU 99420.Zamora-Chinchipe: "Yani-Inzari," AMNH 43259, 43394; Zamora, AMNH 78928.
FRENCH GUIANA:Cayenne: Crique Grégoire, UP 40; Maripa, Oyapok River, UP 72; Oyapok River, UZM 1473.Inini: Crique Gabrielle, UP 118-20.
GUYANA: No specific locality: RMNH 1873(3), ZMB 3102(2).East Demerara: Atkinson Field, ASU 11622.Mazaruni-Potaro: Chinapora River, upper Potaro River, BMNH 1905.11.1.20-1; Kamakusa, AMNH 21416, 21418-9, 21422; Kartabo, AMNH 11689, 11691, 11697-9, 11703, 11706-8, 23107, 39730, 70966, 70969-70, 70973-5, USNM 118057; Moraballi Creek, Essequibo River, BMNH 1930.10.10.47-51; Oko Mountains, FMNH 26692-705; upper Potaro River, Tung District, BMNH 1905.11.1.40; Rockstone, FMNH 26591.North West: Amakura River, Haulover, UMMZ 83735.Rupununi: north of Acaray River, west of New River, KU 69747-8; Kuyuwini Landing, AMNH 46283; Pakaraima Mountains, BMNH 1933.6.19.49; Shudi-kar-wau, AMNH 10665, 39637, 49256(2).West Demerara: Demerara, CAS 54773-4; Demerara Falls, BMNH 72.10.16.16-22, 72.10.16.25-32; Dunoon, MCZ 4834, UMMZ 46736, 52493-4, 52502, 52504-5, 57271; Vryheid, BMNH 78.12.13.18.
PERÚ:Amazonas: Río Cenepa, AMNH 43400.Huanuco: Monte Alegre, Río Pachitea, AMNH 43014, 43019.Loreto: Achinamisa, Río Huallaga, AMNH 42178, 42502; Andoas, Río Pastaza, AMNH 79991; Cashiboya, AMNH 43388, 43453; Estirón, Río Ampiyacu, CAS 93264-74, 93276, 93278-9, 93281, 93283-6, 93289, 93311, 93327; Igarapé Champuia, upper Río Curiuja, MZUSP 10339; Iquitos, AMNH 42204, 42442, 43468, NHMW 6118; Lago de Miraño, mouth of Río Napo, AMNH 42712, 43186; Nauta, ANSP 11399; Ollanta, AMNH 42865; Pampa Hermosa, Río Cushabatay, AMNH 43124, 43146; Pebas, CAS-SU 6375; Pucallpa, MJP 101(2), 140(3); Punga, Río Tapiche, AMNH 43194; "Rancho de Indiana, Iquitos District," MVZ 16890; upper Río Abujao, AMNH 42908; Río Itaya, AMNH 42755; upper Río Pisqui, AMNH 43536; Río Tapiche at Río Contaya, AMNH 42983; Río Utoquinia at Brasilian frontier, AMNH 43137; Sobral, Río Tamaya, AMNH 43242; Yurimaguas, BMNH 84.2.18.50.San Martín: Cainarachi, AMNH 42763; Moyobamba, ZSM 19/1914.
SURINAM: No specific locality, BMNH 70.3.10.67, NHMW 18433.3.Brokopondo: Afobaka, RMNH 16536; Brownsweg, RMNH 16537; Railway km. 121, RMNH 16534.Marowijne: Djai Creek, RMNH 16513-4; Maroni River, ZMB 8240, 8531; Nassaugebergte, RMNH 16517-33; Paloemeu, USNM 159025; Swamp Camp, RMNH 16515.Nickerie: Sipaliwini, RMNH 16538.Saramacca: Left Coppename River, RMNH 16535; Tibiti, RMNH 16516.Suriname: Berlijn, RMNH 15064; Powakka, CM 44226; Zanderij, CM 50568.
VENEZUELA:Amazonas: Cerro Duida, UPR-M 2875; Cerro Marahuaca, UPR-M 114-5; Esmeralda, AMNH 23174; Iniridi, SMF 2640; La Culebra, MCZ 28572, UPR-M 117; Laguna, between Tama Tama and Esmeralda, UPR-M 2760; Río Pescado, AMNH 23177; Tapara, UPR-M 113.Distrito Federal: Caracas, BMNH 51.7.17.182.
Osteocephalus verrucigerus
COLOMBIA:Huila: Acevedo, Río Suaza, FMNH 69709-10.
ECUADOR: No specific locality, ZMB 16589.Napo: Avila, UMMZ 90413; south slope Cordillera del Dué, KU 123176-88, 123189 (skeleton), 124208 (eggs), 124209-11 (tadpoles); L'Alegria, USNM 167472-3; Río Pacayacu, tributary of Río Cotapino, CAS-SU 13150; southeast slope Volcán Sumaco, CAS-SU 11442.Pastaza: Abitagua, CAS-SU 5067, FMNH 25791, 27619, UMMZ 90414, 92092; Alpayaca, Río Pastaza, BMNH 1912.11.1.64; Mera, UMMZ 90412(4).Tungurahua: Baños, NHRM 1960.
PERÚ:Ayacucho: La Mar, Sivia, Río Apurimac, FMNH 39853.Huanuco: Río Pachitea, midway between Puerto Victoria and Puerto Inca, CAS-SU 17745.Junín: Satipo, MJP 38.
LITERATURE CITED
Andersson, L. G.
1945. Batrachians from east Ecuador collected 1937, 1938 by Win. Clarke-MacIntyre and Rolf Blomberg. Arkiv Zool., 37A(2):1-88.
Bokermann, W. C. A.
1964. Field observations on the hylid frogOsteocephalus taurinusFitz. Herpetologica, 20:252-255.
1966. Lista anotada das localidades tipo de anfíbios Brasileiros. São Paulo, 183 pp.
Boulenger, G. A.
1882. Catalogue of the Batrachia Salientia s. Ecaudata in the collection of the British Museum, ed. 2, London, xvi+503 pp.
Cochran, D. M.andC. J. Goin
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Transcriber's NotesExcept for the list of corrections below and minor corrections not listed, the text presented here is that of the original printed version.Typographical CorrectionsPageCorrection5is ⇒ in5buckley ⇒ buckleyi17scaral ⇒ sacral19Provicia ⇒ Provincia25in ⇒ is25metalic ⇒ metallic41bromeiad ⇒ bromeliad
Transcriber's Notes
Except for the list of corrections below and minor corrections not listed, the text presented here is that of the original printed version.
Typographical Corrections