Search

Verdade, Vanessa K., Almeida-Silva, Diego, Rodrigues, Miguel T. (2023): The endotrophic nidicolous tadpole of Cycloramphus eleutherodactylus (Miranda-Ribeiro) (Anura: Cycloramphidae). Zootaxa 5254 (2): 287-294, DOI: 10.11646/zootaxa.5254.2.8, URL: http://dx.doi.org/10.11646/zootaxa.5254.2.8

Barcelos, Lucas A., Verdade, Vanessa K. (2022): A new fossil of Ceratophrys Wied-Neuwied (Ceratophryidae: Anura) from Southeastern Brazil. Zootaxa 5094 (3): 495-500, DOI: https://doi.org/10.11646/zootaxa.5094.3.9

Osteocephalus helenae: BOLIVIA: Santa Cruz: Buena Vista: MZUSP 95529. BRAZIL: Acre: Catuaba (S 10 ��04'00''; W 63 �� 37 '00''): SMNS 14190 -1, 17502, 17520. Amap��: Rio Marac��, boca Igarap�� Camaip��: MZUSP 99990-99999; Seringal Boa Fortuna: MZUSP 74253. Amazonas: Reserva Adolpho Ducke: MZUSP 59516, 74253, 75648, 84581; Reserva INPA- WWF (Rio Preto): MZUSP 58055. Rond��nia: Cachoeira de Nazar��, Rio Machado: MZUSP 63805. UHE Samuel: MZUSP 76921. GUYANA: Cuyuni-Mazaruni: Kartabo: AMNH H- 13492. Demerara-Mahaica: Marudi Creek: AMNH A- 46233. Essequibo: Shudikar-wau: AMNH A- 49252. Essequibo Islands-West Demerara: Dunoon: UMMZ 52681 (holotype). Potaro-Siparuni: Iwokrama Forest: MTD 48360 - 2; Magdalen's Creek, NW bank of Konawaruk River, ca. 40 km WSW Mabura Hill: AMNH A- 166220 -4, 166229. VENEZUELA: Amazonas: SW sector of Cerro Yapacana: AMNH-A 100598., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 181, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

Osteocephalus camufatus sp. nov. (Figs. 1���4) Osteocephalus buckleyi [Ca 1 _ MTR 12779 _ 2748]��� Jungfer et al. 2013 Holotype: MZUSP 142389 (field number MTR 12779), an adult male of 39.5 mm SVL with well-developed nuptial pads from Brazil: Amazonas: Igarap��-a��u (04�� 20 ' 40 ''S 58 �� 38 '06''W), right bank of Rio Abacaxis, 30 m asl., collected by Miguel T. Rodrigues, Sergio Marques de Souza, Jos�� Cassimiro and Jos�� M��rio Guellere, on 9 January 2007 (Figs. 1���4). Paratype: MZUSP 157020 (field number MTR 13147), an adult male with well-developed nuptial pads from Brazil: Amazonas: Areal, left bank of Rio Abacaxis (04�� 35 ' 49 ''S 58 �� 13 ' 14 ''W), 39 m asl., collected by Miguel T. Rodrigues, Sergio Marques de Souza, Jos�� Cassimiro and Jos�� M��rio Guellere, on 20 January 2007. Diagnosis and comparisons. Osteocephalus camufatus may be diagnosed as (1) a small to medium sized species (as defined by Jungfer 2010: 29) in males (females unknown); (2) skin on dorsum of males shagreen with a few irregular small tubercles, only some bearing keratinized tips; (3) skin on flanks coarsely areolate between limb insertions; (4) rounded, tuberculate canthus rostralis curved inward; (5) frontoparietal ridges not visible externally; (6) dentigerous processes of vomers angular; (7) thick tuberculate supratympanic fold from the posterior edge of the orbit sloping in an arch towards the arm insertion, not reaching below tympanum posteroventrally; (8) webbing on inner edge of third finger extending slightly beyond penultimate subarticular tubercle; (9) distal subarticular tubercle on Finger IV bifid; (10) dorsum boldly blotched irregularly; (11) throat, chest and venter creamy tan to creamy white with numerous irregular tan spots; (12) supralabial area with irregular light and dark markings; (13) flanks light with irregular dark markings; (14) vocal sacs paired, protruding ventral to angles of jaws; (15) juvenile colouration unknown; (16) tadpole habitat (most likely in pools along streams) and labial tooth row formula unknown; (17) colour of tibiofibular bones white in preservative. O. camufatus is a member of the O. buckleyi species group as revealed by molecular data. A phylogenetic analysis using both maximum parsimony (MP) and likelihood (ML) inference resulted in trees that showed that O. camufatus was basal to a large and well-supported clade containing frogs from large areas of Amazonia and Guiana. The oldest available name for this clade is O. helenae (Jungfer et al. 2013). Uncorrected p distances of the mitochondrial 16 S gene between O. camufatus and O. helenae from various localities were 1.3���2.6, while genetic divergences within O. helenae, whose subclade relationships are not yet sufficiently resolved, were 0.0��� 2.2 (Jungfer et al. 2013: Appendix S 4 d). The new species shares with several other species of the group the following character states: 1) greenish ground colour, 2) strongly tuberculate canthus rostralis and upper eyelids and 3) posterior surfaces of thighs light with large dark brown spots in preservative. These three characters distinguish it from all frogs in the O. alboguttatus, O. leprieurii, O. planiceps and O. taurinus groups (Jungfer et al. 2013) in which the ground colours are shades of tan or brown, the canthus rostralis is smooth to granulate and the posterior thigh surfaces are uniform. From other species in the O. buckleyi group it differs as follows (characters of O. camufatus in parentheses): Six upland species from the eastern Andean slopes between 600 and 2300 m differ as follows: in Osteocephalus carri, a predominantly brown species (bold blotches of green and brown), the canthus rostralis and upper eyelids are smooth (strongly tuberculate) and the iris is black with golden spots in life (light); O. duellmani is a tan species with or without dark brown blotches (bold blotches of green and brown), with the posterior surfaces of the thighs uniform tan (marbled) and the canthus and upper eyelid bearing a few low tubercles (strongly tuberculate); O. festae exhibits brown ground colour (bold blotches of green and brown), a dark brown iris in life (light, Fig. 4) and uniform tan posterior thigh surfaces (marbled). Males of Osteocephalus mimeticus, O. mutabor (the latter also occurring as low as 150 m asl.) and O. verruciger have a brown ground color lacking green (blotches in shades of green and brown), uniform posterior thigh surfaces (marbled dark brown), and heavily tuberculate dorsa with numerous spinous tubercles bearing keratinized tips during breeding (a few irregular small tubercles, few of them with keratinization). Species of the O. buckleyi group from the lowlands (O. camufatus, differ as follows: Osteocephalus buckleyi has low tubercles or is smooth on the upper eyelid (strongly tuberculate), and has low or lacks tarsal tubercles (prominent). Breeding males of O. buckleyi exhibit dorsal tubercles, many with keratinized tips (few keratinized tips). Osteocephalus cabrerai has a row of tubercles on the lower jaw (absent) and an irregular, deep fringe on the outer edge of Finger IV (low tubercles on proximal segment of Finger IV). Osteocephalus cannatellai from lowlands, but also up to 1290 m asl. (Ron et al. 2012), appears to be larger (38.5���57.2 mm SVL in males, mean, 46.8 mm, Ron et al. 2012) (to 40.4 mm, but note that the sample size is n= 2), and differs by a longer supratympanic fold that reaches the arm insertion (not reaching lower level of tympanum); axillary membrane present (absent); usually a dark venter varying from light grey to brown with or without dark markings (Ron et al. 2012) (yellowish white with small irregular brown spots). Osteocephalus helenae (Ruthven) is a variable frog throughout its range. Specimens from localities closest to the two sites known for O. camufatus, from about 230 km northwest of the type locality, near Manaus and other sites just north and south of the Amazonas in Amazonas, Brazil, exhibit an axillary membrane (absent), a scalloped fringe on the outer edge of Finger IV continued to the ulna (low, indistinct tubercles on proximal segment of Finger IV and single tubercles on ulna), and more webbing on the hand, reaching beyond the distal subarticular tubercle on Finger IV (to distal subarticular tubercle, Fig. 4 a). Specimens from near Rio Branco, Acre, Brazil, roughly 1100 km WSW of the type locality of O. camufatus, have posterior thigh surfaces brown, finely mottled with cream (bold dark brown markings on light ground) and lack tubercles on the eyelids (present). Frogs in the closely related genus Dryaderces are similar in general appearance to some frogs of the O. buckleyi group, but can be distinguished as follows: They differ in exhibiting bold dark ventral reticulation (dark spotting) and uniform dark iris (light iris) in D. pearsoni (Gaige), a longer snout of EN/HL 0.27���0.33 (shorter, EN/ HL 0.25), non-tuberculate canthus rostralis (tuberculate) and light, uniform venter (venter with small dark spots) in Dryaderces sp. (Dryaderces pearsoni [Ca 1 _ MTR 13158 _ 2768] of Jungfer et al. 2013) (KHJ, unpublished data). The inspection of a fresh specimen of ��� Osteocephalus ��� inframaculatus (Boulenger), a species until recently only known from the holotype and tentatively placed with the O. buckleyi group by Jungfer (2010), was placed in Dryaderces by Hoogmoed (2013). It is easily distinguishable from O. camufatus by its bold dark markings on throat and chest and has a longer snout with EN/HL 0.33 (0.25) and larger tympanum with TD/HL 0.26 (0.22��� 0.23). Description of holotype. Snout short (EN/HL 0.245) and blunt, head as long as wide. Nostrils raised, opening laterally. Snout truncate in dorsal and lateral outline. Canthus rostralis well-developed, rounded, describing an inwardly curved line, bearing large tubercles. Skin of head coarsely shagreen, orbits with a few additional small tubercles, some bearing small keratinized tips. A few more scattered keratinized tips on some tubercles of the head and anterior part of the body. No frontoparietal crests evident externally. Loreal region strongly concave bearing low tubercles. Lips flared below the eye. Choanae oblique, elliptical, though slightly angular. Dentigerous processes of vomers short, considerably shorter than choanae width, angular, not in contact with each other, bearing 4 teeth on the left and 3 on the right. The anterior edges of the dentigerous processes are in line with the posterior third of the choanae, their posterior edges well behind the posterior margins of the choanae. Tongue elliptical, slightly wider (by 6 %) than long. The vocal sacs are paired, subgular and protrude ventral to the jaw articulation. A thick glandular supratympanic fold from the posterior part of the orbit medially, covering tympanic annulus dorsally from ��� 11 ��� (when tympanum is thought to be a clock face) to ��� 3 ���, then sloping in a curve towards the arm insertion, reaching no further than the lower one third level of the tympanum. The latter is conspicuous, rounded (barely wider than high), with about 66 % the diameter of the eye. Skin on dorsum coarsely shagreen with a few low, irregularly spaced tubercles. Transversal furrows on the shoulders are absent. Skin smooth between eye and tympanum and areolate posterior to the tympanum above the arm insertion, very coarsely areolate laterally between the limb insertions. Throat and belly areolate. Cloacal opening situated at about half the level of thighs, surrounded by large, irregular warts (except on the cloacal flap, which is smooth). Skin on arms shagreen with a few faint tubercles dorsally, hidden surfaces finely shagreen. A row of low ulnar tubercles also encompasses the proximal segment of Finger IV. Axillary membrane absent. Dorsal surfaces of hind legs shagreen, posterior surfaces of thighs and ventral parts of shank smooth, proximal three fourths of thigh coarsely areolate ventrally. A row of low outer tarsal tubercles from the heel to the proximal subarticular tubercle of Toe V. On the heel, three and four low tubercles, respectively, directed dorsally and laterally. Finger and toe discs elliptical and wider than long. Diameter of disc on Finger III is 74 % of tympanum diameter. Relative finger length is I I basal II 1 ����� 2 �� III 2 ⅓��� 2 IV. On the foot a large elliptical inner metatarsal tubercle and a small rounded plantar tubercle. A larger rounded conical outer metatarsal tubercle in line with the row of tarsal tubercles. The latter form a shallow fringe on the distal part of the proximal toe segment that is continued to the toe disc. A few indistinct supernumerary tubercles on the proximal segments of Toes III���V. Subarticular tubercles single and conical. Relative lengths of appressed toes I I 1���2 - II 1���2 III 1 ��� 1 + IV 1 +��� 1 V. Measurements and proportions. SVL 39.5; HL 14.3; HW 14.3; TL 22.2; FL 26.8; ED 4.7; TD 3.1; FD 2.3; EN 3.5; IN 3.4; TE 2.0. HL/SVL 0.36; HW/SVL 0.36; TL/SVL 0.56; FL/SVL 0.68; TD/ED 0.66; TD/FD 1.35; EN/ HL 0.25; EN/SVL 0.09; HL/HW 1.00; TE/TD 0.65; TD/HL 0.22. Colour in preservative. The ground colour of head and dorsum is light (green in life) with numerous, more or less irregular blotches in tan and dark brown (various shades of brown in life). This pattern extends to the lateral sides of the head and body. One ill-defined dark brown mark somewhat more conspicuous than others has two arms between the eyes (including the orbits), two between the tympana and on midbody the posterior two arms diverging like the lower part of an X and are continued on the flanks. Two additional ill-defined dark brown marks posteriorly to the proximal part of the sacrum, continued to the flanks. Throat and belly with small irregular brown spots. Although there are light areas on the upper lip, there are no discrete labial marks distinguishable from the rest of the pattern. Tympanum tan. Arms dorsally and laterally with irregular dark brown crossbars, with irregular brown spotting ventrally. Legs dorsally with irregular dark brown crossbars, posterior surfaces of thigh and calf marbled dark brown. Ventral surfaces of thigh tan with lighter areolae, of femur tan peppered with darker markings. Webbing tan, slightly marbled. Subcloacal warts are in creamy white and different shades of tan. Colour in life. Colours were taken from photographs (Fig. 4). Dorsal surfaces covered by bold blotches of different shades of green and brown. A light mark posteroventral to the eye is light green, the posterior part of the upper lip dark brown, of the lower lip white. The tympanum is tan. The iris is light tan in its upper half and creamy white in its lower one, the latter bearing a diffuse dark brown median vertical streak. Upper and lower halves are separated by a weakly demarcated reddish brown horizontal streak and are both finely reticulated black. Some light blue coloration can be seen on the hidden surface of the thigh. Outer tarsal and heel as well as ulnar tubercles are tipped white. Variation in the paratype. An adult male slightly larger than the holotype (40.4 mm SVL) with welldeveloped nuptial pads. The snout is bluntly rounded in dorsal aspect. The dentigerous processes bear five teeth on each side. Keratinized tips on tubercles are lacking entirely. Coloration is similar, but the large tan dorsal mark is less conspicuously outlined. The belly is spotted evenly. There is slightly more webbing on the hand with I basal II 1 2 / 3 ��� 2 2 / 3 III 2 ��� 2 IV and little variation on the foot: I 1 +��� 2 II 1���2 - III 1 ��� 1 1 / 3 IV 1 1 / 3 ��� 1 V. There is also little variation in proportions. Measurements and proportions. SVL 40.4; HL 15.0; HW 14.9; TL 21.2; FL 25.9; ED 5.0; TD 3.4; FD 2.1; EN 3.7; IN 3.3; TE 1.9. HL/SVL 0.37; HW/SVL 0.37; TL/SVL 0.53; FL/SVL 0.64; TD/ED 0.68; TD/FD 1.62; EN/ HL 0.25; EN/SVL 0.09; HL/HW 1.01; TE/TD 0.56; TD/HL 0.23. Distribution and habitat. The new species is so far only known from two sites about 54 km apart along the terra firme forests of the right bank of the Rio Abacaxis, a black water river south of the Amazonas that discharges into Paran�� do Arari��, a white water channel delimiting the southern border of Ilha de Tupinambarana (Fig. 5). This is a huge island (about 300 km long) extending between Nova Olinda do Norte on the lower Rio Madeira to Parintins on the Amazonas. Habitat at Igarap��-a��u consisted of typical terra firme forest with higher trees reaching up to 40 m and a diameter at breast height of around 80���100 cm. The understory was particularly rich in low palm trees and the leaf litter was dense. The specimen was obtained close to a small stream perched on a tree about 1 m high. At Areal, a site further upstream (also referred to as Paca or Pacamiri locally) the local habitat consisted of an extensive white sand area (campinarana) with scattered vegetation near the margin of the river with abundant clumps of ground bromeliads, gradually replaced by denser areas covered by palm trees and then terra firme forest. The terra firme forest with large trees became more evident farther from the river margin, where the second specimen was obtained. Etymology. The species name camufatus is the Latinized past participle of Italian camuffare, to disguise or mask, from which the word camouflaged is derived, in allusion to the shape-dissolving pattern of greens and browns of the new species., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on pages 172-178, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624, {"references":["Jungfer, K. - H., Faivovich, J., Padial, J. M., Castroviejo-Fisher, S., Lyra, M. M., Berneck, B. V. M., Iglesias, P. P., Kok, P. J. R., MacCulloch, R. D., Rodrigues, M. T., Verdade, V. K., Torres Gastello, C. P., Chaparro, J. C., Valdujo, P. H., Reichle, S., Moravec, J., Gvozdik, V., Gagliardi-Urrutia, G., Ernst, R., De la Riva, I., Means, D. B., Lima, A. P., Senaris, J. C., Wheeler, W. C. & Haddad, C. F. B. (2013) Systematics of spiny-backed treefrogs (Hylidae: Osteocephalus): an Amazonian puzzle. Zoologica Scripta, 42, 351 - 380 + online appendix. Available from: http: // onlinelibrary. wiley. com / doi / 10.1111 / zsc. 12015 / suppinfo (Accessed 19 May 2016)","Jungfer, K. - H. (2010) The taxonomic status of some spiny-backed treefrogs, genus Osteocephalus (Amphibia: Anura: Hylidae). Zootaxa, 2407, 28 - 50.","Ron, S. R., Venegas, P. J., Toral, E., Read, M., Ortiz, D. A. & Manzano, A. L. (2012) Systematics of the Osteocephalus buckleyi species complex (Anura, Hylidae) from Ecuador and Peru. ZooKeys, 229, 1 - 52. http: // dx. doi. org / 10.3897 / zookeys. 229.3580","Hoogmoed, M. (2013) Rediscovery of the rare tree frog Hyla inframaculata Boulenger, 1882 (Anura: Hylidae), in Amazonian Brazil with notes on variation and distribution, and its generic allocation. Amphibia-Reptilia, 34, 421 - 432. http: // dx. doi. org / 10.1163 / 15685381 - 00002907"]}

Amorim, Dalton S., Santos, Charles Morphy D., Krell, Frank-Thorsten, Dubois, Alain, Nihei, Silvio S., Oliveira, Otto M. P., Pont, Adrian, Song, Hojun, Verdade, Vanessa K., Fachin, Diego A., Klassa, Bruna, Lamas, Carlos José E., Oliveira, Sarah S., De Carvalho, Claudio J. B., Mello-Patiu, Cátia A., Hajdu, Eduardo, Couri, Márcia S., Silva, Vera C., Capellari, Renato S., Falaschi, Rafaela L., Feitosa, Rodrigo M., Prendini, Lorenzo, Pombal Jr, José P., Fernández, Fernando, Rocha, Rosana M., Lattke, John E., Caramaschi, Ulisses, Duarte, Marcelo, Marques, Antonio Carlos (2016): Timeless standards for species delimitation. Zootaxa 4137 (1): 121-128, DOI: http://doi.org/10.11646/zootaxa.4137.1.9

A new species of treefrog of the genus Osteocephalus is described from the Rio Abacaxis, a southern tributary of the Amazonas in the state of Amazonas, Brazil. This member of the O. buckleyi group is characterized by green dorsal colouration with irregular blotches of various shades of brown, light venter with tan spots and bold dark markings on the posterior surfaces of the thighs. It can be distinguished from its closest relative, O. helenae from the same general area, by the lack of an axillary membrane, a few indistinct tubercles on the proximal segment of Finger IV and single ulnar tubercles. Fil: Jungfer, Karl Heinz. Universitat Koblenz. Landau Institut Fur Biologie; Alemania Fil: Verdade, Vanessa K.. Universidade Federal do ABC; Brasil Fil: Faivovich, Julián. Consejo Nacional de Investigaciones Científicas y Técnicas. Oficina de Coordinación Administrativa Parque Centenario. Museo Argentino de Ciencias Naturales ; Argentina. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Departamento de Biodiversidad y Biología Experimental; Argentina Fil: Rodrigues, Miguel Trefaut. Universidade de Sao Paulo; Brasil

Osteocephalus verruciger: COLOMBIA: Caquet��: Municipio de Florencia: Escuela Tarqui, carretera Altamira-Florencia km 48-49: ICN 23648. Municipio de Florencia: 35.2 km. arriba de Florencia: ICN 23943. Municipio de Florencia: Vereda Tarqui 38.8 -39.0 km: ICN 23944 - 5, ICN 23948, ICN 23952, ICN 23954. Municipio de Florencia: Vereda Tarqui 13.2 km arriba de Florencia: ICN 23946. Municipio de Florencia: 39.3 km arriba de Florencia: ICN 23947. Huila: Acevedo, R��o Suaza, R��o Aguas Claras near San Adolfo, 1400 m: FMNH 69709 - 10. Palestina: ICN 0 1542. Putumayo: 10.3 km W El Pepino, 1440 m: KU 169586 -7, 169589-93, 169595-6,169599, 169601 -4, 169606- 7. ECUADOR: No specific locality: ZMH-A 946 (syntype). Cotopaxi: Las Pampas (in error): MHNG 2259.20, 2560.62, 2560.64 - 68. Napo: El Reventador: MHNG 2259.18, 2273.28, 2485.65 - 70, 2560.63. 2 km SSW R��o Reventador, 1700 m: KU 164408, 164414, 164416 -9, 164421, 164423 -4, 164426. 3.2 km NNE Oritoyacu, 1910 m: KU 178839 - 44. R��o Azuela, 1740 m: KU 143210 -2, 143215- 7, 143219-24, 164434, SMNS 14197. R��o Azuela, 9.5 km W of Reventador, 1630 m: KU 217750 - 1. R��o Salado, 1 km upstream from R��o Coca, 1420 m: KU 164437, 164442, 178844, 178846- 7. 0.7 km NE R��o Salado bridge on Lago Agrio road, 1380 m: KU 190054. San Rafael: MHNG 2259.19, 2272.98 - 99. S slope Cordillera del Due, 1150 m: KU 123181, 123186. 11.1 km NE Santa Rosa, 1900 m: KU 19004953. 16.5 km NNE Santa Rosa, 1700 m: KU 143209. Pastaza: 9.5 km NW Mera, 1270 m: KU 178848. Tungurahua: 11 km E R��o Negro, 1170 m: KU 146469 - 70., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 181, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

Osteocephalus camufatus: BRAZIL: Amazonas: Igarap��-a��u, right bank of Rio Abacaxis, 30 m (04�� 20 ' 40 ''S; 58 �� 38 '06''W): MZUSP 142389. Areal, left bank of Rio Abacaxis, 39 m (04�� 35 ' 49 ''S; 58 �� 13 ' 14 ''W). MZUSP 157020., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 180, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

Osteocephalus buckleyi: ECUADOR: Napo: Jatun Sacha: KU 217746, SMNS 13714 - 5. Orellana: Hacienda Primavera, 40 km S Coca: MHNG 2556.58; Yasun�� (S 00�� 41 ���00������; W 76 �� 24 ���00������): SMNS 13713. Pastaza: Canelos: BMNH 1947.2.13.40��� 1947.2. 13.41, 1947.2. 13.43, 1947.2. 13.44 (lectotype), 1947.2. 13.45. Sucumb��os: San Pablo de Kantesiya: MHNG 2366.39���2366.40; Santa Cecilia: KU 105208 ���9, 150492 ���3, 152477, 175500; Shushufindi: MHNG 2560.61. PERU: Loreto: 28 km S Iquitos: MUSM KHJ-F 0 67., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 180, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

Osteocephalus mimeticus: PERU: Ayacucho: San Jos�� on R��o Santa Rosa, 1005 m: KU 196994 ��� 5. Sivia, R��o Apurimac, 760 m: FMNH 39853. Cuzco: Pozo Pagoreni, Comunidad Kirigueti: MUSM 21872. Zona Reservada Nahua Kugapakori: MUSM 23179 ���80, 23188, 23207, 23218, 23226. Hu��nuco: Fundo Flor, R��o Pachitea, 300 m: SMNS 6515. San Mart��n: W slope Abra Tangarana, 7 km NE San Juan de Pacaysapa, 1080 m: KU 212189 ��� 90. Cataratas Ahuashiyacu, 14 km NE Tarapoto, 730 m: KU 212191. R��o Cainarache, 33 km NE Tarapoto on road to Yurimaguas: KU 209454 ��� 5. R��o Cumbaza Valley, 9.4 km N Tarapoto, 390 m: KU 212182 ��� 5. Roque: GNM 469 (lectotype). Pongo de Shilcayo, ca. 4 km NNW Tarapoto, 470 m: KU 212196 ��� 9. 14 km ESE Shapaja, 360 m: KU 212193. 22.7 km NE Tarapoto, 810 m: KU 212200. 28 km NE Tarapoto, 600 m: KU 212201 ��� 3. 29 km NE Tarapoto, 550 m: KU 212181. Tocache, R��o Huallaga: MUSM 10845. Venceremos, 89 km NW Rioja, 1650 m: KU 212186 ��� 7. Pasco: Pozuzo, 770 m: MUSM 20351. Puerto Bermudez: MUSM 17801. Santa Isabel: MUSM 17847 ���52, 18022, 18024. Yaupi, R��o Paucartambo, 1600 m: KU 136312. Ucayali: Boquer��n del Padre Abad: ZFMK 33352, 39614���5, 39748���50, 39752���3, 40152��� 3., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 181, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

Osteocephalus festae: ECUADOR: Morona-Santiago: Valle de Santiago (= lower R��o Zamora): MZUT An. 208 (holotype). Napo: Archidona: MHNG 2560.60. Loreto, ��vila, subcentro Caim��toyacu: EPN 5577 ��� 8. Sucumb��os: Cuyabeno, Campamento Concienti: EPN AA- 5611, AA- 5607. PERU: Cajamarca: San Ignacio: El Sauce: MUSM 19224. San Mart��n: 1 km NW Venceremos, 1600 m: KU 217302. 14 km W Venceremos, 2000 m: KU 217303., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 181, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

Osteocephalus duellmani: ECUADOR: Morona-Santiago: Cordillera del C��ndor, R��o Piuntza, 1910 m asl, about 3 �� 25 'S, 78 �� 27 'W: KU 147171 (paratype), 147172 (holotype)., Published as part of Jungfer, Karl-Heinz, Verdade, Vanessa K., Faivovich, Juli��n & Rodrigues, Miguel T., 2016, A new species of spiny-backed treefrog (Osteocephalus) from Central Amazonian Brazil (Amphibia: Anura: Hylidae), pp. 171-181 in Zootaxa 4114 (2) on page 181, DOI: 10.11646/zootaxa.4114.2.6, http://zenodo.org/record/255624

Haddadus aramunha (Cassimiro, Verdade & Rodrigues, 2008), new comb. Diagnosis. To date, there is no identifiable morphological synapomorphy supporting the monophyly of the genus Haddadus (see discussion). We assign H. aramunha to this genus based on inferred relationships using molecular data, as well as on the overall similarity among males of S. aramunha and H. binotatus. Also, head width (HW)/ snout-vent length (SVL) lower than 49 % is a characteristic distinguishing members of Haddadus from Strabomantis (Cassimiro et al. 2008) (up to 54 % in Strabomantis, Hedges et al. 2008). Haddadus aramunha is characterized by the following combination of traits: (1) largest size for the genus (adult males SVL 37.3–44.9 mm, adult females SVL 51.0– 81.9 mm; Napoli et al. 2010); (2) wide head (HW/SVL 0.40–0.49; HL/SVL 0.37–0.45), its width greater to slightly smaller than its length; (3) adult females bearing welldeveloped frontoparietal crests, delimiting a mid-dorsal longitudinal groove; (4) posterior part of pars fascialis of maxilla deepened; (5) mandibular ramus of the trigeminal nerve passing lateral to the m. levator mandibulae posterior subexternus, (6) finger I much longer (FLI/SVL 0.15–0.23) than finger II (FLII/SVL 0.08 –012); (7) fifth toe smaller than third; (8) dorsum generally smooth, covered with keratin white dots, bearing only one or two granular dorsolateral ridges; (9) vocal sac and vocal slits absent in males; (10) no asperities on thumbs of males; (11) in adult live specimens, dorsal background dark reddish brown. The new combination herein proposed changes Hedges’ et al. (2008) generic diagnosis of Haddadus: (1) head narrower (or broader than body); (3) cranial crests absent (or present and well developed in females); (8) Toe III equal in length, slightly shorter (or clearly longer than Toe V); and (11) range in SVL 17 mm in only known specimen of H. plicifer (82 mm in females of H. aramunha). Comparisons with other species. Haddadus aramunha differs from H. binotatus by always presenting the head wider than long in females, and usually wider than long in males (always longer than wide in H. binotatus); shorter legs (slightly longer than SVL in H. aramunha (thigh + tibia + foot length about 1.05 SVL), and clearly longer than SVL in H. binotatus (thigh + tibia + foot length about 1.15 SVL); conspicuous frontoparietal crests in females, and poorly developed ones in males, delimiting a mid-dorsal longitudinal groove (absent in H. binotatus); Haddadus aramunha also differs from H. binotatus and H. plicifer (species data from Boulenger 1888) by the presence of only one or two granular dorsolateral ridges on dorsum (five symmetrical undulous linear folds on each side of the back in H. binotatus and H. plicifer). The dark reddish brown dorsal background in live adult females of H. aramunha differs this species from H. binotatus, which never presents such a pattern (See Fig. 1 of Napoli et al. 2010). Haddadus aramunha inhabits open areas, living near streams within “campo rupestre” environments, while H. binotatus and H. plicifer are restricted to forested areas of the tropical Atlantic forest.

Made available in DSpace on 2022-04-29T07:12:49Z (GMT). No. of bitstreams: 0 Previous issue date: 2013-11-29 The taxonomic and biogeographic affinities of Strabomantis aramunha from the Campos Rupestres of Brazil are intriguing. A unique skull morphology of females suggest affinities with the broad-headed eleutherodactylines of Northwestern South America in the genus Strabomantis. Male and juvenile morphology nonetheless suggest S. aramunha could be related to members of the recently described genus Haddadus from eastern Brazil. We assess the affinities of S. aramunha using molecular phylogenetic analyses of mitochondrial (12S, tRNAval, 16S, cyt b) and nuclear sequences (RAG-1and rhodopsin). Bayesian inference, likelihood, and parsimony analysis recover a highly supported clade with S. aramunha and H. binotatus as sister taxa. Accordingly, we transfer S. aramunha to Haddadus, and provide a new generic definition of the later. The distribution of species in Haddadus (highlands of the Espinhaço mountain Range and coastal eastern Brazil) is now concordant with the general pattern observed for other species in the area. © 2013 Magnolia Press. Universidade de São Paulo Instituto de Biociências Departamento de Zoologia, Caixa Postal 11.461, 05422-970 São Paulo, São Paulo Universidade Federal do Rio de Janeiro Museu Nacional Departamento de Vertebrados, 20940-040 Rio de Janeiro, Rio de Janeiro Universidade Federal da Bahia Instituto de Biologia Departamento de Zoologia, 40170-115, Salvador, Bahia Universidade Estadual de Feira de Santana Departamento de Ciências Biológicas, 44036-900 Feira de Santana, Bahia Universidade Federal do ABC Centro de Ciências Naturais e Humanas, Avenida dos Estados, n5001, 09210-971 Santo-André, São Paulo Universidade Estadual Paulista Júlio de Mesquita Filho Instituto de Biociências Departamento de Zoologia, 13506-900 Rio-Claro, São Paulo Universidade Estadual Paulista Júlio de Mesquita Filho Instituto de Biociências Departamento de Zoologia, 13506-900 Rio-Claro, São Paulo

Strabomantis aramunha sp. nov. (Figs. 1���3) Holotype: MZUSP 138693 (field number JC- 1212), an adult female, from Serra do Sincor��, Espinha��o range, 13 �� 04' 07" S and 41 �� 20 ' 09" W, 998 m elevation, municipality of Mucug��, State of Bahia, Brazil, collected by J. Cassimiro and F. S. F. Leite on 11 March 2005. Paratypes (N= 6): MZUSP 138687 (JC- 1198), an adult female, 13 �� 00' 33 " S and 41 �� 22 ' 46 " W, elevation 1191 m, on 8 March 2005; MZUSP 138688 (JC- 1206), an adult male, 13 �� 00' 32 " S and 41 �� 22 ' 48 " W, elevation 1207 m, same date; MZUSP 138690 (JC- 1214), an adult male, 13 �� 03' 43 " S and 41 �� 20 ' 26 " W, elevation 947 m, on 11 March 2005; MZUSP 138689 (JC- 1213), juvenile female, 13 �� 04' 07" S and 41 �� 20 ' 09" W, elevation 998 m, same date; MZUSP 138691 (JC- 1251), juvenile male, on 20 March 2005; MZUSP 138692 (JC- 1277), juvenile female, 13 �� 01' 20 " S and 41 �� 21 ' 24 " W, elevation 1068 m, on 26 March 2005. All collected by J. Cassimiro and F. S. F. Leite at the surroundings of the type locality. Variable MZUSP MZUSP MZUSP MZUSP MZUSP MZUSP MZUSP 138693 138687 138688 138690 138691 138689 138692 Etymology: The specific name is from the Tupi language aramunha, meaning giant, in allusion to the large size of the species, and is used as noun in apposition. Diagnosis: A large broad-headed species (adult male 40.3���41.1 mm SVL, adult female 75.2���79.7 mm; HW respectively 44���46 % and 48���49 % of SVL), adult females bearing well-developed frontoparietal crests, posterior part of pars fascialis of maxilla deepened, mandibular ramus of the trigeminal nerve passing lateral to the m. levator mandibulae posterior subexternus, eyelid tubercles present posteriorly, not elongated, skin of venter coarsely areolate, leg relatively short (FL 51���53 % of SVL) finger discs absent, first finger longer than second, poorly developed discs on toes, first and second toes ridged, fifth toe smaller than third, tarsal fold absent, testes white. Comparison to other species: The adult Strabomantis aramunha is promptly diagnosed from all other eleutherodactylines by the combination of large size, broad-head, flared maxilla, frontoparietal bearing welldeveloped crests in females, mandibular ramus of the trigeminal nerve passing lateral to the m. levator mandibulae posterior subexternus, posterior eyelid tubercles present and not elongated, skin of venter coarsely areolate, absence of finger discs, first finger longer than second, poorly developed discs on toes, first and second toes ridged, fifth toe smaller than third, and, tarsal fold absent. These characters place squarely the new species in the Eleutherodactylus sulcatus group (Lynch 1997; Lynch & Duellman 1997), currently in the genus Strabomantis (Hedges et al. 2008). The only other species of Strabomantis occurring in Brazil is S. sulcatus (Lynch 1975, 1997; Frost 2007; Hedges et al. 2008). Strabomantis sulcatus differ from S. aramunha by the smaller size (SVL 28���42 mm; Lynch 1975, 1980), concealed surfaces of thigh and shank presenting light blotches, and by the presence of tarsal fold. From the other species of Strabomantis, S. aramunha differ by the larger size (adult female S. cadenai, SVL 40.6 mm; Lynch 1997), skin of venter areolate (smooth in S. cadenai, S. laticorpus; Lynch 1997), by not presenting elongate conical tubercles on eyelid (present in S. cadenai, S. cerastes, S. ingeri, S. laticorpus, S. necopinus, S. ruizi; Lynch 1997), large flattened warts absent (present in S. helonotus; Lynch 1997), tarsal fold absent (present in S. anomalus, S. cornutus, S. ingeri, S. necopinus, and S. ruizi; Lynch 1997), cranial crests present (absent in S. anomalus, and S. cheiroplethus; Lynch & Myers 1983; Lynch 1990), reduced basal web only between first and third toes (extensive toe webbing in S. anomalus, S. anatipes and S. zygodactylus; Lynch & Myers 1983). Males and the young specimens, by being smaller and presenting narrower heads most resemble the adult Haddadus binotatus (formerly ��� Eleutherodactylus ��� binotatus, Hedges et al. 2008). However, under close examination H. binotatus is promptly differentiated from S. aramunha by the smoother dorsal skin and the presence of tubercles not bearing keratinized tips (present in S. aramunha). Additionally, the head is narrower and longer than wide (wider than long in S. aramunha), the body is slender, the legs are longer, and the supernumerary tubercles are more numerous in H. binotatus than in S. aramunha. Description of holotype: Head wider than long (HW = 36.4 mm; HL = 32.6 mm); a pair of frontoparietal crests externally visible; eyes dorsolateral, median sized (ED = 9.9 mm), diameter similar to the eye-nostril distance (EN = 10.0 mm); snout sub-elliptical in dorsal view, rounded in lateral view, overhanging lower jaw; nostrils small, slightly protuberant, directed laterally and positioned distally at the canthus rostralis; canthus rostralis sharp, accentuated by bony keel; loreal region deeply concave, tuberculate; flared maxilla, with posterior pars fascialis deepened; upper eyelid bearing small tubercles posteriorly, not elongated; interorbital space highly depressed, bearing small tubercles; tympanic membrane smooth; tympanic annulus prominent, higher than long (TYD = 4.5 mm; TYH = 5.5 mm), oblique; tympanum-eye distance less than tympanum diameter (TED = 3.6 mm); supratympanic fold evident, extending from posterior corner of eye to axils. Skin of dorsum spiculate under magnification, with tubercles more concentrated on posterior half; dorsolateral ridges present; skin of venter coarsely areolate; anal opening plicate, surrounded by small tubercles; ulnar ridge slightly evident, evidenced by small and white tubercles; relative finger lengths I> III> II �� IV; subarticular tubercles present; finger tips slightly expanded, not bearing discs; fingers free, not fringed or ridged; dorsal surface of thigh and shank tuberculate, tubercles with keratinized tips; hidden parts of thigh and shank smooth; no calcar ornamentation; sole of foot smooth; toe tips slightly expanded, with poorly developed discs; toe relative lengths I Color in life: Ground color of dorsum light brown, with two dark dorsolateral stripes extending from posterior corner of the eyelids to groin, medial dorsum with undefinite darker mottling, canthus rostralis dark brown, flanks barred from the dorsolateral stripes to venter, venter irregularly mottled with brown; dorsal surface of forearms, thighs and shanks barred; anterior and posterior surface of thigh uniformly faintly pigmented with brown; iris bronze. Color in preservative: Similar in life, although slightly faded. Measurements of holotype: SVL 75.2 mm, HL 32.6 mm, HW 36.4 mm, ED 9.9 mm, EN 10.0 mm, TYD 4.5 mm, TYH 5.5 mm, ID 7.9 mm, HAL 19.3 mm, FL 40.0 mm, TL 38.3 mm, FOL 34.9 mm, IMT 3.7 mm. Variation (N= 6): Adult male and females of S. aramunha differ in size (male SVL 40.3���41.1 mm; female SVL 75.2���79.7 mm), HW/SVL ratio (males 0.44���0.46, females 0.48���0.49), development of flared maxilla, and presence of frontoparietal crests. Females are larger, present more developed flared maxilla, and are unique in presenting cranial crests. Males present the skin of dorsum spiculate with tubercles densely and evenly distributed, while in females, the density of tubercles is more variable, usually more concentrated on the posterior half. There is ontogenetic variation in the development of frontoparietal crests and labial flaring in females. In the smaller female (SVL 40.2 mm) of the type series cranial crests are absent; they are already seen in a slightly enlarged specimen (SVL 46 mm) becoming highly conspicuous in large specimens. Sexual dimorphism and ontogenetic variation observed is congruent with those reported for broad-headed eleutherodactylines (Lynch 1975, 1997; Lynch et al. 1994; Savage & Myers 2002). Distribution and ecology: Strabomantis aramunha is presently known only and supposed to be endemic to the ���campos rupestres��� of the neighborhood of Mucug�� municipality, Serra do Sincor��, Espinha��o Range (Fig. 4). The Espinha��o range is well known by the high level of species endemism (e.g. Bokermann 1956, 1964; Cunha 1966; Bokermann & Sazima 1973, 1978; Sazima & Bokermann 1978, 1983; Caramaschi & Sazima 1984, 1985; Vanzolini & Heyer 1988; Pinna 1992; Wege & Long 1995; Stattersfield et al. 1998; Heyer 1999; Lugli & Haddad 2006 a, b; Napoli & Junc�� 2006; Rodrigues et al. 2006). This mountain complex lies over Eastern Brazil at the transitional limits of the Atlantic forest and the Cerrado biomes with elevations from 700���2000 m a.s.l. (Stattersfield et al. 1998). The region is characterized by open and rocky areas dominated by shrubs and grasses referred to as ���campos rupestres��� or ���campos de altitude��� (for detailed characterization of Brazilian ���campos de altitude��� see Safford 1999 a, b). All specimens of S. aramunha were found at night, motionless on exposed rock outcrops in the ���campos rupestres���. This is also the way broad-headed frogs have been frequently found as reported by Lynch (1997). Despite the rain, we did not hear the frogs calling. Nevertheless, the apparent absence of advertisement call is not infrequent among eleutherodactylines (Savage 1987; Savage & Myers 2002). Remarks: This is the first record of an endemic eleutherodactyline from the ���campos rupestres��� of the Espinha��o range. These frogs are generally leaf-litter inhabitants that deposit their eggs in terrestrial humid habitats (Duellman 1978; Duellman & Trueb 1994). Presently the only recorded eleutherodactyline frog from these high elevation areas is the widespread Ischnocnema ramagii (MTR unpublished data). In Brazil, eleutherodactyline frogs generally are associated with forested habitats, and, even those associated to non-forested biomes, like the Brazilian Cerrado, occur inside riparian forests. The single exception is Ischnocnema juipoca, known to occur in open areas (Haddad & Sazima, 1992). In this sense, the discovery of the new species Strabomantis aramunha from ���campos rupestres��� of the Espinha��o range is highly surprising and may represent a relict lineage evolved along with retraction of forests in this mountain complex (Giulietti & Pirani 1988). The discovery of a large broad-headed eleutherodactyline in the high altitude mountains of Espinha��o range in eastern Brazil is enigmatic and reveals how complex is the understanding of the relationships and biogeography of these mountains. In the absence of an exhaustive phylogenetic study including east and southeastern South American Terrarana (Hedges et al. 2008), we can only rely on comparative examples to study the origin of Strabomantis aramunha. Phylogenetic and biogeographical relationships of endemic species from the Espinha��o range are virtually unexplored but some patterns are evident. Some frogs, like Thoropa megatympanum, Bokermannohyla alvarengai, B. nanuzae, B. oxente, B. saxicola, Rupirana cardosoi, Phasmahyla jandaia, Hylodes otavioi, seem to have their closest relatives in species occurring in the eastern Atlantic forest (Bokermann 1956, 1964; Bokermann & Sazima 1973, 1978; Sazima & Bokermann 1983; Caramaschi & Sazima 1984; Heyer 1999; Lugli & Haddad 2006 a, b). This is also the case of some lizards like the genera Placosoma and Enyalius (Cunha 1966; Rodrigues et al. 2006). A very distinctive and possibly much older pattern is shown by species revealing relationships with species present in high elevation areas of northern and/or western South America. For example, species of Microlophus, the closest outgroup known for the lizard genus Eurolophosaurus, are Andean or transandean (Frost et al. 2001). Similarly, among the species of the cisandean Tropidurus of the torquatus group, Tropidurus mucujensis most resemble Tropidurus bogerti, a species from the Ayuantepui in the Tepui region of Venezuela (Rodrigues 1987). Old Atlantic forest relationships are also disclosed by the relictual presence of the Atlantic forest anguid lizard Diploglossus fasciatus in western Amazon Basin (��vila-Pires 1995). Again among lizards, the Rhachisaurinae, an endemic subfamily of gymnophthalmids restricted to these mountains was recently recognized (Pellegrino et al. 2001). Such examples are indicative of how far we are to understand the faunal relationships of this old, Precambrian ridge of eastern South America. The new species is placed in the genus Strabomantis based on morphological characters. However, males of S. aramunha are similar to Haddadus binotatus. This could indicate the generic misplacement of the species. Phylogenetic studies including species from a wide range of Eleutherodactylines (e.g. Craugastor, Haddadus, Holoaden, Ischnocnema, Oreobates, Pristimantis, Strabomantis, etc.) would be needed to asses this and other hypothesis of relationships within this group of frogs., Published as part of Cassimiro, Jos��, Verdade, Vanessa K. & Rodrigues, Miguel T., 2008, A large and enigmatic new eleutherodactyline frog (Anura, Strabomantidae) from Serra do Sincor��, Espinha��o range, Northeastern Brazil, pp. 59-68 in Zootaxa 1761 on pages 60-66, DOI: 10.5281/zenodo.181933, {"references":["Lynch, J. D. & Duellman, W. E. (1997) Frogs of the genus Eleutherodactylus in western Ecuador. Systematics, ecology, and biogeography. Special Publication, Natural History Museum, University of Kansas, 23, 1 - 236.","Hedges, S. B., Duellman, W. E., & Heinicke, M. P. (2008) New World direct-developing frogs (Anura: Terrarana): Molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182.","Lynch, J. D. (1975) A review of the broad-headed Eleutherodactyline frogs of South America (Leptodactylidae). Occasional Papers of the Museum of Natural History, The University of Kansas, 38, 1 - 46.","Frost, D. R. (2007) Amphibian Species of the World: an online reference. Version 5.1 (10 October, 2007). American Museum of Natural History, New York, USA. Available from http: // research. amnh. org / herpetology / amphibia / index. php (accessed 22 January 2008)","Lynch, J. D. (1980) A taxonomic and distributional synopsis of the Amazonian frogs of the genus Eleutherodactylus. American Museum Novitates, 2696, 1 - 24.","Lynch, J. D. & Myers, C. W. (1983) Frogs of the fitzingeri group of Eleutherodactylus in eastern Panama and Chocoan South America (Leptodactylidae). Bulletin of the American Museum of Natural History, 175, 481 - 572.","Lynch, J. D. (1990) A new large species of streamside Eleutherodactylus from western Colombia (Amphibia: Leptodactylidae). Herpetologica, 46, 135 - 142.","Lynch, J. D., Ruiz-Carranza, P. M. & Ardila-Robayo, M. C. (1994) The identities of the Colombian frogs confused with Eleutherodactylus latidiscus (Boulenger) (Amphibia: Anura: Leptodactylidae). Occasional Papers of the Natural History Museum of the University of Kansas, 170, 1 - 42.","Savage, J. M. & Myers, C. W. (2002) Frogs of the Eleutherodactylus biporcatus group (Leptodactylidae) of Central America and Northern South America, including rediscovered, resurrected and new taxa. American Museum Novitates, 3357, 1 - 48.","Bokermann, W. C. A. (1956) Sobre uma nova especie de Hyla do estado de Minas Gerais, Brasil (Amphibia, Salientia, Hylidae). Papeis Avulsos do Departamento de Zoologia, Secretaria da Agricultura, Sao Paulo, 12, 357 - 362.","Bokermann, W. C. A. (1964) Dos nuevas especies de Hyla de Minas Gerais y notas sobre Hyla alvarengai Bok. (Amphibia, Salientia, Hylidae). Neotropica, 10, 67 - 76.","Cunha, O. R. (1966) Sobre uma nova especie de lagarto do estado de Minas Gerais Placosoma cipoense sp. nov. (Lacertilia, Teiidae). Boletim do Museu Paraense Emilio Goeldi, 61, 1 - 9.","Bokermann, W. C. A. & Sazima, I. (1973) Anfibios da Serra do Cipo, Minas Gerais, Brasil. 1. - Especies novas de Hyla (Anura, Hylidae). Revista Brasileira de Biologia, 33, 329 - 336.","Caramaschi, U. & Sazima, I. (1984) Uma nova especie de Thoropa da Serra do Cipo, Minas Gerais, Brasil (Amphibia, Leptodactylidae). Revista Brasileira de Zoologia, 2, 139 - 146.","Caramaschi, U. & Sazima, I. (1985) Uma nova especie de Crossodactylus da Serra do Cipo, Minas Gerais, Brasil (Amphibia, Leptodactylidae). Revista Brasileira de Biologia, 3, 43 - 49.","Vanzolini, P. E. & Heyer, W. R. (1988) Proceedings of a Workshop on Neotropical Distribution Patterns held 12 - 16 January 1987. Academia Brasileira de Ciencias, Rio de Janeiro, 488 pp.","Pinna, M. C. C. (1992) A new subfamily of Trichomycteridae (Teleostei, Sioluriformes), lower loricaroid relationships and a discussion on the impact of additional taxa for phylogenetic analysis. Zoological Journal of the Linnean Society, 106, 175 - 229.","Wege, D. C. & Long. A. C. (1995) Key Areas for Threatened Birds in the Neotropics. BirdLife International, Cambridge.","Stattersfield, J. A., Crosby, M. J., Long, A. J. & Wege, D. C. (1998) Endemic Bird Areas of the World: Priorities for Biodiversity Conservation. The Burlington Press Ltd., Cambridge, U. K.","Heyer, W. R. (1999) A new genus and species of frog from Bahia, Brazil (Amphibia: Anura: Leptodactylidae) with comments on the zoogeography of the Brazilian campos rupestres. Proceedings of the Biological Society of Washington, 112, 19 - 39.","Lugli, L. & Haddad, C. F. B. (2006 a) New species of Bokermannohyla (Anura, Hylidae) from Central Bahia, Brazil. Journal of Herpetology, 40, 7 - 15.","Lugli, L. & Haddad, C. F. B. (2006 b) A new species of the Bokermannohyla pseudopseudis group from Central Bahia, Brazil (Amphibia, Hylidae). Herpetologica, 62, 453 - 465.","Napoli, M. F. & Junca, F. A. (2006) A new species of the Bokermannohyla circumdata group (Amphibia: Anura: Hylidae) from Chapada Diamantina, State of Bahia, Brazil. Zootaxa, 1244, 57 - 68.","Rodrigues, M. T., Freitas, M. A., Silva, T. F. S. & Bertolotto, C. E. V. (2006) A new species of lizard genus Enyalius (Squamata, Leiosauridae) from the highlands of Chapada Diamantina, State of Bahia, Brazil, with a key to species. Phyllomedusa, 5, 11 - 24.","Safford, H. D. (1999 a) Brazilian Paramos I. An introduction to the physical environment and vegetation of the campos the altitude. Journal of Biogeography, 26, 693 - 712.","Safford, H. D. (1999 b) Notas sobre a ecologia e a preservacao de Notochilus coccineus (Scrophulariaceae), especie endemica do macico do Caparao, MG / ES, Brasil. Acta Botanica Brasilica, 13, 175 - 185.","Savage, J. M. (1987) Systematics and distribution of the Mexican and Central American rainfrogs of the Eleutherodactylus gollmeri group (Amphibia: Leptodactylidae). Fieldiana, 33, 1 - 57.","Duellman, W. E. (1978) The biology of an Equatorial herpetofauna in Amazonian Ecuador. Miscellaneous Publications of the University of Kansas Museum of Natural History, 65, 1 - 352.","Duellman, W. E. & Trueb, L. (1994) Biology of Amphibians. The John Hopkins University Press, Maryland, 696 pp.","Haddad, C. F. B. & Sazima, I. (1992) Anfibios anuros da Serra do Japi. In: Morellato, L. P. C. (Eds.), Historia Natural da Serra do Japi: ecologia e preservacao de uma area florestal no Sudeste do Brasil. Editora da Unicamp, Campinas, pp. 188 - 211.","Giulietti, A. M. & Pirani, J. R. (1988) Patterns of geographic distribution of some plant species from the Espinhaco Range, Minas Gerais and Bahia, Brazil. In: Heyer, W. R. & Vanzolini, P. E. (Eds.), Proceedings of a Workshop on Neotropical Distribution Patterns. Academia Brasileira de Ciencias, Rio de Janeiro, pp. 39 - 69.","Sazima, I. & Bokermann, W. C. A. (1983 \" 1982 \") Anfibios da Serra do Cipo, Minas Gerais, Brasil. 5: Hylodes otavioi sp. n. (Anura, Leptodactylidae). Revista Brasileira de Biologia, 42, 767 - 771.","Frost, D. R., Etheridge, R., Janies, D. & Titus, T. A. (2001) Total evidence, sequence alignment, evolution of polychrotid lizards, and a reclassification of the Iguania (Squamata: Iguania). American Museum Novitates, 3343, 1 - 38.","Rodrigues, M. T. (1987) Sistematica, ecologia e zoogeografia dos Tropidurus do grupo torquatus ao sul do rio Amazonas (Sauria, Iguanidae). Arquivos de Zoologia, 31, 105 - 230.","Avila-Pires, T. C. S. (1995) Lizards of Brazilian Amazonia (Reptilia: Squamata). Zoologische Verhandelingen, Leiden, 299, 1 - 706.","Pellegrino, K. C. M., Rodrigues, M. T., Yonenaga-Yassuda, Y. & Sites, J. W. (2001) A molecular perspective on the evolution of microteiid lizards (Squamata, Gymnophthalmidae), and a new classification for the family. Biological Journal of the Linnean Society, 74, 315 - 338."]}

Strabomantis aramunha sp. nov. (Figs. 1–3) Holotype: MZUSP 138693 (field number JC- 1212), an adult female, from Serra do Sincorá, Espinhaço range, 13 ° 04' 07" S and 41 ° 20 ' 09" W, 998 m elevation, municipality of Mucugê, State of Bahia, Brazil, collected by J. Cassimiro and F. S. F. Leite on 11 March 2005. Paratypes (N= 6): MZUSP 138687 (JC- 1198), an adult female, 13 ° 00' 33 " S and 41 ° 22 ' 46 " W, elevation 1191 m, on 8 March 2005; MZUSP 138688 (JC- 1206), an adult male, 13 ° 00' 32 " S and 41 ° 22 ' 48 " W, elevation 1207 m, same date; MZUSP 138690 (JC- 1214), an adult male, 13 ° 03' 43 " S and 41 ° 20 ' 26 " W, elevation 947 m, on 11 March 2005; MZUSP 138689 (JC- 1213), juvenile female, 13 ° 04' 07" S and 41 ° 20 ' 09" W, elevation 998 m, same date; MZUSP 138691 (JC- 1251), juvenile male, on 20 March 2005; MZUSP 138692 (JC- 1277), juvenile female, 13 ° 01' 20 " S and 41 ° 21 ' 24 " W, elevation 1068 m, on 26 March 2005. All collected by J. Cassimiro and F. S. F. Leite at the surroundings of the type locality. Variable MZUSP MZUSP MZUSP MZUSP MZUSP MZUSP MZUSP 138693 138687 138688 138690 138691 138689 138692 Etymology: The specific name is from the Tupi language aramunha, meaning giant, in allusion to the large size of the species, and is used as noun in apposition. Diagnosis: A large broad-headed species (adult male 40.3–41.1 mm SVL, adult female 75.2–79.7 mm; HW respectively 44–46 % and 48–49 % of SVL), adult females bearing well-developed frontoparietal crests, posterior part of pars fascialis of maxilla deepened, mandibular ramus of the trigeminal nerve passing lateral to the m. levator mandibulae posterior subexternus, eyelid tubercles present posteriorly, not elongated, skin of venter coarsely areolate, leg relatively short (FL 51–53 % of SVL) finger discs absent, first finger longer than second, poorly developed discs on toes, first and second toes ridged, fifth toe smaller than third, tarsal fold absent, testes white. Comparison to other species: The adult Strabomantis aramunha is promptly diagnosed from all other eleutherodactylines by the combination of large size, broad-head, flared maxilla, frontoparietal bearing welldeveloped crests in females, mandibular ramus of the trigeminal nerve passing lateral to the m. levator mandibulae posterior subexternus, posterior eyelid tubercles present and not elongated, skin of venter coarsely areolate, absence of finger discs, first finger longer than second, poorly developed discs on toes, first and second toes ridged, fifth toe smaller than third, and, tarsal fold absent. These characters place squarely the new species in the Eleutherodactylus sulcatus group (Lynch 1997; Lynch & Duellman 1997), currently in the genus Strabomantis (Hedges et al. 2008). The only other species of Strabomantis occurring in Brazil is S. sulcatus (Lynch 1975, 1997; Frost 2007; Hedges et al. 2008). Strabomantis sulcatus differ from S. aramunha by the smaller size (SVL 28–42 mm; Lynch 1975, 1980), concealed surfaces of thigh and shank presenting light blotches, and by the presence of tarsal fold. From the other species of Strabomantis, S. aramunha differ by the larger size (adult female S. cadenai, SVL 40.6 mm; Lynch 1997), skin of venter areolate (smooth in S. cadenai, S. laticorpus; Lynch 1997), by not presenting elongate conical tubercles on eyelid (present in S. cadenai, S. cerastes, S. ingeri, S. laticorpus, S. necopinus, S. ruizi; Lynch 1997), large flattened warts absent (present in S. helonotus; Lynch 1997), tarsal fold absent (present in S. anomalus, S. cornutus, S. ingeri, S. necopinus, and S. ruizi; Lynch 1997), cranial crests present (absent in S. anomalus, and S. cheiroplethus; Lynch & Myers 1983; Lynch 1990), reduced basal web only between first and third toes (extensive toe webbing in S. anomalus, S. anatipes and S. zygodactylus; Lynch & Myers 1983). Males and the young specimens, by being smaller and presenting narrower heads most resemble the adult Haddadus binotatus (formerly “ Eleutherodactylus ” binotatus, Hedges et al. 2008). However, under close examination H. binotatus is promptly differentiated from S. aramunha by the smoother dorsal skin and the presence of tubercles not bearing keratinized tips (present in S. aramunha). Additionally, the head is narrower and longer than wide (wider than long in S. aramunha), the body is slender, the legs are longer, and the supernumerary tubercles are more numerous in H. binotatus than in S. aramunha. Description of holotype: Head wider than long (HW = 36.4 mm; HL = 32.6 mm); a pair of frontoparietal crests externally visible; eyes dorsolateral, median sized (ED = 9.9 mm), diameter similar to the eye-nostril distance (EN = 10.0 mm); snout sub-elliptical in dorsal view, rounded in lateral view, overhanging lower jaw; nostrils small, slightly protuberant, directed laterally and positioned distally at the canthus rostralis; canthus rostralis sharp, accentuated by bony keel; loreal region deeply concave, tuberculate; flared maxilla, with posterior pars fascialis deepened; upper eyelid bearing small tubercles posteriorly, not elongated; interorbital space highly depressed, bearing small tubercles; tympanic membrane smooth; tympanic annulus prominent, higher than long (TYD = 4.5 mm; TYH = 5.5 mm), oblique; tympanum-eye distance less than tympanum diameter (TED = 3.6 mm); supratympanic fold evident, extending from posterior corner of eye to axils. Skin of dorsum spiculate under magnification, with tubercles more concentrated on posterior half; dorsolateral ridges present; skin of venter coarsely areolate; anal opening plicate, surrounded by small tubercles; ulnar ridge slightly evident, evidenced by small and white tubercles; relative finger lengths I> III> II ˜ IV; subarticular tubercles present; finger tips slightly expanded, not bearing discs; fingers free, not fringed or ridged; dorsal surface of thigh and shank tuberculate, tubercles with keratinized tips; hidden parts of thigh and shank smooth; no calcar ornamentation; sole of foot smooth; toe tips slightly expanded, with poorly developed discs; toe relative lengths I Color in life: Ground color of dorsum light brown, with two dark dorsolateral stripes extending from posterior corner of the eyelids to groin, medial dorsum with undefinite darker mottling, canthus rostralis dark brown, flanks barred from the dorsolateral stripes to venter, venter irregularly mottled with brown; dorsal surface of forearms, thighs and shanks barred; anterior and posterior surface of thigh uniformly faintly pigmented with brown; iris bronze. Color in preservative: Similar in life, although slightly faded. Measurements of holotype: SVL 75.2 mm, HL 32.6 mm, HW 36.4 mm, ED 9.9 mm, EN 10.0 mm, TYD 4.5 mm, TYH 5.5 mm, ID 7.9 mm, HAL 19.3 mm, FL 40.0 mm, TL 38.3 mm, FOL 34.9 mm, IMT 3.7 mm. Variation (N= 6): Adult male and females of S. aramunha differ in size (male SVL 40.3–41.1 mm; female SVL 75.2–79.7 mm), HW/SVL ratio (males 0.44–0.46, females 0.48–0.49), development of flared maxilla, and presence of frontoparietal crests. Females are larger, present more developed flared maxilla, and are unique in presenting cranial crests. Males present the skin of dorsum spiculate with tubercles densely and evenly distributed, while in females, the density of tubercles is more variable, usually more concentrated on the posterior half. There is ontogenetic variation in the development of frontoparietal crests and labial flaring in females. In the smaller female (SVL 40.2 mm) of the type series cranial crests are absent; they are already seen in a slightly enlarged specimen (SVL 46 mm) becoming highly conspicuous in large specimens. Sexual dimorphism and ontogenetic variation observed is congruent with those reported for broad-headed eleutherodactylines (Lynch 1975, 1997; Lynch et al. 1994; Savage & Myers 2002). Distribution and ecology: Strabomantis aramunha is presently known only and supposed to be endemic to the “campos rupestres” of the neighborhood of Mucugê municipality, Serra do Sincorá, Espinhaço Range (Fig. 4). The Espinhaço range is well known by the high level of species endemism (e.g. Bokermann 1956, 1964; Cunha 1966; Bokermann & Sazima 1973, 1978; Sazima & Bokermann 1978, 1983; Caramaschi & Sazima 1984, 1985; Vanzolini & Heyer 1988; Pinna 1992; Wege & Long 1995; Stattersfield et al. 1998; Heyer 1999; Lugli & Haddad 2006 a, b; Napoli & Juncá 2006; Rodrigues et al. 2006). This mountain complex lies over Eastern Brazil at the transitional limits of the Atlantic forest and the Cerrado biomes with elevations from 700–2000 m a.s.l. (Stattersfield et al. 1998). The region is characterized by open and rocky areas dominated by shrubs and grasses referred to as “campos rupestres” or “campos de altitude” (for detailed characterization of Brazilian “campos de altitude” see Safford 1999 a, b). All specimens of S. aramunha were found at night, motionless on exposed rock outcrops in the “campos rupestres”. This is also the way broad-headed frogs have been frequently found as reported by Lynch (1997). Despite the rain, we did not hear the frogs calling. Nevertheless, the apparent absence of advertisement call is not infrequent among eleutherodactylines (Savage 1987; Savage & Myers 2002). Remarks: This is the first record of an endemic eleutherodactyline from the “campos rupestres” of the Espinhaço range. These frogs are generally leaf-litter inhabitants that deposit their eggs in terrestrial humid habitats (Duellman 1978; Duellman & Trueb 1994). Presently the only recorded eleutherodactyline frog from these high elevation areas is the widespread Ischnocnema ramagii (MTR unpublished data). In Brazil, eleutherodactyline frogs generally are associated with forested habitats, and, even those associated to non-forested biomes, like the Brazilian Cerrado, occur inside riparian forests. The single exception is Ischnocnema juipoca, known to occur in open areas (Haddad & Sazima, 1992). In this sense, the discovery of the new species Strabomantis aramunha from “campos rupestres” of the Espinhaço range is highly surprising and may represent a relict lineage evolved along with retraction of forests in this mountain complex (Giulietti & Pirani 1988). The discovery of a large broad-headed eleutherodactyline in the high altitude mountains of Espinhaço range in eastern Brazil is enigmatic and reveals how complex is the understanding of the relationships and biogeography of these mountains. In the absence of an exhaustive phylogenetic study including east and southeastern South American Terrarana (Hedges et al. 2008), we can only rely on comparative examples to study the origin of Strabomantis aramunha. Phylogenetic and biogeographical relationships of endemic species from the Espinhaço range are virtually unexplored but some patterns are evident. Some frogs, like Thoropa megatympanum, Bokermannohyla alvarengai, B. nanuzae, B. oxente, B. saxicola, Rupirana cardosoi, Phasmahyla jandaia, Hylodes otavioi, seem to have their closest relatives in species occurring in the eastern Atlantic forest (Bokermann 1956, 1964; Bokermann & Sazima 1973, 1978; Sazima & Bokermann 1983; Caramaschi & Sazima 1984; Heyer 1999; Lugli & Haddad 2006 a, b). This is also the case of some lizards like the genera Placosoma and Enyalius (Cunha 1966; Rodrigues et al. 2006). A very distinctive and possibly much older pattern is shown by species revealing relationships with species present in high elevation areas of northern and/or western South America. For example, species of Microlophus, the closest outgroup known for the lizard genus Eurolophosaurus, are Andean or transandean (Frost et al. 2001). Similarly, among the species of the cisandean Tropidurus of the torquatus group, Tropidurus mucujensis most resemble Tropidurus bogerti, a species from the Ayuantepui in the Tepui region of Venezuela (Rodrigues 1987). Old Atlantic forest relationships are also disclosed by the relictual presence of the Atlantic forest anguid lizard Diploglossus fasciatus in western Amazon Basin (Ávila-Pires 1995). Again among lizards, the Rhachisaurinae, an endemic subfamily of gymnophthalmids restricted to these mountains was recently recognized (Pellegrino et al. 2001). Such examples are indicative of how far we are to understand the faunal relationships of this old, Precambrian ridge of eastern South America. The new species is placed in the genus Strabomantis based on morphological characters. However, males of S. aramunha are similar to Haddadus binotatus. This could indicate the generic misplacement of the species. Phylogenetic studies including species from a wide range of Eleutherodactylines (e.g. Craugastor, Haddadus, Holoaden, Ischnocnema, Oreobates, Pristimantis, Strabomantis, etc.) would be needed to asses this and other hypothesis of relationships within this group of frogs.

Made available in DSpace on 2021-06-25T12:17:32Z (GMT). No. of bitstreams: 0 Previous issue date: 2016-11-23 Villanova Univ, Dept Biol, Villanova, PA 19085 USA Museu Nacl Hist Nat & Ciencia, Lisbon, Portugal Univ Brasilia, Dept Zool, Inst Ciencias Biol, BR-70910900 Brasilia, DF, Brazil Smithsonian Inst, Natl Museum Nat Hist, Washington, DC 20560 USA Sorbonne Univ, Museum Natl Hist Nat, ISYEB, Paris, France Univ Nacl Tucuman, San Miguel De Tucuman, Argentina King Saud Univ, Riyadh, Saudi Arabia Cornell Univ, Ithaca, NY USA Univ Fed Sao Paulo, Diadema, Brazil Upper Austrian Museum, Ctr Biol, Linz, Austria Villanova Univ, Villanova, PA 19085 USA Univ Salzburg, Salzburg, Austria Plazi, Bern, Switzerland Univ Brasilia, Brasilia, DF, Brazil Univ Sao Paulo, Ribeirao Preto, Brazil Zool Forsch Museum A Koenig, Bonn, Germany Museu Paraense Emilio Goeldi, Belem, Para, Brazil Russian Acad Sci, St Petersburg, Russia Pontificia Univ Catolica Rio Grande do Sul, Porto Alegre, RS, Brazil Museo Reg Sci Nat, Turin, Italy Univ Fed Rio Grande do Sul, Porto Alegre, RS, Brazil Royal Museum Cent Africa, Tervuren, Belgium Univ Nacl Autonoma Mexico, Mexico City, DF, Mexico Univ Innsbruck, Inst Ecol, Innsbruck, Austria Hasselt Univ, Hasselt, Belgium Univ Fed Pernambuco, Recife, PE, Brazil Univ Fed Espirito Santo, Vitoria, ES, Brazil Univ Nacl Misiones, Felix De Azara, Argentina Albany Museum, Grahamstown, South Africa Natl Museum, Bloemfontein, South Africa Senckenberg Nat Hist Sammlungen, Dresden, Germany Univ Estadual Paulista, Botucatu, SP, Brazil Stephen F Austin State Univ, Nacogdoches, TX 75962 USA Landesmuseum, Leonding, Austria Univ Camerino, Camerino, Italy Univ Fed ABC, Sao Bernardo, Brazil Univ Estadual Londrina, Museu Zool, Londrina, Parana, Brazil Senckenberg Res Inst, Frankfurt, Germany Museo Civ Storia Nat, Carmagnola, Italy Polskiej Akad Nauk, Muzeum & Inst Zool, Warsaw, Poland Museo Civ Storia Nat, Genoa, Italy Russian Acad Sci, Inst Zool, St Petersburg, Russia Univ Estadual Paulista, Sao Vicente, Brazil Sorbonne Univ, ISYEB, Museum Natl Hist Nat, Paris, France Port Elizabeth Museum, Port Elizabeth, South Africa Sam Noble Museum, Norman, OK USA Harvard Univ, Museum Comparat Zool, Cambridge, MA 02138 USA North West Univ, Potchefstroom, South Africa Museu Nacl, Rio De Janeiro, Brazil Nat Hist Museum, Basel, Switzerland Senckenberg Nat Kundemuseum, Gorlitz, Germany North Carolina Museum Nat Sci, Raleigh, NC USA Univ Rennes 1, Rennes, France Univ Sao Paulo, Museu Zool, Sao Paulo, SP, Brazil Amer Museum Nat Hist, New York, NY 10024 USA Univ Huddersfield, Huddersfield, W Yorkshire, England North Dakota State Univ, Fargo, ND USA Univ Sao Paulo, Sao Paulo, SP, Brazil Fac Sci Sfax, Sfax, Tunisia DPTC PC, Manaus, Amazonas, Brazil Louisiana State Univ, Museum Nat Sci, Baton Rouge, LA 70803 USA Zool Survey India, Kolkata, India Univ Tennessee, Chattanooga, TN USA Ohio State Univ, Columbus, OH 43210 USA Univ Luterana Brasil, Museu Ciencias Nat, Canoas, Brazil Univ Firenze, Florence, Italy Univ Ovidius Constanta, Constanta, Romania Univ Mississippi, Oxford, MS USA Royal Belgian Inst Nat Sci, Brussels, Belgium Univ New Mexico, Albuquerque, NM 87131 USA Inst Nacl de Pesquisas da Amazonia, Manaus, Amazonas, Brazil Univ Stellenbosch, Matieland, South Africa Univ Fed Minas Gerais, Belo Horizonte, MG, Brazil CIBIO Ctr Invest Biodiversidade & Recursos Genet, Vairao, Portugal Museum Hist Nat, Geneva, Switzerland Univ Fed Mato Grosso, Cuiaba, Brazil Univ Austral Chile, Valdivia, Chile Univ Malaysia, Sarawak, Malaysia Indian Inst Sci, Bangalore, Karnataka, India Scottish Assoc Marine Sci, Oban, Argyll, Scotland Sam Houston State Univ, Huntsville, TX 77340 USA Museo Nacl Ciencias Nat, Madrid, Spain Sorbonne Univ, CR2P, Museum Natl Hist Nat, Paris, France Drexel Univ, Acad Nat Sci, Philadelphia, PA 19104 USA Univ Richmond, Richmond, VA 23173 USA Ctr Pesquisas Cacau, Itabuna, Brazil Univ Torino, Turin, Italy Soc Hist Nat Alcide dOrbigny, Aubiere, France Wolfden Sci Consulting, Murcia, Spain Univ Fed Rio de Janeiro, Macae, Brazil Univ Tennessee, Knoxville, TN USA Inst Humboldt, Villa De Leyva, Colombia Escuela Politec Nacl, Quito, Ecuador Calif Acad Sci, San Francisco, CA 94118 USA Museum Dept Hist Nat Var, Toulon, France Nat Environm & Wildlife Soc, Angul, India Univ Kansas, Lawrence, KS 66045 USA Kyoto Univ, Kyoto, Japan Consejo Nacl Invest Cient & Tecn, Museo Argentino Ciencias Nat Bernardino Rivadavia, Buenos Aires, DF, Argentina Univ Fed Paraiba, Joao Pessoa, Paraiba, Brazil Univ Fed Parana, Curitiba, Parana, Brazil Univ Fed Rio de Janeiro, Rio De Janeiro, Brazil Nat Hist Museum Narodini Museum, Prague, Czech Republic Nat Hist Museum Zimbabwe, Bulawayo, Zimbabwe Ateneo Manila Univ, Quezon City, Philippines Pontificia Univ Javeriana, Bogota, Colombia RACINE, St Maugan, France Univ Paris 06, Sorbonne Univ Paris, ISYEB, Paris, France Ocean Univ China, Qingdao, Peoples R China Museo Nacl Hist Nat Paraguay, San Lorenzo, Paraguay Univ Carabobo, Valencia, Venezuela Natl Ctr Biol Sci, Bengaluru, India Univ Nebraska, Lincoln, NE USA CENAK Ctr Nat Kunde, Hamburg, Germany Ohio Hist Connect, Columbus, OH USA Univ Fed Amazonas, Manaus, Amazonas, Brazil Univ Malaga, Malaga, Spain Chinese Acad Sci, Qingdao, Shandong, Peoples R China Benemerita Univ Autonoma Puebla, Puebla, Mexico Natl Polytech Inst, Ctr Res & Adv Studies, Irapuato, Mexico Field Museum Nat Hist, Chicago, IL 60605 USA Univ Texas El Paso, El Paso, TX 79968 USA Univ Fed Lavras, Lavras, Brazil Nat Hist Museum, Vienna, Austria La Sierra Univ, Riverside, CA USA Univ San Francisco Quito, Quito, Ecuador Univ Hong Kong, Hong Kong, Peoples R China CUNY, New York, NY 10021 USA CAS, Inst Vertebrate Biol, Brno, Czech Republic Sorbonne Univ, MECADEV, Museum Natl Hist Nat, Paris, France Univ Estadual Paulista, Rio Claro, Brazil Zool Staatssammlung Munchen, Munich, Germany Univ Michigan, Dearborn, MI 48128 USA Smithsonian Inst, Natl Museum Nat Hist, Washington, DC USA Helmholtz Ctr Environm Res, Leipzig, Germany Santa Rosa Jr Coll, Santa Rosa, CA USA George Washington Univ, Washington, DC 20037 USA Inst Teknol Bandung, Bandung, Indonesia Earlham Coll, Richmond, IN USA Senckenberg Forschungsinst & Nat Museum, Frankfurt, Germany Univ Estadual Londrina, Londrina, Parana, Brazil Univ Koblenz Landau, Koblenz, Germany Nat Hist Museum, London, England Ditsong Natl Museum Nat Hist, Pretoria, South Africa Bombay Nat Hist Soc, Bombay, Maharashtra, India Philipps Univ Marburg, Marburg, Germany Hess Landesmuseum, Darmstadt, Germany Smithsonian Conservat Biol Inst, Washington, DC USA Univ Pannonia, Keszthely, Hungary Staatliches Museum Nat Kunde, Stuttgart, Germany UFT, Tocantins, Portugal Museum Nat Kunde, Berlin, Germany Nat Hist Riksmuseet, Stockholm, Sweden Hikarigaoka, Munakata, Japan Inst Agr & Environm Sci, Tartu, Estonia Univ Los Andes, Merida, Venezuela Bat Jungle, Monteverde, Costa Rica Texas A&M Univ, College Stn, TX USA Yale Univ, New Haven, CT USA Univ Estadual Paulista, Sao Jose Do Rio Preto, Brazil Museo Nacl Hist Nat, Montevideo, Uruguay Consejo Nacl Invest Cient & Tecn, Fdn Miguel Lillo, San Miguel De Tucuman, Argentina Roger Tory Peterson Inst Nat Hist, Jamestown, VA USA Univ Fed Reconcavo Bahia, Cruz Das Almas, Brazil South China Normal Univ, Guangzhou 510631, Guangdong, Peoples R China Museu Ciencias Tecnol PUCRS, Porto Alegre, RS, Brazil Univ Fed Tocantins, Porto Nacl, Brazil Univ Toronto, Scarborough, ON, Canada Univ Nacl Colombia, Inst Ciencias Nat, Bogota, Colombia Natl Museum Philippines, Manila, Philippines NOVA Univ Lisbon, Caparica, Portugal Royal Saskatchewan Museum, Regina, SK, Canada Deutsch Zentrum Marine Biodiversitatsforsch, Hamburg, Germany Ctr Invest Biol Noroeste, La Paz, Mexico Naturalis Biodivers Ctr, Leiden, Netherlands Univ Douala, Douala, Cameroon Vet & Farmaceut Univ Brno, Brno, Czech Republic Univ Swaziland, Kwaluseni, Eswatini Fundacao Oswaldo Cruz, Rio De Janeiro, Brazil Univ Claude Bernard, Lyon, France Museum Vertebrate Zool, Berkeley, CA USA Smithsonian Inst, Washington, DC 20560 USA Nat Hist Museum & Inst, Chiba, Japan Senckenberg Biodiversitat & Klima Forschunsgzentr, Frankfurt, Germany Marquette Univ, Milwaukee, WI 53233 USA Univ Firenze, Museo Storia Nat, Florence, Italy Univ Fed Bahia, Salvador, BA, Brazil Fed Univ ABC, Sao Bernardo, Brazil Univ Fed Goias, Goiania, Go, Brazil Univ Alaska Museum, Fairbanks, AK USA Egerton Univ, Egerton, Kenya Museo Hist Nat, Lima, Peru IKIAM Univ Reg Amazon, Tena, Ecuador Univ Hyogo, Sanda, Japan Inst Rech Dev, Paris, France Univ Sao Paulo, Piracicaba, Brazil Niedersachs Landesbetrieb Wasserwirtschaft Kusten, Hannover, Germany Univ Complutense Madrid, Madrid, Spain Consejo Nacl Invest Cient & Tecn, Ctr Nacl Patagon, Puerto Madryn, Argentina Roosevelt Univ, Coll Profess Studies, Chicago, IL 60605 USA Western Kentucky Univ, Bowling Green, KY 42101 USA Univ Naples Federico II, Naples, Italy Univ Fed Acre, Rio Branco, Acre, Brazil Univ Fed Santa Catarina, Florianopolis, SC, Brazil Inst Bio & Geociencias Noroeste Argentino, Salta, Argentina Inst Venezolano Invest Cient, Caracas, Venezuela Umweltbundesamt, Vienna, Austria Penclen, Plumelec, France CNRS MNHN UPMC, Ctr Rech Paleobiodivers & Paleoenvironm, Paris, France Staatl Museum Nat Kunde, Stuttgart, Germany Consejo Nacl Invest Cient & Tecn, Museo Argentino Ciencias Nat Bernardino Rivada, Buenos Aires, DF, Argentina Ctr Biol Gest Populat INRA, Montferrier Sur Lez, France Loyola Univ Chicago, Chicago, IL USA Pontificia Univ Catolica Rio do Sul, Porto Alegre, RS, Brazil Durban Museum Nat Sci, Durban, South Africa Univ Estado Amazonas, Manaus, Amazonas, Brazil Brigham Young Univ, Provo, UT 84602 USA Museo Hist Nat La Salle, Caracas, Venezuela Univ Vet Med Hannover, Hannover, Germany Australian Museum, Sydney, NSW, Australia Texas Tech Univ, Lubbock, TX 79409 USA Francis Marion Univ, Florence, SC USA Univ Washington, Seattle, WA 98195 USA Univ Connecticut, Groton, CT USA Fed Univ ABC, Santo Andre, SP, Brazil Univ Fed Uberlandia, Uberlandia, MG, Brazil Fed Univ Para, Belem, Para, Brazil Yale Peabody Museum, New Haven, CT USA Univ Texas Arlington, Arlington, TX 76019 USA Senckenberg Deutsch Entomol Inst, Muncheberg, Germany CSIRO, Natl Res Collect, Canberra, ACT, Australia Univ Venda, Thohoyandou, South Africa Univ Ulsan, Ulsan, South Korea Univ Estadual Sao Paulo, Botucatu, SP, Brazil Senckenberg Forsch Inst & Nat Museum, Frankfurt, Germany Ocean Univ China, Inst Marine Biodivers & Evolut, Qingdao, Shandong, Peoples R China Univ Fed Para, Altamira, Brazil Univ Estadual Paulista, Assis, Brazil UPMC, Ctr Rech Paleobiodiversite & Paleoenvironm, CNRS, MNHN, Paris, France Tech Univ Kaiserslautern, Kaiserslautern, Germany Charles Univ Prague, Dept Zool, Prague, Czech Republic Univ Nacl Expt los Llanos Occident Ezequiel Zamor, Guanare, Venezuela Long Isl Univ, Brooklyn, NY USA Univ Ryukyus, Okinawa, Japan Pontificia Univ Catolica Ecuador, Museo Zool, Escuela Ciencias Biol, Quito, Ecuador Indiana Univ Penn, Indiana, PA USA IRD, Dakar, Senegal State Museum Nat Hist Karlsruhe, Karlsruhe, Germany Univ Toulouse III Paul Sabatier, Toulouse, France Univ Ghent, Ghent, Belgium Univ Antwerp, Antwerp, Belgium Karlsruhe Inst Technol, Karlsruhe, Germany Ctr Cellular & Mol Biol, Hyderabad, India Univ Fed ABC, Santo Andre, Brazil Yale Peabody Museum Nat Hist, New Haven, CT USA Univ Innsbruck, Mondsee, Austria Bethune Cookman Univ, Daytona Beach, FL USA Natl Museum Nat & Sci, Tokyo, Japan Zool Garten Koln, Cologne, Germany Univ Estadual Paulista, Botucatu, SP, Brazil Univ Estadual Paulista, Sao Vicente, Brazil Univ Estadual Paulista, Rio Claro, Brazil Univ Estadual Paulista, Sao Jose Do Rio Preto, Brazil Univ Estadual Paulista, Assis, Brazil