Rivulus amanapira Costa, Material examined

Transcription

1 Material examined Brazil: Estado do Amazonas: UFRJ 5925, holotype; UFRJ 5926, 36 paratypes; UFRJ 5927, 4 paratypes (c&s); MCP 34858, 2 paratypes; São Gabriel da Cachoeira, stream near igarapé Iá, km 9.4 of the road São Gabriel da Cachoeira to Cucuí, upper rio Negro drainage, rio Amazonas basin; W. J. E. M. Costa, S. Lima and L. Silva, 30 Aug UFRJ 5928, 9; São Gabriel da Cachoeira, upper rio Negro drainage, rio Amazonas basin, stream tributary to igarapé Miuá, km 13.6 of the road São Gabriel da Cachoeira to Cucuí, S W; W. J. E. M. Costa, S. Lima and L. Silva, 30 Aug Wilson J. E. M. Costa Diagnosis Distinguished from all other congeners by the combination of the following features: anterior portion of trunk slightly deeper than wide, jaws short, snout blunt, tip of anal fin slightly pointed in male, caudal fin rounded in male, pelvic-fin tip reaching middle of anal-fin base in male, dorsal-fin origin on vertical between base of 8th and 9th anal-fin rays, dorsal-fin rays 7-8, anal-fin rays 11-12, frontal squamation S-patterned, frontal scales arranged transversely, canal preopercular absent, contact organs on flank scales in male, longitudinal series of scales 32-33, gill rakers of first branchial arch , oblique rows of red dots on flank in male, transverse dark brown bar through the chin, and dark grey to black spot on dorsal portion of caudal-fin in both sexes. Fig. 37. Rivulus uakti, UFRJ 5925, male, holotype, 24.0 mm SL (two days after collection); Brazil: Amazonas: São Gabriel da Cachoeira. Photo by W. J. E. M. Costa. Fig. 38. Rivulus uakti, UFRJ 5926, female, paratype, 27.9 mm SL (two days after collection); Brazil: Amazonas: São Gabriel da Cachoeira. Photo by W. J. E. M. Costa. Fig. 39. Brazil: Amazonas: São Gabriel da Cachoeira; creek at the forest border, the habitat of Rivulus uakti. Photo by W. J. E. M. Costa. Also differs from all other congeners of the Owiyeye in possessing two unique colour patterns: dorsal fin with bright blue distal zone in male, and caudal spot forming a black bar posteriorly bordered by pale yellow bar in female (Figs 37-38). Distribution Upper rio Negro basin (Fig. 22). Habitat Moderately sunny places of shallow streams within the forest, sandy bottom and reddish hyaline water (Fig. 39). Rivulus amanapira Costa, 2004 Rivulus amanapira Costa, 2004f: 7 (type locality: São Gabriel da Cachoeira, pools near igarapé Palestina, Airport road, upper rio Negro drainage, rio Amazonas basin, S W, altitude 110 m, Estado do Amazonas, Brazil; holotype: UFRJ 5929). Material examined Brazil: Estado do Amazonas: UFRJ 5929, holotype; UFRJ 5930, 11 paratypes; UFRJ 5931, 3 paratypes (c&s); MCP 34859, 2 paratypes; São Gabriel da Cachoeira, pools near igarapé Palestina, Airport road, upper rio Negro drainage; W. J. E. M. Costa, S. Lima and L. Silva, 30 Aug Diagnosis Distinguished from all other congeners by the 159 aqua vol. 11 no

2 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin combination of the following features: anterior portion of trunk slightly deeper than wide, jaws short, snout blunt, tip of anal fin slightly pointed in male, caudal fin truncate in male, pelvic-fin tip reaching anterior portion of anal-fin base in male, dorsal-fin origin on vertical between base of 11 th and 13 th analfin rays, dorsal-fin rays 8, anal-fin rays 14-15, frontal squamation S-patterned, frontal scales arranged transversely, canal preopercular absent, contact organs on flank scales in male, longitudinal series of scales 48-49, gill rakers of first branchial arch 1 + 8, longitudinal rows of red dots on flank in male, transverse dark brown bar through the chin, and ovoid dark grey to black spot on dorsal portion of caudal fin in both sexes. Additionally distinguished from all other members of Owiyeye in having entire marginal region of caudal fin bright yellow in male (Fig. 40). Distribution Upper rio Negro drainage (Fig. 23). Fig. 40. Rivulus amanapira, UFRJ 5929, male, holotype, 38.6 mm SL (one day after collection); Brazil: Amazonas: São Gabriel da Cachoeira. Photo by W. J. E. M. Costa. Habitat notes Small shallow temporary pools in the forest, about m in diameter and about 15 cm deep (Fig. 41). Rivulus tecminae Thomerson, Nico & Taphorn, 1992 Rivulus tecminae Thomerson, Nico & Taphorn, 1992: 290 (type locality: open savanna pools about 500 m from left bank, río Guayapo, about 83 km above confluence with río Sipapo, Orinoco basin, Amazonas Federal Territory, Venezuela, 4 16 N W; holotype: MCNG 23886). Material examined Venezuela: Território Federal del Amazonas: MCNG 23886, holotype; MCNG 21369, 33 paratypes; open savanna pools about 500 m from left bank, río Guayapo, about 83 km above confluence with río Sipapo, Orinoco basin; L. Nico & E. Guayamare, 28 May Brazil: Estado do Amazonas: UFRJ 2085, 2; UFRJ 2095, 1 (c&s); Inambu, right bank of upper rio Negro; D. Teixeira, 26 Aug Fig. 41. Brazil: Amazonas: São Gabriel da Cachoeira; shallow temporary pool within the forest, habitat of Rivulus amanapira. Photo by W. J. E. M. Costa. Diagnosis Distinguished from all other congeners by the combination of the following features: anterior portion of trunk slightly deeper than wide, jaws short, snout blunt, tip of anal fin rounded in male, caudal fin truncate in male, pelvic-fin tip reaching middle of anal-fin base in male, dorsal-fin origin on vertical between base of 9 th and 10 th anal-fin rays, dorsalfin rays 8-10, anal-fin rays 12-15, frontal squamation S-patterned, frontal scales arranged transversely, canal preopercular absent, contact organs on flank scales in male, longitudinal series of scales 37-41, gill rakers of first branchial arch 1 + 7, red stripes on aqua vol. 11 no

3 Wilson J. E. M. Costa Fig. 42. Jaw suspensorium and opercular apparatus of Rivulus pictus, UFRJ 5959, female, 27.7 mm SL. AA = angulo-articular; DE = dentary; HY = hyomandibula; IO = interopercle; MS = mesopterygoid; MT = metapterygoid; MX = maxilla; OP = opercle; PL = autopalatine; PM = premaxilla; PO = preopercle; QU = quadrate; RA = retro-articular; RC = rostral cartilage; SO = subopercle; SY = sympletic. Arrows indicate: 1 = short and pointed dorsal portion of preopercle (synapomorphy 25.1), diagnostic for Melanorivulus; 2 = vestigial ventral process of the angulo-articular (synapomorphy 18.3), diagnostic for a clade including R. pictus and R. apiamici. Larger stippling indicates cartilage. Scale bar 1 mm. flank, transverse black bar through the chin, and vertically elongated dark grey to black spot on dorsal portion of caudal-fin in both sexes. Distribution Upper Orinoco river basin, Venezuela, and upper Negro river, Brazil (Fig. 19). Habitat Temporary pools in savannas (D. Teixeira, pers. comm.). Melanorivulus, new subgenus Type species: Rivulus punctatus Boulenger, Diagnosis Distinguished from the remaining subgenera of Rivulus in having the following apomorphic features: dorsal portion of preopercle short and pointed (25.1) (Fig. 42), dorsal and anal fins slightly pointed in male (72.1, 73.1; also occurring in Laimosemion) (Fig. 43), two oblique bars on post-orbital region (97.1) (Fig. 43), melanophores concentrated on margins of unpaired and pelvic fins in female (111.1) (Fig. 44), and female with black spot on upper portion of caudal fin not close to fin margin, overlapping caudal fin bars (113.2) (Fig. 44). All included species except R. modestus have oblique chevron-like rows of red dots or bars on flank in males (90.2) (Fig. 43). Etymology From the Latin, melania (black pigmentation on the skin) and rivulus (stream), referring to the black margins of unpaired and pelvic fins, a condition unique among congeners. Gender masculine. Included species Rivulus apiamici Costa, 1989, R. cyanopterus Costa, 2005, R. dapazi Costa, 2005, R. decoratus Costa, 1989, R. egens Costa, 2005, R. litteratus Costa, 2005, R. modestus Costa, 1991, R. paracatuensis Costa, 2003, R. parnaibensis Costa, 2003, R. pictus Costa, 1989, R. pinima Costa, 1989, R. punctatus Boulenger, 1895, R. rossoi Costa, 2005, R. rutilicaudus Costa, 2005, R. scalaris Costa, 2005, R. violaceus Costa, 1991, R. vittatus Costa, 1989, and R. zygonectes Myers, Distribution Paraná-Paraguay-Uruguay, upper Tapajós, upper and middle Xingu, upper and middle Tocantins, 161 aqua vol. 11 no

4 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin São Francisco and Parnaíba river basins, in central and north-eastern Brazil, south-eastern Bolivia, Paraguay and northern Argentina. Greatest diversification concentrated in the central Brazilian Plateau (Costa, 2005a). Rivulus modestus Costa, 1991 Rivulus modestus Costa, 1991: 329 (type locality: rio Mutum, rio Tapajós basin, Mato Grosso, Brazil; holotype: MNRJ 11670). Material examined Brazil: Estado de Mato Grosso: MNRJ 11670, holotype; MNRJ 11671, 5 paratypes; UFRJ 2102, 6; UFRJ 2103, 5 (c&s); small stream tributary of rio Mutum, at crossing with road BR-364, 51 km from Vilhena, rio Juruena drainage, rio Tapajós basin; K. Tanizaki, M. T. Lacerda, S. O. Kullander & A. Hogerborn-Kullander, 16 Oct Diagnosis Distinguished from all other congeners by the combination of the following features: anterior portion of trunk deeper than wide, jaws short, snout blunt, tip of anal fin slightly pointed in male, caudal fin rounded in male, pelvic-fin tip reaching anterior portion of anal-fin base in male, dorsal-fin origin on vertical between base of 8th and 9th anal-fin rays, dorsal-fin rays 10-11, anal-fin rays 13-14, frontal squamation F-patterned, frontal scales arranged circularly, canal preopercular absent, contact organs absent, longitudinal series of scales 31-33, gill rakers of first branchial arch , red marks of flank absent, lower jaw black, and round black spot on dorsal portion of caudal-fin in female. Distribution Upper rio Juruena drainage (Fig. 10). Habitat notes Shallow creeks with slow current, in moderately sunny region. Rivulus zygonectes Myers, 1927 Rivulus zygonectes Myers, 1927: 127 (type locality: Vereda Extrema, into Cannabrava [now Cana Brava, at rio Tocantins left bank], Goyas [now Estado de Goiás], Brazil; lectotype: CAS 76314, designated by Huber, 1992: 464). Material examined Brazil: Tocantins-Araguaia basin: Estado de Goiás: CAS 76314, lectotype; Vereda Extrema into Cana Brava; C. Ternetz, 14 Jan UFRJ 1575, 3; stream 15 km E of Aruanã; W. J. E. M. Costa et al., 29 Aug UFRJ 1748, 6; stream tributary to rio Verde, 30 km E of São Miguel do Araguaia; W. J. E. M. Costa et al., 25 Aug UFRJ 1567, 5; stream 98 km W of Jussara; W. J. E. M. Costa et al., 30 Aug UFRJ 1482, 22; UFRJ 2108, 2 (c&s); stream tributary to rio Verde, 32 km N of São Miguel do Araguaia; W. J. E. M. Costa et al., 25 Aug MZUSP 35416, 20; MZUSP 38334, 55; pool in Aruanã; W. J. E. M. Costa et al., 28 Jan Estado do Tocantins: UFRJ 2106, 6; UFRJ 2107, 4 (c&s); stream 20 km S of Santa Rosa; W. J. E. M. Costa et al., 15 Feb MZUSP 38375, 2; stream tributary to córrego Dona Francisquinha, Porto Nacional; W. J. E. M. Costa et al., 31 Jan UFRJ 1712, 7; stream tributary to rio Verde, 50 km N of São Miguel do Araguaia; W. J. E. M. Costa et al., 25 Aug UFRJ 1597, 2; stream 12 km N of Sandolândia; W. J. E. M. Costa et al., 27 Aug MZUSP 37212, 28 (6 c&s); road between road BR-153 and Formoso do Araguaia, 29 km S of Gurupi; P. S. Santos-Filho, Jun MZUSP 45223, 9; rio Água Fria, road Araguaçu- Barreira do Piqui, 27 km N of Araguaçu; F. C. T. Lima, Feb UFRJ 1374, 13; shallow lagoon 2 km W of rio das Mortes at the road Água Boa-Cocalinhos; W. J. E. M. Costa et al., 20 Feb UNT 555, 5; river at the road BR-153 between Filadelphia and Araguaina; E. L. Beerli, 2 Nov UNT 556, 3; rio Brejão, Araguaina; E. L. Beerli & L, M, Lima, 22 Nov UNT 2045, 1; córrego Gorgulho, Porto Nacional; NEAMB, 14 Sep UNT 2046, 1; córrego Gorgulho, Porto Nacional; NEAMB, 14 Sep UNT 2047, 3; córrego Marimbondo, Tupirantins; NEAMB, 27 Oct UNT 2048, 1; córrego Barreiro, Guaraí; NEAMB, 20 Oct UNT 2049, 2; córrego Gorgulho, Porto Nacional; NEAMB, 14 Sep UNT 2050, 1; córrego Gorgulho, Porto Nacional; NEAMB, 14 Sep UNT 2051, 1; córrego Lageado, Paranã; NEAMB, 22 Mar UNT 2052, 9; córrego Água Suja, Tupirantins; NEAMB, 27 Oct UFRJ 5150, 1; floodplains of left bank of rio Tocantins, about 1200 m from the river margin, SSE Sampaio, S W; G. C. Brasil, 8 Jun Estado do Maranhão: UFRJ 5152, 16; swamp about 500 m from the right margin of rio Tocantins, S W; G. C. aqua vol. 11 no

5 Wilson J. E. M. Costa Brasil, 8 Jun Rio Xingu basin, Estado de Mato Grosso: UFRJ 1386, 1; stream 67 km N of Paranatinga; W. J. E. M. Costa et al., 10 Feb UFRJ 1173, 5; stream at the road BR-080, 9 km E of São José do Xingu; W. J. E. M. Costa et al., 16 Feb UFRJ 1417, 1; stream at the road BR- 080, 2 km E of São José do Xingu; W. J. E. M. Costa et al., 18 Feb UFRJ 1389, 6; stream at the road BR-080, 41 km E of São José do Xingu; W. J. E. M. Costa et al., 17 Feb UFRJ 1420, 6; stream at the road BR-080, 42 km E of São José do Xingu; W. J. E. M. Costa et al., 17 Feb UFRJ 1392, 7; stream at the road BR-080, 36 km E of São José do Xingu; W. J. E. M. Costa et al., 17 Feb UFRJ 1357, 20; stream at the road Paranatinga-Canarana, 158 km of Paranatinga; W. J. E. M. Costa et al., 12 Feb UFRJ 1414, 20; UFRJ 2109; stream at the road BR-080, 7 km E of rio Xingu margin; W. J. E. M. Costa et al., 16 Feb UFRJ 6265, 1; stream near rio Xingu, Altamira; W. J. E. M. Costa et al., 16 Jun Rio Tapajós basin, Estado de Mato Grosso: MZUSP 45304, 10; upper rio Preto, road Cuiabá-Santarém; N. Menezes et al., 24 Oct MZUSP 45321, 2; stream tributary to rio Preto, road to São Francisco; N. Menezes et al., 24 Oct Diagnosis Distinguished from all other congeners by the combination of the following features: anterior portion of trunk slightly deeper than wide, jaws short, snout blunt, tip of anal fin slightly pointed in male, caudal fin rounded in male, pelvic-fin tip reaching anterior portion of anal-fin base in male, dorsal-fin origin on vertical between base of 8 th and 10 th analfin rays, dorsal-fin rays 9-11, anal-fin rays 13-15, frontal squamation F-patterned, frontal scales arranged circularly, canal preopercular absent, contact organs absent, longitudinal series of scales 33-35, gill rakers of first branchial arch , oblique rows of red dots sometimes united forming chevron-like bars on flank in male, lower jaw black, and round black spot on dorsal portion of caudal fin in female. Distribution Tocantins, Araguaia, Xingu and Tapajós river basins (Fig. 22). Habitat notes Shallow creeks with slow current, in gallery forest of savanna-like region. 163 Rivulus violaceus Costa, 1991 Rivulus violaceus Costa, 1991: 331 (type locality: rio das Mortes, rio Araguaia-Tocantins basin, Mato Grosso, Brazil; holotype: MNRJ 11672). Material examined Brazil: Estado de Mato Grosso, rio das Mortes basin: MNRJ 11672, holotype; MNRJ 11673, 5 paratypes; UFRJ 2104, 20; rio das Mortes when crossed by BR-163 at km 313; K. Tanizaki, M. T. Lacerda, S. O. Kullander & A. Hogerborn-Kullander, 19 Oct UFRJ 143, 16; UFRJ 2105, 7 (c&s); small stream tributary to rio Perdidos, close to BR-070, about 10 km W of Primavera do Leste; K. Tanizaki, M. T. Lacerda, S. O. Kullander & A. Hogerborn-Kullander, 19 Oct UFRJ 1207, 4; rio Suspiro, 57 km S of Paranatinga; W. J. E. M. Costa, C. P. Bove, R. D. Cunha & C. Muratori, 9 Feb UFRJ 4284, 13; idem; M. Britto, C. Moreira & R. Cunha, 28 Oct UFRJ 4283, 3; stream at the road Primavera do Leste-Paranatinga; M. Britto, C. Moreira & R. Cunha, 28 Oct Diagnosis Distinguished from all other congeners by the combination of the following features: anterior portion of trunk slightly deeper than wide, jaws short, snout blunt, tip of anal fin slightly pointed in male, caudal fin rounded in male, pelvic-fin tip reaching urogenital papilla in male, dorsal-fin origin on vertical between base of 8 th and 9 th anal-fin rays, dorsalfin rays 10-11, anal-fin rays 14-15, frontal squamation F-patterned, frontal scales arranged circularly, canal preopercular absent, contact organs absent, longitudinal series of scales 30-32, gill rakers of first branchial arch , oblique chevron-like red bars on flank in male, lower jaw black, and round black spot on dorsal portion of caudal fin in female. Distribution Upper rio das Mortes drainage (Fig. 10). Habitat notes Shallow creeks with slow current, in gallery forest of savanna-like region. Rivulus litteratus Costa, 2005 Rivulus sp.: Lacerda, 1989: 25 (Ponte Branca, Rio Araguaia, central Brazil). aqua vol. 11 no

6 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin Rivulus pictus (non Costa): Costa et al., 2003: 143 (misidentification of specimens from Alto Araguaia). Rivulus litteratus Costa, 2005a: 75 (type locality: Município de Alto Araguaia, Córrego do Sapo, upper Rio Araguaia basin, road MT-100, 31 km S of Alto Araguaia, 17º S 53º W, altitude 750 m, Estado de Mato Grosso, Brazil; holotype: UFRJ 5956). Material examined Brazil: Estado de Mato Grosso: Município de Alto Araguaia, upper rio Araguaia basin: UFRJ 5956, holotype; Município de Alto Araguaia, córrego do Sapo, upper rio Araguaia basin, road MT-100, 31 km S of Alto Araguaia; W. J. E. M. Costa, B. B. Costa and C. P. Bove, 14 Jan Diagnosis Distinguished from all other congeners by the combination of the following features: anterior portion of trunk slightly deeper than wide, jaws short, snout blunt, tip of anal fin slightly pointed in male, caudal fin rounded in male, pelvic-fin tip reaching anterior portion of anal-fin base in male, dorsal-fin origin on vertical between base of 7 th 8 th l-fin rays, dorsal-fin rays 9-10, anal-fin rays 13-14, frontal squamation F-patterned, frontal scales arranged circularly, canal preopercular absent, contact organs absent, longitudinal series of scales 31-32, gill rakers of first branchial arch , overlapped red marks of variable shape on flank in male, lower jaw dark grey, and round black spot on dorsal portion of caudal-fin in female. Also distinguished from the remaining species of Melanorivulus by possessing a colour pattern consisting of red marks variable in shape and highly overlapped (Fig. 43). Distribution Upper rio Araguaia drainage (Fig. 23). Habitat notes Shallow creeks with slow current, in gallery forest of savanna-like region (Fig. 45). Discussion The present study confirms a weakly supported genus Rivulus containing clades corroborated both by morphological (e.g. Huber 1992, Costa 1998a, the present study) and molecular data (e.g. Murphy et al. 1999, rbek & Larson 1999), and recognized as subgenera. No unambiguous, apomorphic morphological condition was found to be shared by all species presently assigned to Rivulus and molecular studies indicate that Rivulus is a paraphyletic assemblage (Murphy et al. 1999, Hrbek & Larson 1999). However, a group comprising all species of Rivulus except the subgenus Rivulus is well supported by morphology, their members having a uniquely derived pattern of arrangement of rostral neuromasts (i.e. transversely placed), and a long neural prezygapophysis in caudal vertebrae as already discussed in former studies (e.g. Costa 1998a), also occurring in Kryptolebias. Fig. 43. Rivulus litteratus, UFRJ 5956, male, holotype, 26.9 mm SL (one day after collection); Brazil: Mato Grosso: Alto Araguaia. Photo by W. J. E. M. Costa. Fig. 44. Rivulus litteratus, UFRJ 5957, female, paratype, 25.6 mm SL (one day after collection); Brazil: Mato Grosso: Alto Araguaia. Photo by W. J. E. M. Costa. Fig. 45. Brazil: Mato Grosso: Alto Araguaia; savanna creek close to córrego do Sapo, habitat of Rivulus litteratus. Photo by W. J. E. M. Costa. aqua vol. 11 no

7 Wilson J. E. M. Costa The subgenus Rivulus, as herein defined, is also corroborated by a molecular analysis (Murphy et al. 1999: 295). In other molecular analyses, R. roloffi is considered to be more closely related to other nominal species of Rivulus than to R. cylindraceus (e.g. Hrbek & Larson 1999). As R. roloffi, which was not accurately examined in the present study (only examined at the field just after collection), have more than half caudal fin scaled and a dark green humeral spot, which constitute two synapomorphies of the subgenus Rivulus, R. roloffi is inserted in this subgenus. Although weakly supported by morphological features, Cynodonichthys as proposed here is also corroborated by molecular data (Murphy et al. 1999: 295). However, some molecular analyses do not support monophyly of this assemblage (e.g. Hrbek & Larson 1999). Further analyses should be conducted in order to test the monophyly of this geographically widespread assemblage. Three species of Cynodonichthys from the Brazilian Amazonas basin were found in collections examined in the present study: R. urophthalmus, R. taeniatus and R. micropus. A fourth species, R. xanthonotus, has been reported for the Brazilian Amazon. However, this species, which seems to be similar to R. urophthalmus, is only known from its poor original description (Ahl 1926), where the type locality is inaccurately described (i.e. Amazon Strom ). Huber (1992) reported that types exhibit an uncommon posteriorly positioned dorsal fin, thus constituting a valid species. No specimen potentially identifiable as R. xanthonotus was examined in the present study. Rivulus urophthalmus was found in an extensive region encompassing the lower Amazonas basin and tributaries, and adjacent coastal areas to East, including Maranhão, the type locality R. auratus. This latter species is therefore considered a synonym of R. urophthalmus, as already proposed by Huber (1992). Rivulus poey was described by Steindachner (1876) based on specimens from Belém. The brief description includes characters congruent with R. urophthalmus, previously described from the same locality. As the types of R. poey are unknown, this highly probable synonymy cannot be confirmed. In order to solve this problem, a neotype from the city of Belém is herein designated for R. poey, becoming a clear synonym of R. urophthalmus, as already proposed by other authors (e.g. Garman 1895, Huber 1991). A geographically widespread species of the central and western Brazilian Amazon is here tentatively identified as R. taeniatus, a species first described from the Colombian Amazon and still poorly known. However, the species collected in some distant points of the Brazilian Amazon also fits well with other nominal species from the Peruvian and Ecuadorian Amazon, such as R. rubrolineatus and R. limoncochae, which possibly are synonyms of R. taeniatus. These species have been distinguished mainly on the basis of the number of rows of red dots on the caudal peduncle (i.e. three or five) (e.g. Huber 1992). However, this character was variable within populations both of R. urophthalmus and R. taeniatus (see descriptions above). Steindachner (1863) described R. micropus based upon a single specimen collected in the rio Negro, Brazilian Amazon. The identity of this species was obscure until publication of photos and new morphological data on the holotype (Huber 1991). Henn (1916) described a similar species, R. compressus, from Manaus, a city at the confluence of rio Negro and rio Solimões, central Brazilian Amazon. At the beginning of the description of R. compressus, Henn (1916: 111) noted that: This may be R. micropus Steidachner, but it seems to differ in the more forward position of the dorsal, the longer head, etc.. However, Huber (1992) examined types of both species and concluded that these diagnostic characters are similar in both nominal species. This synonymy follows Costa (2003b), supported by material collected in the type locality region, having pointed snout and posteriorly positioned dorsal fin as observed in holotypes of R. micropus (Huber, 1991: fig. 2) and R. compressus (Henn, 1916: fig. 1). Anablepsoides was first proposed to include only R. atratus (Huber, 1992). However, morphological data (Costa 1998a, the present study) consistently corroborated a clade containing both R. atratus and R. ornatus. This hypothesis has not yet been tested by molecular analyses, since both included species were not simultaneously included in these studies (e.g. Murphy et al. 1999, Hrbek & Larson 1999). Examination of types revealed that R. ornatus and R. obscurus, respectively, refer to the male and female of the same species, thus constituting synonyms (Costa 2003b). Rivulus ornatus has been identified as R. obscurus (e.g. Huber 1992), whereas an undescribed species from Peru has been misidentified as R. ornatus (e.g. Fels & de Rham 1982, Huber 1992). The specimen mentioned by Garman (1895: 139) under the description of R. ornatus, as possibly belonging to another species, is in fact a member of the poeciliid genus Fluviphylax Whitley (Costa 1996, Costa & Le Bail 1999). Benirivulus is erected to R. beniensis, which does not 165 aqua vol. 11 no

8 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin fit with any other subgenus of Rivulus. The present phylogenetic hypothesis indicates that Benirivulus is the sister group to a clade including three subgenera, Laimosemion, Owiyeye, and Melanorivulus, each of them corroborated by both morphological and molecular data. The only species of Benirivulus, R. beniensis, was poorly known until now, and consequently was not included in molecular analyses. Myers (1927b) erected R. beniensis on the basis of specimens collected in the drainage of the Beni- Mamoré drainage, rio Madeira basin, and previously misidentified by Pearson (1925) as R. strigatus. Based on putative minor differences in dorsal-fin origin position, Myers (1927b) recognized two subspecies from distinct localities, R. beniensis beniensis and R. beniensis lacustris. Seegers (1985) described R. bolivianus also from the río Mamoré drainage. Examination of material from distant localities along the rio Madeira basin, including the Mamoré, Beni and Guaporé river drainages, revealed that R. beniensis is a widespread species, indistinguishable from R. beniensis lacustris and R. bolivianus, thus confirming the synonymy proposed by Costa (2003b). Huber (1999) erected Laimosemion to include species of the R. geayi superspecies as proposed by Huber (1992). Molecular studies (Murphy et al. 1999, Hrbek et al. 2004) corroborated monophyly of a group comprising species of the R. geayi superspecies, R. breviceps superspecies, and R. frenatus superspecies, which is in accordance to the present study. Two species of Laimosemion are found in the Brazilian Amazonas river: R. strigatus and R. dibaphus. Rivulus strigatus was described by Regan (1912) based on a single specimen donated by Arnold. This material was obtained from an aquarium fish shipment from the city of Pará (now Belém, capital of Estado do Pará) (Arnold 1913). Huber (1992) mentioned rio da Prata, a tributary of rio Jari, Pará, Brazil, as the true type locality of R. strigatus, based on a misinterpreted citation by von Ihering (1931). The latter author, under a brief diagnosis of R. strigatus, cited: Distribuição: (typo) Amazonas, sem mais indicação além do nome do colleccionador Arnold, mas cujo material provem principalmente do rio da Prata (Distribution: (type) Amazonas, without further indication besides the collector name Arnold, but that material mainly originates from rio da Prata) (von Ihering 1931: 263). In fact, von Ihering referred to the río de La Plata basin, from where R. punctatus Boulenger was often misidentified as R. strigatus in the past (e.g., Ringuelet et al. 1967), not to the small rio da Prata in Pará, unknown for most ichthyologists and still almost inaccessible at present. The geographic position of the rio da Prata mentioned by von Ihering may be clarified when examining other parts of his paper. For example, among other localities, he cited the occurrence of Phalloceros caudimaculatus (Hensel) for the rio da Prata, Paraguay (von Ihering 1931: 246). Huber (1992) considered both R. strigatus and R. dibaphus as synonyms of R. geayi, a species first described from Amapá, northern Brazil, but also occurring in the Guianas (e.g. Huber 1991, 1992). According to Huber (1991), the three nominal species would have near type localities, but in fact the type locality of R. geayi is about 480 km and 560 km in a straight line from the type locality of R. strigatus and R. dibaphus, respectively, and type locality of R. strigatus is about 710 km from the type locality of R. dibaphus. Comparison of material of R. strigatus with material from the type locality region of R. geayi revealed that R. strigatus is a distinct and valid species, easily distinguished from R. geayi by having a rounded caudal fin in the male (vs. subtruncate) and frontal squamation F-patterned (vs. usually E-patterned). Rivulus dibaphus is also a valid species. Examination of the type material of R. dibaphus and material collected in the type locality region revealed that it may be readily distinguished from R. strigatus and R. geayi by the unique colour pattern on the caudal peduncle in the male (Figs 31-32) and that the frontal squamation is always E-patterned. Owiyeye corresponds to the R. rectocaudatus species group diagnosed by Thomerson et al. (1992) and the R. rectocaudatus superspecies diagnosed by Huber (1992). This clade is well supported both by morphological (the present study) and molecular data (Murphy et al. 1999, Hrbek & Larson 1999, Hrbek et al. 2004). Owiyeye and Anablepsoides share two derived conditions (frontal squamation S-patterned and a transverse stripe through chin), but it is parsimoniously considered to a member of a clade also including Laimosemion and Melanorivulus, mainly diagnosed by the absence of second pharyngobranchial teeth. Seven species of Owiyeye are found in the Brazilian Amazon, all described in recent years: R. romeri, R. kirovskyi, R. uatuman, R. uakti, R. amanapira, and R. tecminae. Rivulus kirovskyi and R. duckensis are identical and were collected at the same locality. Both original descriptions were published in 2004, but the name R. kirovskyi has chronological priority over R. duckensis, since the former was published on 30 April, the latter on 20 August. Melanorivulus corresponds to the R. punctatus aqua vol. 11 no

9 Wilson J. E. M. Costa superspecies diagnosed by Huber (1992) or the Rivulus punctatus species-complex diagnosed by Costa (1995a). This clade is strongly supported by morphology, and is also corroborated by molecular studies (e.g. Murphy et al. 1999, Hrbek & Larson 1999). It includes four species endemic to the Brazilian Amazonas basin: R. modestus, R. zygonectes, R. violaceus, and R. litteratus. As already noted in previous studies (e.g. Costa 1998a, Murphy et al. 1999, Hrbek & Larson 1999), species of Rivulus endemic to the Amazonas basin do not form a monophyletic assemblage. Species from southern Amazonas tributaries, running on the Brazilian Shield (i.e., belonging to Melanorivulus) are more closely related to species endemic to basins to South and East (i.e., Paraná, Paraguay, São Francisco and Parnaíba river basins). On the other hand, species occurring in northern and western Amazonas tributaries arising from the Andes or the Guyana Shield (i.e., belonging to Cynodonichthys, Laimosemion and Owiyeye) are more closely related to species endemic to the Orinoco and Guianas basin. This is probably a historic consequence of the past isolation of rivers draining the Brazilian Shield before the formation of the present Amazonas basin, as described in geological studies (e.g. Beurlen 1970, Lundberg et al. 1998). Field observation on the habitat of Amazonian species of Rivulus revealed that different lineages occupy distinct kinds of habitat. This is most clear for the clade Benirivulus + Laimosemion + Owiyeye + Melanorivulus, in which members of each genus are found in a particular kind of aquatic environment, thus suggesting divergent specializations during the evolution of the group. In Benirivulus, individuals were collected near the margin of stagnant streams and lakes (about cm deep), with dense aquatic vegetation, at the forest border (Fig. 25). Species of Laimosemion, Owiyeye and Melanorivulus were collected in shallower places (about 5-40 cm). Possibly, the ecological preference noted for Benirivulus is the primitive condition for the clade Benirivulus + Laimosemion + Owiyeye + Melanorivulus, since species of Anablepsoides, the putative sister group to the above mentioned clade, live in similar habitats. Amazonian species of Laimosemion were uniquely found in clear water streams, with moderate current and sandy bottom (Figs 29 and 33). No other species from the Amazonas basin inhabits a similar environment, but R. xiphidius and R. geayi, two species of Laimosemion from Guyana, seem to live in the same environmental conditions (Huber 1979). However, R. agilae, another species of Laimosemion from Guyana, has been reported as living in sunny places (Huber 1979). Only species of Owiyeye were found in extremely shallow temporary pools (about 5-10 cm deep) with tea-coloured water (Figs 36 and 41). Since some species of Owiyeye were uniquely found in temporary pools, they possibly are semiannual fishes. An exception is R. uakti, which inhabit marginal zones of creeks at the forest border (Fig. 39). Species of Melanorivulus are found in sunny creeks (Fig. 45) in savanna-like environments. However, this is not a unique condition among Amazonian Rivulus, since R. urophthalmus, R. taeniatus, and R. micropus are also found in sunny creeks, often near the border of forests (Figs 11, 17 and 20). These conditions are interpreted as independently derived, because Melanoorus and Cynodonichthys are not closely related taxa, and basal rivulids (i.e. basal species of Kryptolebias and Prorivulus) inhabit dark forested habitats (Costa 2004a, 2004c, Vermeulen & Hrbek 2005). Acknowledgements Thanks are due to E. Araujo, F. Autran, S. Barrera, D. Belote, C. P. Bove, G. C. Brasil, M. Britto, C. Campinha, B. B. Costa, R. D. Cunha, R. D Arrigo, M. I. Landim, S. M. Q. Lima, C. Moreira, R. Paiva, A. Pinto, D. Ramos, A. Sarraf, K. Tanizaki, E. Vicente, and L. Villa-Verde, for help in collecting trips; to H. Berkenkamp, G. Brasil, H. Britski, J. Carvalho, D. Catania, J. L. Figueiredo, K. Hartel, J. Huber, M. Kottelat, S. Kullander, M. Lacerda, P. Lucinda, N. Menezes, D. Moraes Jr., G. Nunan, O. Oyakawa, L. Parenti, C. Pereira, L. Py-Daniel, U. Römer, J. Sarmiento, D. Taphorn, D. Teixeira, and J. Zuanon, for the loan, exchange, or donation of specimens, or for support during visits to their institutions. Collecting trips were supported by Fundação o Boticário de Proteção à Natureza. This study was funded by CNPq-MCT and FAPERJ. Material was collected with permits /98 and /02 from IBAMA (Instituto Brasileiro do Meio Ambiente e dos Recursos Naturais Renováveis Ministério do Meio Ambiente, dos Recursos Hídricos e da Amazônia Legal). References ARNOLD, J. P Rivulus strigatus, sp. n. Rgn. Wochenschrift für Aquarien-und Terrarienkunde 10: BERKENKAMP, H. O. &. ETZEL, V Die Rivulus-Arten 167 aqua vol. 11 no

10 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin von Panama 2: Wiederbeschreibung von Rivulus brunneus Meek & Hildebrand, 1913 und Beschreibung von Rivulus birkhahni spec. nov. von Chiquiri Grande, West-Panama. Deutsche Killifisch Gemeinschaft Journal 24: BERKENKAMP, H. O. &. ETZEL, V. 1993a. Die Rivulus-Arten von Panama 3: Rivulus kuelpmanni spec. nov., eine neue Bachlingsart von Punta Peña, West-Panama. Deutsche Killifisch Gemeinschaft Journal 25: BERKENKAMP, H. O. &. ETZEL, V. 1993b. Die Rivulus-Arten von Panama 4: Rivulus frommi spec. nov., eine neue Bachlingsart von El Valle, Zentral-Panama. Deutsche Killifisch Gemeinschaft Journal 25: BERKENKAMP, H. O. &. ETZEL, V Die Rivulus-Arten von Panama 5: Rivulus monikae spec. nov., eine weitere neue Bachlingsart aus den bewaldeten und bergigen Teilen der Provinz Coclé, Zentral-Panama. Deutsche Killifisch Gemeinschaft Journal, 27: 1-7. BERKENKAMP, H. O. &. ETZEL, V Die Rivulus-Arten von Panama 6: Zur Identität von Rivulus montium Hildebrand, 1938 mit der Beschreibung einer neuen Art, Rivulus villwocki spec. nov. Deutsche Killifisch Gemeinschaft Journal 29: BERKENKAMP, H. O. &. ETZEL, V Die Rivulus-Arten von Panama 7: Rivulus wassmanni spec. nov., eine neue Bachlingsart der Rivulus birkhahni-kuelpmanni-gruppe aus der Provinz Bocas del Toro, Nordost-Panamá. Aquaristik Aktuell 1999: BEURLEN, K Geologie von Brasilien. Bornträger, Berlin, 444 pp. BUSSING, W. A Status of the cyprinodontid fish genus Rivulus in Costa Rica, with descriptions of new endemic species. Brenesia 17: COSTA, W. J. E. M Descrição de cinco novas espécies de Rivulus das bacias dos rios Paraná e São Francisco (Cyprinodontiformes, Rivulidae). Revista Brasileira de Zoologia 6: COSTA, W. J. E. M Análise filogenética da família Rivulidae (Cyprinodontiformes, Aplocheiloidei). Revista Brasileira de Biologia 50: COSTA, W. J. E. M. 1991a. Redescrição do gênero Rivulus (Cyprinodontiformes: Rivulidae), com notas sobre R. stellifer e R. compactus e a descrição de duas novas espécies do Brasil central. Revista Brasileira de Biologia 51: Costa, W. J. E. M. 1991b. Description of two new species of the genus Rivulus (Cyprinodontiformes: Rivulidae) from eastern South American coastal plains. Revue Suisse de Zoologie 98: COSTA, W. J. E. M. 1995a. Revision of the Rivulus punctatus species-complex (Cyprinodontiformes: Rivulidae). Ichthyological Exploration of Freshwaters 3: COSTA, W. J. E. M. 1995b. Pearl killifishes the Cynolebiatinae: systematics and biogeography of the neotropical annual fish subfamily. TFH, Neptune City, 128 pp. COSTA, W. J. E. M Relationships, monophyly and three new species of the neotropical miniature poeciliid genus Fluviphylax (Cyprinodontiformes: Cyprinodontoidei). Ichthyological Exploration of Freshwaters 7: COSTA, W. J. E. M. 1998a. Phylogeny and classification of Rivulidae revisited: evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei). Journal of Comparative Biology 3: COSTA, W. J. E. M. 1998b. Phylogeny and classification of the Cyprinodontiformes (Euteleostei: Atherinomorpha): a reappraisal. In: Malabarba, L. R., Reis, R. E., Vari, R. P., Lucena, Z. M. S. & Lucena, C. A. S. (Eds.). Phylogeny and classification of neotropical fishes, Edipucrs, Porto Alegre, pp COSTA, W. J. E. M The neotropical annual fish genus Cynolebias (Cyprinodontiformes: Rivulidae): phylogenetic relationships, taxonomic revision and biogeography. Ichthyological Exploration of Freshwaters 12: COSTA, W. J. E. M. 2003a. Family Rivulidae (South American annual fishes). In: R. E. Reis, Kullander, S. O. & Ferraris, C. J., Jr. (eds) Check list of the freshwater fishes of South and Central America, Edipucrs, Porto Alegre, Pp COSTA, W. J. E. M. 2003b. A new miniature rivuline fish from the upper Negro river basin, northern Brazil (Teleostei, Cyprinodontiformes, Rivulidae). Arquivos do Museu Nacional, Rio de Janeiro 61: COSTA, W. J. E. M. 2003c. Rivulus paracatuensis n. sp. (Cyprinodontiformes: Rivulidae): a new rivuline species from the Rio São Francisco basin, Brazil. aqua Journal of Ichthyology and Aquatic Biology 7: COSTA, W. J. E. M. 2004a. Relationships and redescription of Fundulus brasiliensis (Cyprinodontiformes: Rivulidae), with description of a new genus and notes on the classification of the Aplocheiloidei. Ichthyological Exploration of Freshwaters 15: COSTA, W. J. E. M. 2004b. Kryptolebias, a substitute name for Cryptolebias Costa, 2004 and Kryptolebiatinae, a substitute name for Cryptolebiatinae Costa, 2004 (Cyprinodontiformes: Rivulidae). Neotropical Ichthyology 2: COSTA, W. J. E. M. 2004c. A new killifish genus and species from the coastal plains of northeastern Brazil (Teleostei: Cyprinodontiformes: Rivulidae). Zootaxa 642: COSTA, W. J. E. M. 2004d. Rivulus kirovskyi, a new killifish from the central Amazon, Brazil (Cyprinodontiformes: Rivulidae). Neotropical Ichthyology 2: COSTA, W. J. E. M. 2004e. Rivulus uatuman sp. n. (Teleostei: Cyprinodontiformes: Rivulidae): a new miniature killifish from the central Amazon. Zootaxa 696: 1-8. COSTA, W. J. E. M. 2004f. Rivulus uakti sp. n. and R. amanapira sp. n. (Teleostei: Cyprinodontiformes: Rivulidae): two new species from the upper Rio Negro, Brazilian Amazon. Zootaxa 465: COSTA, W. J. E. M. 2004g. Rivulus simplicis n. sp. (Cyprinodontiformes: Rivulidae): a new killifish from the coastal plains of south-eastern Brazil. aqua Journal of Ichthyology and Aquatic Biology 8: COSTA, W. J. E. M. 2005a. Seven new species of the killifish genus Rivulus (Cyprinodontiformes: Rivulidae) from the Paraná, Paraguay and upper Araguaia river basins, central Brazil. Neotropical Ichthyology 3: COSTA, W. J. E. M. 2005b. The Neotropical annual killifish genus Pterolebias Garman (Teleostei: Cyprinodontiformes: Rivulidae): phylogenetic relationships. descriptive morphol- aqua vol. 11 no

11 Wilson J. E. M. Costa ogy, and taxonomic revision. Zootax 1067: COSTA, W. J. E. M. & BRASIL, G. C Description of a new species of Rivulus (Cyprinodontiformes: Rivulidae) from the coastal plains of eastern Brazil. Ichthyological Exploration of Freshwater 1: COSTA, W. J. E. M. & LE BAIL, P.-Y Fluviphylax palikur: a new poeciliid from the rio Oiapoque basin, northern Brazil (Cyprinodontiformes: Cyprinodontoidei), with comments on miniaturization in Fluviphylax and other Neotropical freshwater fishes. Copeia 1999: ETZEL, V. & BERKENKAMP, H. O Die Rivulus-Arten von Panama 1: Untersuchungen zur identität von Rivulus chucunaque Breder, Deutsche Killifisch Gemeinschaft Journal 24: FELS, J. F. & HUBER, J. H Descriptión d une espèce nouvelle de Rivulus du Pérou, R. derhami n. sp. et nouvelle description de R. beniensis Myers, 1927 (Pisces, Cyprinodontidae). Revue Française d Aquariologie 12: FELS, J. F. & DE RHAM, P Récentes collectes de Rivulus (Cyprinodontidés) au Pérou, avec description de six espèces nouvelles, deuxième partie. Revue Française d Aquariologie 8: FELSENSTEIN, J Confidence limits on phylogenies: an approach using the bootstrap. Evolution 39: FOWLER, H. W Freshwater fishes from North-western Colombia. Proceedings of the Academy of Natural Sciencies of Philadelphia 96: GARMAN, S The cyprinodonts. Memoirs of the Museum of Comparative Zoology 19: HENN, A. W IX On various South American poeciliid fishes. Annals of the Carnegie Museum 10: plates. HOEDEMAN, J. J The frontal scalation pattern in some groups of toothcarps (Pisces, Cyprinodontiformes). Bulletin of Aquatic Biology 1: HOEDEMAN, J. J Rivulid fishes of Suriname and other Guyanas, with a preliminary review of the genus Rivulus. Studies on the Fauna of Suriname and Other Guyanas 3: HOEDEMAN, J. J Studies on Cyprinodontiform fishes, 8: Preliminary key to the species and subspecies of the genus Rivulus. Bulletin of Aquatic Biology 2: HRBEK, T., DE DEUS, C. P. & FARIAS, I. P Rivulus duckensisi (Teleostei; Cyprinodontiformes): new species from the Tarumã basin of Manaus, Amazonas, Brazil, and its relationships to other neotropical Rivulidae. Copeia 2004: HRBEK, T. & LARSON, A The diapause in the killifish family Rivulidae (Atherinomorpha, Cyprinodontiformes): a molecular phylogenetic and biogeographic perspective. Evolution 53: HUBER, J. H A propos de quatre nouvelles collections de Rivulus des Guyanes, avec description de Rivulus xiphidius n. sp. et Rivulus amphoreus n. sp. Revue Française d Aquariologie 6: HUBER, J. H Revue des espèces de Rivulus de Guyane française, avec descriptions de Rivulus cladophorus n. sp. et de R. igneus n. sp., et nouvelles descriptions de R. urophthalmus Günther et de R. micropus Steindachner. Revue Française d Aquariologie 18: HUBER, J. H Review of Rivulus: ecobiogeografia - relationships. Cybium, Société Française d Ichtyologie, Paris, pp. HUBER, J. H Updates to the phylogeny and systematics of the Neotropical cyprinodont genus Rivulus and its allied (Cyprinodontiformes: Rivulinae). Cybium 23: KITCHING, I. J., FOREY, P. L., HUMPHRIES, C. J. & WILLIAMS, D. M Cladistics, the theory and practice of parsimony analysis (2nd edition), Oxford University Press, Oxford- New York-Tokyo, 228 pp. LASSO, C., TAPHORN, D. C. & THOMERSON, J. E Rivulus gransabanae, a new species of killifish from Venezuela (Cyprinodontiformes: Rivulidae). Ichthyological Exploration of Freshwaters 2: LUNDBERG, J. G., MARSHALL, L. G., GUERRERO, J., HOR- TON, B., MALABARBA, M. C. S. L. & WESSELINGH, F The stage for Neotropical fish diversification: a history of tropical South American rivers. In: Malabarba, L.R., Reis, R. E., Vari, R. P., Lucena, Z. M. S. & Lucena, C. A. S. (Eds.). Phylogeny and classification of neotropical fishes, Edipucrs, Porto Alegre, pp MEEK, S. E The freshwater fishes of Mexico North of the Isthmus of Tchuantepec. Publications of the Field Columbian Museum, Zoology 5: MENEZES, N. A. & WEITZMAN, S. H Two new species of Mimagoniates (Teleostei: Characidae: Glandulocaudinae), their phylogeny and biogeography and a key to glandulocaudin fishes of Brazil and Paraguay. Proceedings of the Biological Society of Washington 103: MURPHY, W. J. & COLLIER, G. E Phylogenetic relationships within the aplocheiloid fish genus Rivulus (Cyprinodontiformes, Rivulidae): Implications for Caribbean and Central American Biogeography. Molecular Biology and Evolution 14: MURPHY, W. J. & COLLIER, G. E A molecular phylogeny for aplocheiloid fishes (Atherinomorpha, Cyprinodontiformes): the role of vicariance and the origins of annualism. Molecular Biology and Evolution 14: MURPHY, W. J., THOMERSON, J. E. & COLLIER, G. E Phylogeny of the neotropical killifish family Rivulidae (Cyprinodontiformes, Aplocheiloidei) inferred from mitochondrial DNA sequences. Molecular and Phylogenetic Evolution 13: MYERS, G. S. 1927a. An analysis of the genera of neotropical killifishes allied to Rivulus. Annals and Magazine of Natural History, ser. 9 19: MYERS, G. S. 1927b. Descriptions of new South American fresh-water fishes collected by Dr. Carl Ternetz. Bulletin of the Museum of Comparative Zoology, Harvard 68: PARENTI, L. R A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha). Bulletin of the American Museum of Natural History 168: PEARSON, N. E The fishes of the Rio Beni basin, Bolivia, collected by the Mullford Expedition. Indiana Uni- 169 aqua vol. 11 no

12 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin versity Studies 64: POEY, F Poissons de Cuba, espèces nouvelles. Memorias sobre la Historia Natural de la Isla de Cuba, 2: RAMOS, T. C Tree Gardener (computer program), Version 2.3beta. Univ. São Paulo, São Paulo. REGAN, C. T VI Diagnoses of new Central-American freshwater fishes of the families Cyprinodontidae and Mugilidae. Annals and Magazine of Natural History, ser. 7 19: REGAN, C. T A revision of the poeciliid fishes of the genera Rivulus, Pterolebias, and Cynolebias. Annals and Magazine of Natural History, ser. 8 10: REIS, R. E Revision of the neotropical catfish genus Hoplosternum (Ostariophysi: Siluriformes: Callichthyidae), with the description of two new genera and three new species. Ichthyological Exploration of Freshwaters, 7: RINGUELET, R. A., ARAMBURU, R. H. & ARAMBURU, A Los peces argentinos de agua dulce. Comisión de Investigaciones Científicas de la Provincia de Buenos Aires, La Plata, 602 pp. SEEGERS, L Zur Revision der Rivulus-Arten Südost- Brasiliens, mit einer Neubeschreibung von Rivulus luelingi n. sp. und Rivulus caudomarginatus n. sp. (Pisces: Cyprinodontidae: Rivulinae). Zoologische Beiträge 28: SEEGERS, L Ein neuer Rivulide aus Bolivien: Rivulus bolivianus n. sp. (Pisces: Cyprinodontiformes: Rivulidae). Bonner Zoologische Beitrage 39: STEINDACHNER, F Beiträge zur Kenntniss der Sciaenoiden Brasiliens und der Cyprinodonten Mejicos. Sitzungsberichte der kaiserlichen Akademie der Wissenschaften Classe K. 48: , plates 1-4. SWOFFORD, D. L PAUP*. Phylogenetic analysis using parsimony (*and other methods), Beta version 4.0b2. Sinauer, Sunderland, Massachusetts. TAYLOR, W. R. & VAN DYKE, G. C Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium 9: THOMERSON, J. E., BERKENKAMP, H. O. & TAPHORN, D. C. 1991a. Rivulus lyricauda, a new species from the Guyana Shield in eastern Venezuela (Cyprinodontiformes: Rivulidae). Ichthyological Exploration of Freshwaters 1: THOMERSON, J. E., NICO, L. G. & TAPHORN, D. C. 1991b. Rivulus immaculatus, a new killifish from Venezuela (Cyprinodontiformes, Rivulidae). Copeia 1991: THOMERSON, J. E., NICO, L. G. & TAPHORN, D. C Rivulus tecminae, a new killifish from Amazonas Territory, Venezuela (Cyprinodontiformes: Rivulidae). Ichthyological Exploration of Freshwaters 2: THOMERSON, J. E. & TAPHORN, D. C Two new annual killifishes from Amazonas Territory, Venezuela (Cyprinodontiformes: Rivulidae). Ichthyological Exploration of Freshwaters 3: WALLACE, A. R Peixes do Rio Negro/Fishes of the Rio Negro. EDUSP, São Paulo, 517 pp. WEITZMAN, S. H. & WOURMS, J. P South American cyprinodont fishes allied to Cynolebias with the description of a new species of Austrofundulus from Venezuela. Copeia 1967: WIENS, J. J. (ed.) Phylogenetic analysis of morphological data, Smithsonian Institution Press, Washington and London, 220 pp. VERMEULEN, F. B. M. & HRBEK, T Kryptolebias sepia n. sp. (Actinopterygii: Cyprinodontiformes: Rivulidae), a new killifish from the Tapanahony River drainage in southeast Surinam. Zootaxa 928: VERMEULEN, F. B. M. & ISBRÜCKER, I. J. H Rivulus torrenticola n. sp. (Actinopterygii: Cyprinodontiformes: Rivulidae), a new killifish from highlands in the Guyana Shield. Beaufortia 50: VON IHERING, R Cyprinodontes brasileiros (peixes guarús ): sistemática e informações biologicas. Archivos do Instituto Biologico 4: aqua vol. 11 no

13 Wilson J. E. M. Costa Appendix 1 The list below includes material examined, except belonging to species of the Brazilian Amazonas basin, which are listed in the Taxonomic accounts above. Data on material is organized in the following sequence: catalogue number, number of specimens, locality. Abbreviations are: c&s, specimens cleared and stained for bone and cartilage, H, holotype, and P, paratype(s). Institutional acronyms are listed in material and methods. Aplocheilidae: Aplocheilus panchax: UFRJ 3140, 2; UFRJ 3141, 2 (c&s); Indonesia: Sulawesi: Desa Radda. Nothobranchiidae: Scriptaphyosemion guignardi: UFRJ 3883, 8; UFRJ 4110, 4 (c&s); Guinea: Dalaba. Rivulidae: Aphyolebias boticarioi: UFRJ 5986, H; UFRJ 5987, 11 P; UFRJ 5988, 5 P; Brazil: Estado do Acre: Porto Acre. Gnatholebias hoignei: MCNG 1116, 18; Venezuela: Portuguesa: La Trinidad. UFRJ 6116, 6; UFRJ 6117, 5 (c&s); Venezuela: Portuguesa: Papelón. Kryptolebias brasiliensis: UFRJ 3458, 32; UFRJ 3682, 1 (c&s); UFRJ 4603, 2 ex. (c&s); UFRJ 5332, 6 ex. (c&s); Brazil: Rio de Janeiro: Magé. Micromoema xiphophora: MCNG 26440, 20; Venezuela: Amazonas: Isla Ratón. UFRJ 3165, 1 (c&s); aquarium material. Moema apurinan: UFRJ 5980, H; UFRJ 5981, 9 P; UFRJ 5982, 7 P (c&s); Brazil: Estado do Acre: Porto Acre. Neofundulus paraguayensis: UFRJ 3647, 10; UFRJ 3648, 4 (c&s); Brazil: Mato Grosso do Sul, about 70 km NW from de Aquidauana. Pituna compacta: UFRJ 3563, 33; UFRJ 3564, 4 (c&s); Brazil: Tocantins: Barreira do Piqui. Prorivulus auriferus: UFRJ 5932, H; UFRJ 5933, 4P; UFRJ 5934, 3P (c&s); Brazil: Bahia: Valença. Rachovia maculipinnis: MCNG 35565, 2; Venezuela: Portuguesa: Sabaneta. UFRJ 6118, 7; UFRJ 6119, 4 (c&s); Venezuela: Portuguesa: Papelón. R. stellifer: MCNG 25828, 9; Venezuela: Portuguesa: La Capilla. UFRJ 245, 5 (c&s); Venezuela: Cojedes: 1 km N of Caño Benito. Renova oscari: MCNG 35926, 2; Venezuela: Amazonas: Isla Ratón. UFRJ 4606, 3 (c&s): aquarium material. Rivulus amphoreus: UFRJ 3889, 2, UFRJ 4606, 3 (c&s); Surinam: Tafelberg. R. apiamici: MZUSP 39976, H; MZUSP 39977, 3P (c&s); UFRJ 5971, 4; UFRJ 5972, 3 (c&s); Brazil: Mato Grosso do Sul; Bataguaçu. R. bahianus: UFRJ 3167, 44; UFRJ 4602, 2 (c&); UFRJ 277, 1 (c&s); Brazil: Bahia: Busca-Vida. R. brunneus: MZUSP uncatalogued, 10 (2 c&s); Panama: Isla de Barro Colorado. R. chucunaque: USNM , 68 (1c&s); Panama. R. cladophorus: UFRJ 643, 4P (2c&s); Guyana: Fourgassier. R. cryptocallus: UFRJ 359, 3; UFRJ 2126, 1 (c&s); Martinique: Ravine Vilaine. R. cyanopterus: UFRJ 5911, H; UFRJ 5913, 36P; UFRJ 5914, 5P (c&s); Brazil: Mato Grosso: Jaciara. R. cylindraceus: MHNC uncat., 6; Cuba: Zapata. USNM uncat., 1 (c&s); Cuba: La Habana. R. dapazi: UFRJ 5915, H; UFRJ 5916, 3P; UFRJ 5917, 2P (c&s); Brazil: Mato Grosso do Sul: Sonora. R. decoratus: MZUSP 39982, H; MZUSP 39983, 4P; UFRJ 2135, 3 (c&s); Brazil: Bahia: Ibiraba. R. derhami: UFRJ 392, 2 (c&s); Peru: Tingo Maria. R. depressus: UFPB 2213, H; UFPB 1749, 17 (2 c&s); UFRJ 2118, 1 (c&s); Brazil: Bahia: Porto Seguro. R. elongatus: MUSM 336, 6; MZUSP 26211, 3; Peru: Ucayali: Pucallpa. R. egens: UFRJ 5973, H; UFRJ 5974, 17P; UFRJ 5975, 3P (c&s); Brazil: Mato Grosso do Sul: Camapuã. R. erberi: UFRJ 358, 3; Ecuador: Napo: Coca. R. fuscolineatus: USNM , 10P; Costa Rica: Guanacaste: Tilaran. R. geayi: MZUSP uncatalogued Brazil: Amapa: Serra do Navio. R. haraldsiolii: UFRJ 125, 6; UFRJ 6295, 2 (c&s); Brazil: Santa Catarina: Joinville. R. hartii: MCZ 26092, 3; Trinidad: Port of Spain. MZUSP 37204, 3; MZUSP 38472, 2 (c&s); Venezuela: Nueva Esparta: Isla de Margarita. R. hildebrandi: USNM 92958, 1P; Panama: Chiriqui: Boquete. R. holmiae: USNM 66302, 1P; Guiana: Holmia. R. igneus: UFRJ 3888, 6; UFRJ 4595, 2 (c&s); Suriname: Creek Colibri. R. immaculatus: USNM , 2P; Venezuela: Bolivar: Elena de Uairen. R. janeiroensis: UFRJ 5333, 8; UFRJ 130, 2 P (c&s); UFRJ 5416, 7 (c&s); Brazil: Rio de Janeiro: Magé. R. luelingi: UFRJ 161, 8, UFRJ 127, 5 (c&s); Brazil: Santa Catarina: Araquari. R. lungi: ZSM 27825, 15; Brazil: Amapá: rio Flechal. R. nicoi: MCNG 23891, H; MCNG 23892, 1P; MCNG 23893, 1P; Venezuela: Amazonas: río Ventuari floodplains. R. nudiventris: MZUSP 40283, H; MZUSP 40284, 3P; MNRJ 11740, 2P (c&s); Brazil: Espírito Santo: Itapemirim. R. paracatuensis: MCP 29639, H; MCP 29640, 1P; UFRJ 2290, 3P (c&s); Brazil: Minas Gerais: Brasilândia de Minas. R. parnaibensis: MCP 29639, H; UFRJ 4962, 25P; UFRJ 5449, 4P (c&s); Brazil: Piauí: São Dimas. R. pictus: MNRJ 11550, H; MNRJ 11551, 2P; UFRJ 5959, 2 ex. (c&s); Brazil: Distrito Federal: Planaltina. R. pinima: MZUSP 39978, H; MZUSP 39984, 2P; UFRJ 5960, 20; UFRJ 5961, 4 (c&s); Brazil: Goiás: Rio Verde. R. punctatus: UFRJ 975, 9; UFRJ 2110, 4 (c&s); Brazil: Mato Grosso do Sul: Aquidauana. R. rossoi: UFRJ 5976, H; UFRJ 5977, 7P; UFRJ, 5978, 4P (c&s); Brazil: Mato Grosso do Sul: Campo Grande. R. rubrolineatus: MZUSP 26371, 3; MUSM 1465, 6; Peru: Loreto: Jenaro Herrera. R. rutilicaudus: UFRJ 5965, H; UFRJ 5966, 12P; UFRJ 5967, 3P (c&s); Brazil: Goiás: Serranópolis. R. santensis: UFRJ 5441, 11; UFRJ 6294, 5 (c&s); Brazil: São Paulo: Boracéia. R. simplicis: UFRJ 5940, H; UFRJ 5976, 3P; UFRJ 5942, 5P (c&s); Brazil: Rio de Janeiro: Parati. R. scalaris: UFRJ 5968, H; UFRJ 5969, 19P; UFRJ 5970, 4P (c&s); Brazil: Mato Grosso do Sul: Costa Rica. R. stagnatus: UFRJ 3890, 3; UFRJ 4605, 4 (c&s); Suriname: Wageningen. R. tenuis: UFRJ 4600, 4; UFRJ 4601, 2 (c&s); Guatemala: Alta Verapaz: Sebol. R. vittatus: MZUSP 39981, H; MZUSP 39981, 6P; UFRJ 2206, 15; UFRJ 5962, 4 (c&s); Brazil: Goiás: Cachoeira Alta. R. xiphidius: UFRJ 4608, 3 (c&s); Guyana (specimens born in aquarium). Trigonectes rubromarginatus: UFRJ 3553, 13; UFRJ 3554, 3 (c&s); Brazil: Tocantins: Barreira do Piqui. Appendix 2 Characters (in brackets) and character states (in parentheses) used to erect the phylogenetic hypothesis among species of Rivulus and other rivulids are listed below, with the respective reference to papers where the character is first described or discussed. Distribution of character states among terminal taxa is presented in the data matrix in Appendix 3. Superficial dermal bones and neurocranium [1] Lachrymal (Parenti 1981; Costa 1998a, 1998b) (CI: 1.00; RI: 1.00): (0) flat, posterior rim wide; (1) slightly twisted, posterior rim reduced, bone formed mainly by canal; (2) very twisted and narrow, slender, canal vestigial. [2] Ventral portion of lachrymal (Costa 1998a) (CI: 0.50; RI: 0.66): (0) short; (1) expanded. [3] Dermosphenotic (Costa 2004d): (CI: 1.00; RI: 1.00): (0) present; (1) absent. 171 aqua vol. 11 no

14 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin [4] Vomerine teeth (Costa 1998a) (CI: 0.40; RI: 0.80): (0) usually 1-4, sometimes 5-6; (1) 6-12; (2) teeth absent [not ordered]. [5] Anterior retrorse process of lateral ethmoid (Costa 1990) (CI: 1.00; RI: 1.00): (0) short; (1) moderate to elongate. [6] Lateral process of sphenotic (Costa 2005b) (CI: 0.50; RI: 0.50): (0) narrow; (1) wide. [7] Anterolateral process of parasphenoid (Costa 1998a) (CI: 0.16; RI: 0.78): (0) short, free; (1) long, attached to pterosphenoid. [8] Posterior portion of parasphenoid (Costa 2005b) (CI: 0.50; RI: 0): (0) wide; (1) narrow. [9] Lateral border of frontal (Costa 1998a) (CI: 1.00; RI: 1.00): (0) well-ossified, approximately straight; (1) poorly ossified, concave. Jaws, jaw suspensorium and opercular apparatus [10] General shape of the premaxilla and dentary (Costa 1998a) (CI: 0.25; RI: 0): (0) elongate, snout profile sharply pointed; (1) short, snout profile blunt. [11] General shape of ascending process of premaxilla (modified from Parenti, 1981, Costa 1998b) (CI: 1.00; RI: 1.00): (0) curved, posterior portion medially directed; (1) approximately straight. [12] Ascending process of premaxilla (Costa 2005b) (CI: 0.50; RI: 0): (0) wide; (1) narrow. [13] Ventral process of maxilla (Parenti 1981, Costa 1998a) (CI: 1.00; RI: 1.00): (0) slightly curved, anterior margin rounded; (1) bent, anterior margin triangular. [14] Maxilla (modified from Parenti 1981) (CI: 0.40; RI: 0.72): (0) approximately straight; (1) slightly twisted; (2) greatly twisted. [15] Rostral cartilage (Costa 1998a) (CI: 0.25; RI: 0.40): (0) approximately rounded; state 1: longitudinal length longer than transversal length. [16] Posterior portion of rostral cartilage (Costa 2004a) (CI: 0.20; RI: 0.69): (0) not distinctively narrowed; (1) distinctively narrowed. [17] Coronoid process of dentary (Costa 1998a) (CI: 1.00; RI: 1.00): (0) broad; (1) narrow. [18] Ventral process of angulo-articular (modified from Costa 1990, 1998a) (CI: 0.66, RI: 0.81): (0) large and broad; (1) large, somewhat narrowed; (2) moderate, narrow and pointed; (3) vestigial; (4) long and somewhat narrowed [not ordered]. [19] Curvature of ventral process of angulo-articular (Costa 1998a) (CI: 1.00, RI: 1.00): (0) straight; (1) curved. [20] External medial teeth of premaxilla and dentary (Costa 1998a) (CI: 0.33; RI: 0.50): (0) approximately directed as other teeth; (1) laterally displaced, strongly contrasting to other teeth. [21] Ventral portion of palatine (Costa 1998a) (CI: 1.00; RI: 1.00): (0) long, overlapping dorsal portion of quadrate; state 1: short, not or slightly contacting quadrate. [22] Metapterygoid (modified from Costa 1998a) (CI: 0.66; RI: 0.90): (0) about rectangular, dorsal and ventral portions wide and approximately equal in width; (1) dorsal portion slightly constricted; (2) about triangular, dorsal portion strongly constricted. [23] Posterior process of quadrate (Costa 1998a) (CI: 0.50; RI: 0.50): (0) short, about 50% of quadrate length; (1) long, about 70% of quadrate length. [24] Preopercle (Costa 1990) (CI: 1.00; RI: 1.00): (0) robust, L-shaped, with a well-developed anteromedian rim; (1) thin, C- shaped, with a reduced anteromedian rim. [25] Dorsal arm of preopercle (Costa 1990) (CI: 0.50; RI: 0.85): (0) broad; (1) narrow and pointed. Hyoid and branchial arches [26] Anterior process of urohyal (Costa 1998a) (CI: 1.00; RI: 1.00): (0) short; (1) elongate. [27] Dorsal process of urohyal length (CI: 1.00; RI: 1.00): (0) present; (1) absent; (?) very short to absent. [28] Dorsal process of urohyal length (Costa 1998a) (CI: 0.25; RI: 0.85): (0) short; (1) elongate; (?) process absent. [29] Basihyal (Costa 1998a) (CI: 0.50; RI: 0): (0) shorter than space occupied by basibranchials; (1) longer than space occupied by basibranchials. [30] Basihyal cartilage extent, as percentage of basihyal length (modified from Costa 1998a) (CI: 0.40; RI: 0.62): (0) %; (1) %; (2) %. [31] Interhyal (Parenti 1981) (CI: 1.00; RI: 1.00): (0) ossified; (1) cartilaginous. [32] Interhyal (Costa 2005b) (CI: 1.00; RI: 1.00): (0) large; (1) minute. [33] Number of branchiostegal rays (Costa 2004e) (CI: 0.33; RI: 0): (0) 6; (1) 5. [34] Main axis of first epibranchial (Costa 1998a) (CI: 1.00; RI: 1.00): (0) approximately straight; (1) curved. [35] Subdistal process of second epibranchial (Costa 2004a) (CI: 1.00; RI: 1.00): (0) present; (1) absent. [36] Uncinate process of third epibranchial (modified from Costa 1998a, b) (CI: 1.00; RI: 1.00): (0) long; (1) moderate; (2) short. [37] Interarcual cartilage (modified from Parenti 1981) (CI: 1.00; RI: 1.00): (0) not reduced; (1) reduced (CI: 1.00; RI: 1.00). [38] Number and arrangement of second pharyngobranchial teeth (Costa 2004a) (CI: 0.50; RI: 0.88): (0) numerous teeth arranged in two rows; (1) few teeth arranged in single row; (2) teeth absent [not ordered]. [39] Proximal edge of first hypobranchial (Costa 1998a) (CI: 0.33; RI: 0.33): (0) plain, terminating in single cartilage united to second basibranchial; (1) bifid, terminating in cartilage united to second basibranchial and another smaller cartilage united to first basibranchial. [40] Distal edge of first hypobranchial (Costa 2004a) (CI: 1.00; RI: 1.00): (0) articular face restricted to cartilaginous head of first ceratobranchial; (1) articular face anteriorly expanded. [41] Orientation of anterior tip of fifth ceratobranchial (Costa 2005b) (CI: 0.50; RI: 0.50): (0) anterior; (1) anterolateral. Vertebrae and caudal skeleton [42] Pointed, anteriorly directed process on first vertebra (modified from Costa 1990) (CI: 1.00; RI: 1.00): (0) absent; (1) present. [43] Epipleural ribs (Parenti 1981, Costa 1998a) (CI: 0.50; RI: 0.66): (0) rod-like; (1) bifid. [44] Neural prezygapophyses (Costa 1990) (CI: 0.33; RI: 0.84): (0) short; (1) long. [45] Hypurals (modified from Costa 1998a) (CI: 0.42; RI: 0.33): (0) two dorsal plates and one ventral plate separated by aqua vol. 11 no

15 Wilson J. E. M. Costa gap; (1) two plates separated by wide gap; (2) two plates in close proximity, sometimes ankylosed; (3) single plate. [46] Proximal end of parahypural (Costa 1998b) (CI: 1.00; RI: 1.00): (0) robust with paired dorsal processes overlapping preural centrum; (1) shortened and laminar, without dorsal paired process, not contacting preural centrum. [47] Hemal spine of preural centrum two (Costa 1998b) (CI: 0.50; RI: 0): (0) distinctively wider than hemal spines anterior to it; (1) slightly wider or equal in width to hemal spines anterior to it. [48] Number of vertebrae (modified from Costa 1990) (CI: 0.18; RI: 0.52): (0) 29-32; (1): 33-35; (2) [49] Number of caudal-fin rays (modified from Costa 1990) (CI: 0.18; RI: 0.59): (0) 26-31; (1) 32-36; (2) [not ordered]. Dorsal- and anal-fin skeleton [50] First dorsal-fin ray (modified from Parenti, 1981) (CI: 1.00; RI: 1.00): (0) single long first ray connected to two proximal radials; (1) long fin ray connected to two proximal radials, preceded by one or two short fin rays. [51] Anterior proximal radials of dorsal and anal fins (Costa 2005b) (CI: 1.00; RI: 1.00): (0) slender; (1): wide. [52] Number of anal-fin rays (modified from Costa 1998a) (CI: 0.28; RI: 0.44): (0) 9-15; (1) 16-20; (2) [53] Orientation of anterior proximal radials of anal fin (modified from Costa 1998a) (CI: 1.00; RI: 1.00): (0) anteriorly or dorsally directed; (1) posteriorly directed. Shoulder and pelvic girdle [54] Pectoral-fin insertion (Costa 1998b) (CI: 1.00; RI: 1.00): (0) lateral; (1) ventrolateral. [55] Supracleithrum and posttemporal (modified from Parenti 1981, Costa 1998b) (CI: 0.66; RI: 0.50): (0): separated; (1) co-ossified, limits almost inconspicuous; (2) fused to form a single structure. [56] Keel along supracleithrum-posttemporal (Costa 2005b) (CI: 1.00; RI: 1.00): (0) absent; (1) present. [57] Ventral process of posttemporal (Parenti 1981, Costa 1998a, b) (CI: 0.20; RI: 0.81): (0) present; (1) absent. [58] Posterior flange of cleithrum (Costa 1998a) (CI: 1.00; RI: 1.00): (0) present; (1) absent. [59] First postcleithrum (Parenti 1981) (CI: 1.00; RI: 1.00): (0) present; (1) absent. [60] Fourth pectoral radial (Costa 1998a) (CI: 0.33; RI: 0.75): (0) not expanded; (1) ventrally expanded. [61] Ischial process of pelvic girdle (Costa 2005b) (CI: 1.00; RI: 1.00): (0) prominent; (1) vestigial. [62] Number of pectoral-fin rays (modified from Costa 1998a): (0) 13-15; (1) [63] Number of pelvic-fin rays (Costa 1990) (CI: 1.00; RI: 1.00): (0) six; (1) seven; (2) eight. External morphology of body and head [64] Mouth (Costa 1998b) (CI: 1.00; RI: 1.00): (0) terminal; (1) superior. [65] Orbital rim (modified from Parenti 1981) (CI: 1.00; RI: 1.00): (0) free dorsally, attached ventrally; (1) completely attached. [66] Branchiostegal and opercular membranes (Parenti 1981) (CI: 1.00; RI: 1.00): (0) separated by long fold; (1) continuous. [67] Skin fold on corner of preopercular region (Costa 2005b) (CI: 1.00; RI: 1.00): (0) present; (1) absent. External morphology of fins [68] Pectoral fin (Costa 1990) (CI: 0.50; RI: 0.75): (0) rounded; (1) pointed. [69] Pectoral-fin length (modified from Parenti 1981) (CI: 0.33; RI: 0.75): (0) % SL; (1) % SL. [70] Extent of pelvic fin in males (Costa 1998a) (CI: 0.25; RI: 0.25): (0) short, tip not surpassing anterior portion of anal fin; (1) long, its tip reaching the central or the posterior portion of the anal fin. [71] Pelvic-fin (Costa 2005b) (CI: 1.00; RI: 1.00): (0) bases separated or in contact; (1) bases united; (2) pelvic fins united along proximal portion of medial margin. [72] Dorsal fin (modified from Costa 1998a) (CI: 0.33; RI: 0.84): (0) short, tip rounded; (1) somewhat elongated, tip pointed; (2) long, tip sharply pointed. [73] Anal fin (modified from Costa 1998a) (CI: 0.22; RI: 0.75): (0) short, tip rounded; (1) somewhat elongated, tip pointed; (2) long, tip sharply pointed. [74] Filaments on tip of dorsal and anal fins in males (Costa 1998a) (CI: 0.50; RI: 0.66): (0) absent; (1) present. [75] Filaments on posterior border of caudal fin in males (Costa 1998a) (CI: 0.33; RI: 0.60): (0) absent; (1) present. [76] Caudal-fin length in males (Costa 1990) (CI: 0.40; RI: 0.70): (0) ; (1) ; (2) [77] Caudal-fin shape in males (CI: 0.33; RI: 0.40): (0) rounded; (1) subtruncate; (2) truncate; (3) acuminate [not ordered]. [78] Dorsal and ventral extensions on caudal fin in males (Costa 1998a) (CI: 0.50; RI: 0): (0) absent; (1) present. Squamation [79] General arrangement of frontal scales (Hoedeman 1958) (CI: 0.50; RI: 0.85): (0) transverse; (1) circular; (2) irregular [not ordered]. [80] Predominant frontal squamation-pattern (Hoedeman 1958) (CI: 0.28; RI: 0.52): (0) G; (1) E; (2) D; (3) F; (4) S [not ordered]. [81] Arrangement of E-scales (Costa 1990) (CI: 1.00; RI: 1.00): (0) overlapped; (1) not overlapped. [82] Caudal-fin squamation in older males (Costa 1990, 1998a) (CI: 0.25; RI: 0.57): (0) approximately on anterior % of fin; (1) approximately on 40 % of fin; (2) approximately on % of fin. [83] Anal-fin base squamation in males (Costa 2005b) (CI: 1.00; RI: 1.00): (0) no scales on anal-fin base; (1) 1-7 rows of scales on anal-fin base. Laterosensory system [84] Cephalic canals (modified from Parenti 1981) (CI: 0.75; RI: 0.92): (0) anterior and posterior infraorbital, preopercular and posterior mandibular closed, supraorbital, rostral, anterior mandibular open with skin trenches around neuromasts; (1) anterior and posterior infraorbital, and preopercular closed, supraorbital, rostral, anterior and posterior mandibular open with skin trenches around neuromasts; (2) anterior and posterior infraorbital, dorsal preopercular, supraorbital, rostral, anterior and posterior mandibular open with skin trenches around neuromasts; (3) no vestige of canals, all neuromasts completely exposed. 173 aqua vol. 11 no

16 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin [85] Number of anterior supraorbital neuromasts (Costa 1990) (CI: 0.50; RI: 0.33): (0) 3; (1) 4; (2) 6 7. [86] Arrangement of anterior supraorbital neuromasts (Costa 2004d) (CI: 1.00; RI: 1.00): (0) continuous row; (1) posteriormost neuromast separated by space covered by scale. [87] Arrangement of anterior and posterior rostral neuromast (CI: 1.00; RI: 1.00): (0) longitudinal; (1) transverse. Contact organs [88] Contact on flank scales in males (Costa 2005b) (CI: 0.14; RI: 0.66): (0) absent; (1) present. Male colour patterns [89] Red marks on male flank (CI: 0.16; RI: 0.44): (0) absent; (1) present. [90] Kind of red marks on male flank (CI: 0.44; RI: 0.69): (0) vertical rows of dots, sometimes forming stripes; (1) vertical rows of dots on anterior portion, chevron-like bars on posterior; (2) dots on chevron-like series on entire flank, sometimes forming oblique bars; (3) overlapping longitudinal and oblique rows; (4) red marks irregularly arranged, not forming any distinguishable pattern; (?) red marks absent [not ordered]. [91] Dark pigmentation pattern on humeral region (modified from Costa 1998a) (CI: 0.33; RI: 0.50): (0) not distinctively coloured; (1) forming distinct blotch. [92] Dark pigmentation pattern on upper portion of caudalfin base (Costa 2005b) (CI: 0.50; RI: 0.40): (0) not distinctively coloured; (1) forming distinct blotch. [93] Dark pigmentation on anterior half of flank when fish is exposed to sunlight (modified from Costa 1998a) (CI: 0.20; RI: 0.80): (0) not distinctively marked; (1) forming grey vertical bars; (2) forming dark grey broad stripe; (3) forming dark grey oblique bars [not ordered]. [94] Dark pigmentation between dorsum and flank (modified from Costa, 1998a) (CI: 1.00; RI: 1.00): (0) not distinctively marked; (1) forming longitudinal dark zone. [95] Dark pigmentation pattern on iris (Parenti 1981) (CI: 1.00; RI: 1.00): (0) no distinctive mark; (1) bar crossing eye. [96] Pigmentation pattern on suborbital region (Costa 1998a) (CI: 0.66; RI: 0.75): (0) no distinctive mark; (1) grey to black suborbital bar; (2) red suborbital spot [not ordered]. [97] Pigmentation pattern on post-orbital and preopercular region (Costa 1998a) (CI: 0.40; RI: 0.66): (0) no distinctive pattern; (1) two oblique bars; (2) post-orbital vertical bar; (3) two oblique stripes [not ordered]. [98] Dark pigmentation pattern on lower jaw (Costa 2004d) (CI: 0.33; RI: 0.84): (0) not distinctively concentrated; (1) jaw dark grey to black; (2) transverse stripe through chin [not ordered]. [99] Dark blue to dark purplish blue iridescence on opercular and infraorbital region (Costa 1998a) (CI: 0.25; RI: 0.57): (0) absent; (1) present. [100] Dark pigmentation on distal portion of male anal fin (Costa 1998a) (CI: 0.20; RI: 0.50): (0) not distinctive concentrated; (1) concentrated to form stripe. [101] Yellow to orange pigmentation pattern on ventral portion of caudal fin (modified from Costa 1998a) (CI: 0.10; RI: 0.55). (0) not distinctive concentrated; (1) concentrated to form stripe; (?) variable. [102] Yellow to orange pigmentation pattern on dorsal portion of caudal fin (modified from Costa 1998a) (CI: 0.11; RI: 0.50). (0) not distinctive concentrated; (1) concentrated to form stripe; (?) variable. [103] Dark pigmentation of ventral margin of male caudal fin (modified from Costa 1998a) (CI: 0.33; RI: 0.75): (0) not distinctive concentrated; (1) concentrated to form stripe. [104] Red pigmentation pattern on subventral portion of caudal fin (CI: 0.33; RI: 0.66): (0) not distinctive concentrated; (1) concentrated to form stripe. [105] Red pigmentation on caudal fin (modified from Costa 1998a) (CI: 0.33; RI: 0.66): (0) not distinctive arranged; (1) forming bars. [106] Dark pigmentation on dorsal portion of caudal fin (Costa 1998a) (CI: 0.50; RI: 0.75): (0) not distinctive concentrated; (1) concentrated to form dark brown stripe; (?) variable. [107] Melanophore pattern on pectoral fin (Costa 1998a) (CI: 0.33; RI: 0): (0) no distinctive marks; (1) spots; (2) bars [unordered]. [108] Colour of pectoral fin (CI: 0.33; RI: 0.50): (0) hyaline; (1) yellow. Female colour patterns [109] Dark pigmentation pattern on flank and fins (Costa 1998a) (CI: 0.50; RI: 0): (0) forming bars, stripes or spots, according to the pattern occurring in males; (1) flank almost plain, pigmentation reduced to minute dots, not presenting the general pattern as in males. [110] Red marks on flank (CI: 0.25; RI: 0.84): (0) absent; (1) present. [111] Melanophore pattern on unpaired fin margins (Costa 1995a) (CI: 1.00; RI: 1.00): (0) not distinctively concentrated; (1) forming black zones of marginal regions. [112] Dark pigmentation pattern on upper portion of caudalfin base (Costa 1990) (CI: 0.14; RI: 0.71): (0) no distinctive mark; (1) one black spot. [113] Kind of caudal blotch (CI: 0.66; RI: 0.85): (0) black blotch close to fin margin, surrounded by light area to form an ocellate spot; (1) black blotch close to or contacting fin margin, anteriorly bordered by triangular light spot; (2) black blotch apart from fin margin, with light area surrounding anterior, dorsal and posterior margin; (?) caudal blotch absent. Appendix 3 Matrix of 113 characters for 49 aplocheiloid species. Characters and states are according to Appendix 2. Autapomorphies were excluded from the analysis; 0 = plesiomorphic state; 1-4 = apomorphic states;? = not pertinent or unknown state. aqua vol. 11 no

17 Wilson J. E. M. Costa A. panchax ? ? S. guignardi ? K. brasiliensis ? ? ? P. auriferus ? ? ? A. boticarioi ? G. hoignei ? ? ? M. xiphophora ? M. apurinan 200? ? N. paraguayensis ? P. compacta ? ? P. longipinnis ?? ? R. maculipinnis ? ? ? R. stellifer ? R. oscari R. amanapira R. amphoreus R. apiamici R. atratus ? ? R. bahianus R. beniensis R. brunneus R. cladophorus ? R. cryptocallus R. cylindraceus R. depressus R. dibaphus ? ? R. hartii R. igneus ? R. janeiroensis ? R. kirovskyi ? R. litteratus R. luelingi R. micropus R. modestus ? R. ornatus ? R. pictus R. punctatus R. romeri ? R. santensis ? ? R. strigatus ? ? R. taeniatus R. tecminae R. tenuis ? R. uakti 201? R. uatuman ? R. urophthalmus R. violaceus R. zygonectes T. rubromarginatus ? 175 aqua vol. 11 no

18 aqua vol. 11 no

19 1. Manuscript preparation: manuscripts must be submitted in English. In exceptional cases aqua may provide translations of manuscripts written in French, German, Italian, or Spanish. Manuscripts must be word-processed in Microsoft WORD and submitted in an electronic form. Generic, specific, and sub-specific names must be italicised. All papers must conform to the International Code of Zoological Nomenclature. Authors are strongly advised to follow the format set out in previous publications of aqua. 2. Title: the title must be short and should precisely identify the main topic of the article. Names of genera or species are followed by the systematic group to which they belong. Author name(s) are given in full beneath the title, followed by the complete mailing and address(es). 3. Abstract: the abstract should not exceed 250 words and draw attention to the principal conclusions. It should not contain any uncommon abbreviations or literature citations. The inclusion of abstracts in other languages is optional. 5. Subject matter: the text of the manuscript is usually arranged in four main sections: Introduction, Materials and methods (including a key to abbreviations), Results, and Discussion. Other subdivisions may be chosen depending on the material presented. Acknowledgements should be placed between the text and references. Scientific names of genera, species, and subspecies should be followed by the name(s) of author(s) and the year of publication on first mention. A description of a new taxon must contain the following sections: Material, Diagnosis, Description, and Affinities. Holotype and paratypes must be clearly identified, the institution in which they have been deposited named, and the catalogue numbers given. Private collections are not acceptable as repositories for holotypes. Synonyms must be arranged in chronological order. Identification keys must be dichotomous. The metric system and SI units must be used. Temperatures are given in C. Fractions should not be used. 6. References to literature: the name-year system must be used. The list of references should be placed at the end of the paper, alphabetically arranged according to author name. Only those publications cited in the paper may be included. Titles of journals must be given in full. Examples of correct reference formats: Guidelines for Authors BLABER, S. J. M Fish of the Trnity inlet system of North Queensland, with notes on the ecology of fish faunas of tropical Indo-Pacific estuaries. Australian Journal of Marine and Freshwater Research 31: DAY, J. H., BLABER, S. J. M., & WALLACE, J. H Estuarine fishes. In: Estuarine Ecology with Particular Reference to Southern Africa. (Ed. J.H. Day.): A. A. Balkema, Rotterdam. DIMMICH, W. W Ultrastructure of North American cyprinid maxillary barbels. Copeia 1988 (1): TREWAVAS, E Tilapiine Fishes of the Genera Sarotherodon, Oreochromis and Danakilia. British Museum (Natural History), London, 583 pp. 7. Submission of manuscript and illustrations: The manuscript and illustrations must be submitted digitally to the Scientific Editor: Dr. Friedhelm Krupp Senckenberg Research Institute Senckenberganlage Frankfurt am Main, Germany f.krupp@senckenberg.de Tel: , Fax: to whom all subsequent correspondence shall be addressed. Texts, tables, and graphs should be in Microsoft-compatible electronic form. After the paper has been accepted for publication, illustrations as high-resolution TIF files or original photographs (ideally transparencies; otherwise glossy prints, preferably in the size in which they will appear - the type area of aqua is 158 x 224 mm, one column is 76 mm wide) must be sent to: Aquapress, The Managing Editor Via G. Falcone 11, Miradolo Terme (Pavia), Italy aqua@aquapress-bleher.it Authors should retain copies of all materials for reference. Proofs of accepted papers will be sent as PDF files by e- mail attachment to the corresponding author. 8. Evaluation of manuscripts: manuscripts will be evaluated by the editors and referees. Papers are accepted on the understanding that they have not and will not be published elsewhere. 9. Reprints: Authors will receive one free copy of the issue in which their paper appears and an e-print in PDF format. Additional copies may be ordered from Aquapress.

20 aqua Journal of Ichthyology and Aquatic Biology Vol. 11 (4), October 2006 Contents: Wilson J. E. M. Costa: Relationships and taxonomy of the killifish genus Rivulus (Cyprinodontiformes: Aplocheiloidei: Rivulidae) from the Brazilian Amazonas river basin, with notes on historical ecology Papers appearing in this journal are indexed in: Zoological Record; Biolis Biologische Literatur Information Senckenberg; Cover photo: Rivulus urophthalmus, UFRJ 6264, male, 29.5 mm SL (one day after collection); Brazil: Pará: Altamira. Photo by W. J. E. M. Costa. Brazil: Pará: Primavera; pool close to creek near forest border, typical habitat of Rivulus urophthalmus. Photo by W. J. E. M. Costa. See p