Wilson J. E. M.

Costa

Material examined
Brazil: Estado do Amazonas: UFRJ 5925, holo-
type; 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. 2003.
UFRJ 5928, 9; São Gabriel da Cachoeira, upper rio
Negro drainage, rio Amazonas basin, stream tribu-
tary to igarapé Miuá, km 13.6 of the road São
Gabriel da Cachoeira to Cucuí, 0°2’58.7”S
66°57’48.6”W; W. J. E. M. Costa, S. Lima and L.
Silva, 30 Aug. 2003.

Diagnosis Fig. 39. Brazil: Amazonas: São Gabriel da Cachoeira; creek

Distinguished from all other congeners by the at the forest border, the habitat of Rivulus uakti. Photo by
combination of the following features: anterior por- W. J. E. M. Costa.
tion of trunk slightly deeper than wide, jaws short,
snout blunt, tip of anal fin slightly pointed in male, Also differs from all other congeners of the Owiyeye
caudal fin rounded in male, pelvic-fin tip reaching in possessing two unique colour patterns: dorsal fin
middle of anal-fin base in male, dorsal-fin origin on with bright blue distal zone in male, and caudal spot
vertical between base of 8th and 9th anal-fin rays, forming a black bar posteriorly bordered by pale yel-
dorsal-fin rays 7-8, anal-fin rays 11-12, frontal squa- low bar in female (Figs 37-38).
mation S-patterned, frontal scales arranged trans-
versely, canal preopercular absent, contact organs on Distribution
flank scales in male, longitudinal series of scales 32- Upper rio Negro basin (Fig. 22).
33, gill rakers of first branchial arch 1 + 7-8, oblique
rows of red dots on flank in male, transverse dark Habitat
brown bar through the chin, and dark grey to black Moderately sunny places of shallow streams within
spot on dorsal portion of caudal-fin in both sexes. 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, 0°9’19.2”S 66°59’58.9”W,
Fig. 37. Rivulus uakti, UFRJ 5925, male, holotype, 24.0 altitude 110 m, Estado do Amazonas, Brazil; holo-
mm SL (two days after collection); Brazil: Amazonas: São type: UFRJ 5929).
Gabriel da Cachoeira. Photo by W. J. E. M. Costa.
Material examined
Brazil: Estado do Amazonas: UFRJ 5929, holo-
type; 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. 2003.
Fig. 38. Rivulus uakti, UFRJ 5926, female, paratype, 27.9
mm SL (two days after collection); Brazil: Amazonas: São Diagnosis
Gabriel da Cachoeira. Photo by W. J. E. M. Costa. Distinguished from all other congeners by the

159 aqua vol. 11 no. 4 - 2006

 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin

combination of the following features: anterior por- transversely, canal preopercular absent, contact
tion of trunk slightly deeper than wide, jaws short, organs on flank scales in male, longitudinal series of
snout blunt, tip of anal fin slightly pointed in male, scales 48-49, gill rakers of first branchial arch 1 + 8,
caudal fin truncate in male, pelvic-fin tip reaching longitudinal rows of red dots on flank in male, trans-
anterior portion of anal-fin base in male, dorsal-fin verse dark brown bar through the chin, and ovoid
origin on vertical between base of 11th and 13th anal- dark grey to black spot on dorsal portion of caudal
fin rays, dorsal-fin rays 8, anal-fin rays 14-15, frontal fin in both sexes. Additionally distinguished from all
squamation S-patterned, frontal scales arranged 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).

Habitat notes
Small shallow temporary pools in the forest,
about 0.5-2.0 m in diameter and about 15 cm deep
Fig. 40. Rivulus amanapira, UFRJ 5929, male, holotype, (Fig. 41).
38.6 mm SL (one day after collection); Brazil: Amazonas:
São Gabriel da Cachoeira. Photo by W. J. E. M. Costa. 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
67°20’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 conflu-
ence with río Sipapo, Orinoco basin; L. Nico & E.
Guayamare, 28 May 1989. Brazil: Estado do Ama-
zonas: UFRJ 2085, 2; UFRJ 2095, 1 (c&s);
Inambu, right bank of upper rio Negro; D. Teixeira,
26 Aug. 1991.

Diagnosis
Distinguished from all other congeners by the
combination of the following features: anterior por-
tion 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 9th and 10th anal-fin rays, dorsal-
fin rays 8-10, anal-fin rays 12-15, frontal squama-
tion S-patterned, frontal scales arranged transversely,
Fig. 41. Brazil: Amazonas: São Gabriel da Cachoeira; shal- canal preopercular absent, contact organs on flank
low temporary pool within the forest, habitat of Rivulus scales in male, longitudinal series of scales 37-41, gill
amanapira. Photo by W. J. E. M. Costa. rakers of first branchial arch 1 + 7, red stripes on

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 Wilson J. E. M. Costa

Fig. 42. Jaw suspensorium and opercular apparatus of Rivulus pictus, UFRJ 5959, female, 27.7 mm SL. AA = angulo-artic-
ular; 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 carti-
lage; 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 overlapping caudal fin bars (113.2) (Fig. 44). All
vertically elongated dark grey to black spot on dor- included species except R. modestus have oblique
sal portion of caudal-fin in both sexes. chevron-like rows of red dots or bars on flank in
males (90.2) (Fig. 43).
Distribution
Upper Orinoco river basin, Venezuela, and upper Etymology
Negro river, Brazil (Fig. 19). From the Latin, melania (black pigmentation on
the skin) and rivulus (stream), referring to the black
Habitat margins of unpaired and pelvic fins, a condition
Temporary pools in savannas (D. Teixeira, pers. unique among congeners. Gender masculine.
comm.).
Included species
Melanorivulus, new subgenus Rivulus apiamici Costa, 1989, R. cyanopterus Costa,
Type species: Rivulus punctatus Boulenger, 1895. 2005, R. dapazi Costa, 2005, R. decoratus Costa,
1989, R. egens Costa, 2005, R. litteratus Costa,
Diagnosis 2005, R. modestus Costa, 1991, R. paracatuensis
Distinguished from the remaining subgenera of Costa, 2003, R. parnaibensis Costa, 2003, R. pictus
Rivulus in having the following apomorphic fea- Costa, 1989, R. pinima Costa, 1989, R. punctatus
tures: dorsal portion of preopercle short and pointed Boulenger, 1895, R. rossoi Costa, 2005, R. rutili-
(25.1) (Fig. 42), dorsal and anal fins slightly pointed caudus Costa, 2005, R. scalaris Costa, 2005, R. vio-
in male (72.1, 73.1; also occurring in Laimosemion) laceus Costa, 1991, R. vittatus Costa, 1989, and R.
(Fig. 43), two oblique bars on post-orbital region zygonectes Myers, 1927.
(97.1) (Fig. 43), melanophores concentrated on
margins of unpaired and pelvic fins in female Distribution
(111.1) (Fig. 44), and female with black spot on Paraná-Paraguay-Uruguay, upper Tapajós, upper
upper portion of caudal fin not close to fin margin, and middle Xingu, upper and middle Tocantins,

161 aqua vol. 11 no. 4 - 2006

 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin

São Francisco and Parnaíba river basins, in central Material examined

and north-eastern Brazil, south-eastern Bolivia, Brazil: Tocantins-Araguaia basin: Estado de Goiás:
Paraguay and northern Argentina. Greatest diversi- CAS 76314, lectotype; Vereda Extrema into Cana
fication concentrated in the central Brazilian Brava; C. Ternetz, 14 Jan. 1924. UFRJ 1575, 3;
Plateau (Costa, 2005a). stream 15 km E of Aruanã; W. J. E. M. Costa et al.,
29 Aug. 1993. UFRJ 1748, 6; stream tributary to
Rivulus modestus Costa, 1991 rio Verde, 30 km E of São Miguel do Araguaia; W.
J. E. M. Costa et al., 25 Aug. 1993. UFRJ 1567, 5;
Rivulus modestus Costa, 1991: 329 (type locality: rio stream 98 km W of Jussara; W. J. E. M. Costa et al.,
Mutum, rio Tapajós basin, Mato Grosso, Brazil; 30 Aug. 1993. UFRJ 1482, 22; UFRJ 2108, 2
holotype: MNRJ 11670). (c&s); stream tributary to rio Verde, 32 km N of São
Miguel do Araguaia; W. J. E. M. Costa et al., 25
Material examined Aug. 1993. MZUSP 35416, 20; MZUSP 38334,
Brazil: Estado de Mato Grosso: MNRJ 11670, 55; pool in Aruanã; W. J. E. M. Costa et al., 28 Jan.
holotype; MNRJ 11671, 5 paratypes; UFRJ 2102, 1986. Estado do Tocantins: UFRJ 2106, 6; UFRJ
6; UFRJ 2103, 5 (c&s); small stream tributary of rio 2107, 4 (c&s); stream 20 km S of Santa Rosa; W. J.
Mutum, at crossing with road BR-364, 51 km from E. M. Costa et al., 15 Feb. 1994. MZUSP 38375, 2;
Vilhena, rio Juruena drainage, rio Tapajós basin; K. stream tributary to córrego Dona Francisquinha,
Tanizaki, M. T. Lacerda, S. O. Kullander & A. Porto Nacional; W. J. E. M. Costa et al., 31 Jan.
Hogerborn-Kullander, 16 Oct. 1989. 1986. UFRJ 1712, 7; stream tributary to rio Verde,
50 km N of São Miguel do Araguaia; W. J. E. M.
Diagnosis Costa et al., 25 Aug. 1993. UFRJ 1597, 2; stream 12
Distinguished from all other congeners by the km N of Sandolândia; W. J. E. M. Costa et al., 27
combination of the following features: anterior por- Aug. 1993. MZUSP 37212, 28 (6 c&s); road
tion of trunk deeper than wide, jaws short, snout between road BR-153 and Formoso do Araguaia, 29
blunt, tip of anal fin slightly pointed in male, caudal km S of Gurupi; P. S. Santos-Filho, Jun. 1976.
fin rounded in male, pelvic-fin tip reaching anterior MZUSP 45223, 9; rio Água Fria, road Araguaçu-
portion of anal-fin base in male, dorsal-fin origin on Barreira do Piqui, 27 km N of Araguaçu; F. C. T.
vertical between base of 8th and 9th anal-fin rays, Lima, 21-26 Feb. 1993. UFRJ 1374, 13; shallow
dorsal-fin rays 10-11, anal-fin rays 13-14, frontal lagoon 2 km W of rio das Mortes at the road Água
squamation F-patterned, frontal scales arranged cir- Boa-Cocalinhos; W. J. E. M. Costa et al., 20 Feb.
cularly, canal preopercular absent, contact organs 1993. UNT 555, 5; river at the road BR-153
absent, longitudinal series of scales 31-33, gill rakers between Filadelphia and Araguaina; E. L. Beerli, 2
of first branchial arch 1 + 7-8, red marks of flank Nov. 2003. UNT 556, 3; rio Brejão, Araguaina; E.
absent, lower jaw black, and round black spot on L. Beerli & L, M, Lima, 22 Nov. 2003. UNT 2045,
dorsal portion of caudal-fin in female. 1; córrego Gorgulho, Porto Nacional; NEAMB, 14
Sep. 2001. UNT 2046, 1; córrego Gorgulho, Porto
Distribution Nacional; NEAMB, 14 Sep. 2001. UNT 2047, 3;
Upper rio Juruena drainage (Fig. 10). córrego Marimbondo, Tupirantins; NEAMB, 27
Oct. 2000. UNT 2048, 1; córrego Barreiro, Guaraí;
Habitat notes NEAMB, 20 Oct. 2000. UNT 2049, 2; córrego
Shallow creeks with slow current, in moderately Gorgulho, Porto Nacional; NEAMB, 14 Sep. 2001.
sunny region. UNT 2050, 1; córrego Gorgulho, Porto Nacional;
NEAMB, 14 Sep. 2001. UNT 2051, 1; córrego
Rivulus zygonectes Myers, 1927 Lageado, Paranã; NEAMB, 22 Mar. 1999. UNT
2052, 9; córrego Água Suja, Tupirantins; NEAMB,
Rivulus zygonectes Myers, 1927: 127 (type locality: 27 Oct. 2000. UFRJ 5150, 1; floodplains of left
Vereda Extrema, into Cannabrava [now Cana bank of rio Tocantins, about 1200 m from the river
Brava, at rio Tocantins left bank], Goyas [now margin, SSE Sampaio, 5°23’00”S 47°51’41”W; G.
Estado de Goiás], Brazil; lectotype: CAS 76314, C. Brasil, 8 Jun. 2000. Estado do Maranhão: UFRJ
designated by Huber, 1992: 464). 5152, 16; swamp about 500 m from the right mar-
gin of rio Tocantins, 5°12’33”S 48°27’59”W; G. C.

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 Wilson J. E. M. Costa

Brasil, 8 Jun. 2000. Rio Xingu basin, Estado de Rivulus violaceus Costa, 1991
Mato Grosso: UFRJ 1386, 1; stream 67 km N of
Paranatinga; W. J. E. M. Costa et al., 10 Feb. 1993. Rivulus violaceus Costa, 1991: 331 (type locality: rio
UFRJ 1173, 5; stream at the road BR-080, 9 km E das Mortes, rio Araguaia-Tocantins basin, Mato
of São José do Xingu; W. J. E. M. Costa et al., 16 Grosso, Brazil; holotype: MNRJ 11672).
Feb. 1993. UFRJ 1417, 1; stream at the road BR-
080, 2 km E of São José do Xingu; W. J. E. M. Costa Material examined
et al., 18 Feb. 1993. UFRJ 1389, 6; stream at the Brazil: Estado de Mato Grosso, rio das Mortes
road BR-080, 41 km E of São José do Xingu; W. J. basin: MNRJ 11672, holotype; MNRJ 11673, 5
E. M. Costa et al., 17 Feb. 1993. UFRJ 1420, 6; paratypes; UFRJ 2104, 20; rio das Mortes when
stream at the road BR-080, 42 km E of São José do crossed by BR-163 at km 313; K. Tanizaki, M. T.
Xingu; W. J. E. M. Costa et al., 17 Feb. 1993. UFRJ Lacerda, S. O. Kullander & A. Hogerborn-Kullan-
1392, 7; stream at the road BR-080, 36 km E of São der, 19 Oct. 1989. UFRJ 143, 16; UFRJ 2105, 7
José do Xingu; W. J. E. M. Costa et al., 17 Feb. (c&s); small stream tributary to rio Perdidos, close
1993. UFRJ 1357, 20; stream at the road Para- to BR-070, about 10 km W of Primavera do Leste;
natinga-Canarana, 158 km of Paranatinga; W. J. E. K. Tanizaki, M. T. Lacerda, S. O. Kullander & A.
M. Costa et al., 12 Feb. 1993. UFRJ 1414, 20; Hogerborn-Kullander, 19 Oct. 1989. UFRJ 1207,
UFRJ 2109; stream at the road BR-080, 7 km E of 4; rio Suspiro, 57 km S of Paranatinga; W. J. E. M.
rio Xingu margin; W. J. E. M. Costa et al., 16 Feb. Costa, C. P. Bove, R. D. Cunha & C. Muratori, 9
1993. UFRJ 6265, 1; stream near rio Xingu, Feb. 1993. UFRJ 4284, 13; idem; M. Britto, C.
Altamira; W. J. E. M. Costa et al., 16 Jun. 2004. Rio Moreira & R. Cunha, 28 Oct. 1996. UFRJ 4283,
Tapajós basin, Estado de Mato Grosso: MZUSP 3; stream at the road Primavera do Leste-Para-
45304, 10; upper rio Preto, road Cuiabá-Santarém; natinga; M. Britto, C. Moreira & R. Cunha, 28
N. Menezes et al., 24 Oct. 1992. MZUSP 45321, 2; Oct. 1996.
stream tributary to rio Preto, road to São Francisco;
N. Menezes et al., 24 Oct. 1992. Diagnosis
Distinguished from all other congeners by the
Diagnosis combination of the following features: anterior por-
Distinguished from all other congeners by the tion of trunk slightly deeper than wide, jaws short,
combination of the following features: anterior por- snout blunt, tip of anal fin slightly pointed in male,
tion of trunk slightly deeper than wide, jaws short, caudal fin rounded in male, pelvic-fin tip reaching
snout blunt, tip of anal fin slightly pointed in male, urogenital papilla in male, dorsal-fin origin on verti-
caudal fin rounded in male, pelvic-fin tip reaching cal between base of 8th and 9th anal-fin rays, dorsal-
anterior portion of anal-fin base in male, dorsal-fin fin rays 10-11, anal-fin rays 14-15, frontal squama-
origin on vertical between base of 8th and 10th anal- tion F-patterned, frontal scales arranged circularly,
fin rays, dorsal-fin rays 9-11, anal-fin rays 13-15, canal preopercular absent, contact organs absent,
frontal squamation F-patterned, frontal scales longitudinal series of scales 30-32, gill rakers of first
arranged circularly, canal preopercular absent, con- branchial arch 1 + 7-8, oblique chevron-like red bars
tact organs absent, longitudinal series of scales 33- on flank in male, lower jaw black, and round black
35, gill rakers of first branchial arch 1 + 7-8, oblique spot on dorsal portion of caudal fin in female.
rows of red dots sometimes united forming
chevron-like bars on flank in male, lower jaw black, Distribution
and round black spot on dorsal portion of caudal Upper rio das Mortes drainage (Fig. 10).
fin in female.
Habitat notes
Distribution Shallow creeks with slow current, in gallery forest
Tocantins, Araguaia, Xingu and Tapajós river of savanna-like region.
basins (Fig. 22).
Rivulus litteratus Costa, 2005
Habitat notes
Shallow creeks with slow current, in gallery forest Rivulus sp.: Lacerda, 1989: 25 (Ponte Branca, Rio
of savanna-like region. Araguaia, central Brazil).

163 aqua vol. 11 no. 4 - 2006

 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin

Rivulus pictus (non Costa): Costa et al., 2003: 143 of first branchial arch 1 + 7-8, overlapped red marks
(misidentification of specimens from Alto Ara- of variable shape on flank in male, lower jaw dark
guaia). grey, and round black spot on dorsal portion of cau-
Rivulus litteratus Costa, 2005a: 75 (type locality: dal-fin in female. Also distinguished from the
Município de Alto Araguaia, Córrego do Sapo, remaining species of Melanorivulus by possessing a
upper Rio Araguaia basin, road MT-100, 31 km S colour pattern consisting of red marks variable in
of Alto Araguaia, 17º33’38.5”S 53º18’33.1”W, shape and highly overlapped (Fig. 43).
altitude 750 m, Estado de Mato Grosso, Brazil;
holotype: UFRJ 5956). Distribution
Upper rio Araguaia drainage (Fig. 23).
Material examined
Brazil: Estado de Mato Grosso: Município de Alto Habitat notes
Araguaia, upper rio Araguaia basin: UFRJ 5956, Shallow creeks with slow current, in gallery forest
holotype; Município de Alto Araguaia, córrego do of savanna-like region (Fig. 45).
Sapo, upper rio Araguaia basin, road MT-100, 31
km S of Alto Araguaia; W. J. E. M. Costa, B. B. Discussion
Costa and C. P. Bove, 14 Jan. 2004. The present study confirms a weakly supported
genus Rivulus containing clades corroborated both
Diagnosis by morphological (e.g. Huber 1992, Costa 1998a,
Distinguished from all other congeners by the the present study) and molecular data (e.g. Murphy
combination of the following features: anterior por- et al. 1999, rbek & Larson 1999), and recognized as
tion of trunk slightly deeper than wide, jaws short, subgenera. No unambiguous, apomorphic morpho-
snout blunt, tip of anal fin slightly pointed in male, logical condition was found to be shared by all
caudal fin rounded in male, pelvic-fin tip reaching species presently assigned to Rivulus and molecular
anterior portion of anal-fin base in male, dorsal-fin studies indicate that Rivulus is a paraphyletic assem-
origin on vertical between base of 7th 8th l-fin rays, blage (Murphy et al. 1999, Hrbek & Larson 1999).
dorsal-fin rays 9-10, anal-fin rays 13-14, frontal However, a group comprising all species of Rivulus
squamation F-patterned, frontal scales arranged cir- except the subgenus Rivulus is well supported by
cularly, canal preopercular absent, contact organs morphology, their members having a uniquely
absent, longitudinal series of scales 31-32, gill rakers derived pattern of arrangement of rostral neuro-
masts (i.e. transversely placed), and a long neural
prezygapophysis in caudal vertebrae as already dis-
cussed 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, Fig. 45. Brazil: Mato Grosso: Alto Araguaia; savanna creek
25.6 mm SL (one day after collection); Brazil: Mato close to córrego do Sapo, habitat of Rivulus litteratus.
Grosso: Alto Araguaia. Photo by W. J. E. M. Costa. Photo by W. J. E. M. Costa.

aqua vol. 11 no. 4 - 2006 164

 Wilson J. E. M. Costa

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

165 aqua vol. 11 no. 4 - 2006

 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin

fit with any other subgenus of Rivulus. The present gists and still almost inaccessible at present. The geo-
phylogenetic hypothesis indicates that Benirivulus is graphic position of the “rio da Prata” mentioned by
the sister group to a clade including three subgenera, von Ihering may be clarified when examining other
Laimosemion, Owiyeye, and Melanorivulus, each of parts of his paper. For example, among other locali-
them corroborated by both morphological and mol- ties, he cited the occurrence of Phalloceros caudimac-
ecular data. The only species of Benirivulus, R. ulatus (Hensel) for the “rio da Prata, Paraguay” (von
beniensis, was poorly known until now, and conse- Ihering 1931: 246).
quently was not included in molecular analyses. Huber (1992) considered both R. strigatus and R.
Myers (1927b) erected R. beniensis on the basis of dibaphus as synonyms of R. geayi, a species first
specimens collected in the drainage of the Beni- described from Amapá, northern Brazil, but also
Mamoré drainage, rio Madeira basin, and previously occurring in the Guianas (e.g. Huber 1991, 1992).
misidentified by Pearson (1925) as R. strigatus. Based According to Huber (1991), the three nominal
on putative minor differences in dorsal-fin origin species would have near type localities, but in fact the
position, Myers (1927b) recognized two subspecies type locality of R. geayi is about 480 km and 560 km
from distinct localities, R. beniensis beniensis and R. in a straight line from the type locality of R. strigatus
beniensis lacustris. Seegers (1985) described R. boli- and R. dibaphus, respectively, and type locality of R.
vianus also from the río Mamoré drainage. Examina- strigatus is about 710 km from the type locality of R.
tion of material from distant localities along the rio dibaphus. Comparison of material of R. strigatus with
Madeira basin, including the Mamoré, Beni and material from the type locality region of R. geayi
Guaporé river drainages, revealed that R. beniensis is a revealed that R. strigatus is a distinct and valid species,
widespread species, indistinguishable from R. benien- easily distinguished from R. geayi by having a
sis lacustris and R. bolivianus, thus confirming the rounded caudal fin in the male (vs. subtruncate) and
synonymy proposed by Costa (2003b). frontal squamation F-patterned (vs. usually E-pat-
Huber (1999) erected Laimosemion to include terned). Rivulus dibaphus is also a valid species.
species of the “R. geayi superspecies” as proposed by Examination of the type material of R. dibaphus and
Huber (1992). Molecular studies (Murphy et al. material collected in the type locality region revealed
1999, Hrbek et al. 2004) corroborated monophyly of that it may be readily distinguished from R. strigatus
a group comprising species of the “R. geayi super- and R. geayi by the unique colour pattern on the cau-
species”, “R. breviceps superspecies”, and “R. frenatus dal peduncle in the male (Figs 31-32) and that the
superspecies”, which is in accordance to the present frontal squamation is always E-patterned.
study. Two species of Laimosemion are found in the Owiyeye corresponds to the “R. rectocaudatus species
Brazilian Amazonas river: R. strigatus and R. dibaphus. group” diagnosed by Thomerson et al. (1992) and
Rivulus strigatus was described by Regan (1912) the “R. rectocaudatus superspecies” diagnosed by
based on a single specimen donated by Arnold. This Huber (1992). This clade is well supported both by
material was obtained from an aquarium fish ship- morphological (the present study) and molecular
ment from the city of Pará (now Belém, capital of data (Murphy et al. 1999, Hrbek & Larson 1999,
Estado do Pará) (Arnold 1913). Huber (1992) men- Hrbek et al. 2004). Owiyeye and Anablepsoides share
tioned rio da Prata, a tributary of rio Jari, Pará, Brazil, two derived conditions (frontal squamation S-pat-
as the true type locality of R. strigatus, based on a mis- terned and a transverse stripe through chin), but it is
interpreted citation by von Ihering (1931). The lat- parsimoniously considered to a member of a clade
ter author, under a brief diagnosis of R. strigatus, also including Laimosemion and Melanorivulus,
cited: “Distribuição: (typo) Amazonas, sem mais indi- mainly diagnosed by the absence of second pharyn-
cação além do nome do colleccionador Arnold, mas cujo gobranchial teeth. Seven species of Owiyeye are found
material provem principalmente do rio da Prata” (Dis- in the Brazilian Amazon, all described in recent years:
tribution: (type) Amazonas, without further indica- R. romeri, R. kirovskyi, R. uatuman, R. uakti, R.
tion besides the collector name Arnold, but that amanapira, and R. tecminae. Rivulus kirovskyi and R.
material mainly originates from rio da Prata) (von duckensis are identical and were collected at the same
Ihering 1931: 263). In fact, von Ihering referred to locality. Both original descriptions were published in
the río de La Plata basin, from where R. punctatus 2004, but the name R. kirovskyi has chronological
Boulenger was often misidentified as R. strigatus in priority over R. duckensis, since the former was pub-
the past (e.g., Ringuelet et al. 1967), not to the small lished on 30 April, the latter on 20 August.
rio da Prata in Pará, unknown for most ichthyolo- Melanorivulus corresponds to the “R. punctatus

aqua vol. 11 no. 4 - 2006 166

 Wilson J. E. M. Costa

superspecies” diagnosed by Huber (1992) or the the same environmental conditions (Huber 1979).
“Rivulus punctatus species-complex” diagnosed by However, R. agilae, another species of Laimosemion
Costa (1995a). This clade is strongly supported by from Guyana, has been reported as living in sunny
morphology, and is also corroborated by molecular places (Huber 1979).
studies (e.g. Murphy et al. 1999, Hrbek & Larson Only species of Owiyeye were found in extremely
1999). It includes four species endemic to the shallow temporary pools (about 5-10 cm deep) with
Brazilian Amazonas basin: R. modestus, R. zygonectes, tea-coloured water (Figs 36 and 41). Since some
R. violaceus, and R. litteratus. species of Owiyeye were uniquely found in tempo-
As already noted in previous studies (e.g. Costa rary pools, they possibly are semiannual fishes. An
1998a, Murphy et al. 1999, Hrbek & Larson 1999), exception is R. uakti, which inhabit marginal zones
species of Rivulus endemic to the Amazonas basin of creeks at the forest border (Fig. 39).
do not form a monophyletic assemblage. Species Species of Melanorivulus are found in sunny creeks
from southern Amazonas tributaries, running on the (Fig. 45) in savanna-like environments. However,
Brazilian Shield (i.e., belonging to Melanorivulus) this is not a unique condition among Amazonian
are more closely related to species endemic to basins Rivulus, since R. urophthalmus, R. taeniatus, and R.
to South and East (i.e., Paraná, Paraguay, São Fran- micropus are also found in sunny creeks, often near
cisco and Parnaíba river basins). On the other hand, the border of forests (Figs 11, 17 and 20). These
species occurring in northern and western Ama- conditions are interpreted as independently derived,
zonas tributaries arising from the Andes or the because Melanoorus and Cynodonichthys are not
Guyana Shield (i.e., belonging to Cynodonichthys, closely related taxa, and basal rivulids (i.e. basal
Laimosemion and Owiyeye) are more closely related species of Kryptolebias and Prorivulus) inhabit dark
to species endemic to the Orinoco and Guianas forested habitats (Costa 2004a, 2004c, Vermeulen
basin. This is probably a historic consequence of the & Hrbek 2005).
past isolation of rivers draining the Brazilian Shield
before the formation of the present Amazonas basin, Acknowledgements
as described in geological studies (e.g. Beurlen 1970, Thanks are due to E. Araujo, F. Autran, S. Barrera,
Lundberg et al. 1998). D. Belote, C. P. Bove, G. C. Brasil, M. Britto, C.
Field observation on the habitat of Amazonian Campinha, B. B. Costa, R. D. Cunha, R. D’Arrigo,
species of Rivulus revealed that different lineages M. I. Landim, S. M. Q. Lima, C. Moreira, R. Paiva,
occupy distinct kinds of habitat. This is most clear A. Pinto, D. Ramos, A. Sarraf, K. Tanizaki, E.
for the clade Benirivulus + Laimosemion + Owiyeye + Vicente, and L. Villa-Verde, for help in collecting
Melanorivulus, in which members of each genus are trips; to H. Berkenkamp, G. Brasil, H. Britski, J.
found in a particular kind of aquatic environment, Carvalho, D. Catania, J. L. Figueiredo, K. Hartel, J.
thus suggesting divergent specializations during the Huber, M. Kottelat, S. Kullander, M. Lacerda, P.
evolution of the group. Lucinda, N. Menezes, D. Moraes Jr., G. Nunan, O.
In Benirivulus, individuals were collected near the Oyakawa, L. Parenti, C. Pereira, L. Py-Daniel, U.
margin of stagnant streams and lakes (about 50-150 Römer, J. Sarmiento, D. Taphorn, D. Teixeira, and J.
cm deep), with dense aquatic vegetation, at the for- Zuanon, for the loan, exchange, or donation of spec-
est border (Fig. 25). Species of Laimosemion, Owiy- imens, or for support during visits to their institu-
eye and Melanorivulus were collected in shallower tions. Collecting trips were supported by Fundação o
places (about 5-40 cm). Possibly, the ecological pref- Boticário de Proteção à Natureza. This study was
erence noted for Benirivulus is the primitive condi- funded by CNPq-MCT and FAPERJ. Material was
tion for the clade Benirivulus + Laimosemion + collected with permits 02001.001660/98 and
Owiyeye + Melanorivulus, since species of Anablep- 02.022005956/02 from IBAMA (Instituto Brasileiro
soides, the putative sister group to the above men- do Meio Ambiente e dos Recursos Naturais Ren-
tioned clade, live in similar habitats. ováveis – Ministério do Meio Ambiente, dos Recur-
Amazonian species of Laimosemion were uniquely sos Hídricos e da Amazônia Legal).
found in clear water streams, with moderate current
and sandy bottom (Figs 29 and 33). No other References
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167 aqua vol. 11 no. 4 - 2006

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Hoplosternum (Ostariophysi: Siluriformes: Callichthyidae), annual killifishes from Amazonas Territory, Venezuela
with the description of two new genera and three new (Cyprinodontiformes: Rivulidae). Ichthyological Exploration
species. Ichthyological Exploration of Freshwaters, 7: 299-326. of Freshwaters 3: 377-384.
RINGUELET, R. A., ARAMBURU, R. H. & ARAMBURU, A. WALLACE, A. R. 2002. Peixes do Rio Negro/Fishes of the Rio
1967. Los peces argentinos de agua dulce. Comisión de Inves- Negro. EDUSP, São Paulo, 517 pp.
tigaciones Científicas de la Provincia de Buenos Aires, La WEITZMAN, S. H. & WOURMS, J. P. 1967. South American
Plata, 602 pp. cyprinodont fishes allied to Cynolebias with the description
SEEGERS, L. 1984. Zur Revision der Rivulus-Arten Südost- of a new species of Austrofundulus from Venezuela. Copeia
Brasiliens, mit einer Neubeschreibung von Rivulus luelingi 1967: 89-100.
n. sp. und Rivulus caudomarginatus n. sp. (Pisces: Cyprin- WIENS, J. J. (ed.). 2000. Phylogenetic analysis of morphological
odontidae: Rivulinae). Zoologische Beiträge 28: 271-320. data, Smithsonian Institution Press, Washington and Lon-
SEEGERS, L. 1988. Ein neuer Rivulide aus Bolivien: Rivulus don, 220 pp.
bolivianus n. sp. (Pisces: Cyprinodontiformes: Rivulidae). VERMEULEN, F. B. M. & HRBEK, T. 2005 Kryptolebias sepia n.
Bonner Zoologische Beitrage 39: 171-177. sp. (Actinopterygii: Cyprinodontiformes: Rivulidae), a new
STEINDACHNER, F. 1863. Beiträge zur Kenntniss der Sci- killifish from the Tapanahony River drainage in southeast
aenoiden Brasiliens und der Cyprinodonten Mejicos. Surinam. Zootaxa 928: 1-20.
Sitzungsberichte der kaiserlichen Akademie der Wissenschaften VERMEULEN, F. B. M. & ISBRÜCKER, I. J. H. 2000. Rivulus
Classe K. 48: 162-185, plates 1-4. torrenticola n. sp. (Actinopterygii: Cyprinodontiformes:
SWOFFORD, D. L. 1998. PAUP*. Phylogenetic analysis using Rivulidae), a new killifish from highlands in the Guyana
parsimony (*and other methods), Beta version 4.0b2. Sinauer, Shield. Beaufortia 50: 185-190.
Sunderland, Massachusetts. VON IHERING, R. 1931. Cyprinodontes brasileiros (peixes
TAYLOR, W. R. & VAN DYKE, G. C. 1985. Revised proce- “guarús”): sistemática e informações biologicas. Archivos do
dures for staining and clearing small fishes and other verte- Instituto Biologico 4: 242-280.

aqua vol. 11 no. 4 - 2006 170

 Wilson J. E. M. Costa

Appendix 1 aldsiolii: UFRJ 125, 6; UFRJ 6295, 2 (c&s); Brazil: Santa Cata-
The list below includes material examined, except belonging to rina: Joinville. R. hartii: MCZ 26092, 3; Trinidad: Port of Spain.
species of the Brazilian Amazonas basin, which are listed in the MZUSP 37204, 3; MZUSP 38472, 2 (c&s); Venezuela: Nueva
“Taxonomic accounts” above. Data on material is organized in Esparta: Isla de Margarita. R. hildebrandi: USNM 92958, 1P;
the following sequence: catalogue number, number of speci- Panama: Chiriqui: Boquete. R. holmiae: USNM 66302, 1P;
mens, locality. Abbreviations are: c&s, specimens cleared and Guiana: Holmia. R. igneus: UFRJ 3888, 6; UFRJ 4595, 2 (c&s);
stained for bone and cartilage, H, holotype, and P, paratype(s). Suriname: Creek Colibri. R. immaculatus: USNM 308411, 2P;
Institutional acronyms are listed in material and methods. Venezuela: Bolivar: Elena de Uairen. R. janeiroensis: UFRJ 5333,
8; UFRJ 130, 2 P (c&s); UFRJ 5416, 7 (c&s); Brazil: Rio de
Aplocheilidae: Aplocheilus panchax: UFRJ 3140, 2; UFRJ Janeiro: Magé. R. luelingi: UFRJ 161, 8, UFRJ 127, 5 (c&s);
3141, 2 (c&s); Indonesia: Sulawesi: Desa Radda. Notho- Brazil: Santa Catarina: Araquari. R. lungi: ZSM 27825, 15;
branchiidae: Scriptaphyosemion guignardi: UFRJ 3883, 8; UFRJ Brazil: Amapá: rio Flechal. R. nicoi: MCNG 23891, H; MCNG
4110, 4 (c&s); Guinea: Dalaba. Rivulidae: Aphyolebias boticarioi: 23892, 1P; MCNG 23893, 1P; Venezuela: Amazonas: río Ven-
UFRJ 5986, H; UFRJ 5987, 11 P; UFRJ 5988, 5 P; Brazil: tuari floodplains. R. nudiventris: MZUSP 40283, H; MZUSP
Estado do Acre: Porto Acre. Gnatholebias hoignei: MCNG 1116, 40284, 3P; MNRJ 11740, 2P (c&s); Brazil: Espírito Santo:
18; Venezuela: Portuguesa: La Trinidad. UFRJ 6116, 6; UFRJ Itapemirim. R. paracatuensis: MCP 29639, H; MCP 29640, 1P;
6117, 5 (c&s); Venezuela: Portuguesa: Papelón. Kryptolebias UFRJ 2290, 3P (c&s); Brazil: Minas Gerais: Brasilândia de
brasiliensis: UFRJ 3458, 32; UFRJ 3682, 1 (c&s); UFRJ 4603, Minas. R. parnaibensis: MCP 29639, H; UFRJ 4962, 25P;
2 ex. (c&s); UFRJ 5332, 6 ex. (c&s); Brazil: Rio de Janeiro: UFRJ 5449, 4P (c&s); Brazil: Piauí: São Dimas. R. pictus:
Magé. Micromoema xiphophora: MCNG 26440, 20; Venezuela: MNRJ 11550, H; MNRJ 11551, 2P; UFRJ 5959, 2 ex. (c&s);
Amazonas: Isla Ratón. UFRJ 3165, 1 (c&s); aquarium material. Brazil: Distrito Federal: Planaltina. R. pinima: MZUSP 39978,
Moema apurinan: UFRJ 5980, H; UFRJ 5981, 9 P; UFRJ 5982, H; MZUSP 39984, 2P; UFRJ 5960, 20; UFRJ 5961, 4 (c&s);
7 P (c&s); Brazil: Estado do Acre: Porto Acre. Neofundulus Brazil: Goiás: Rio Verde. R. punctatus: UFRJ 975, 9; UFRJ
paraguayensis: UFRJ 3647, 10; UFRJ 3648, 4 (c&s); Brazil: 2110, 4 (c&s); Brazil: Mato Grosso do Sul: Aquidauana. R.
Mato Grosso do Sul, about 70 km NW from de Aquidauana. rossoi: UFRJ 5976, H; UFRJ 5977, 7P; UFRJ, 5978, 4P (c&s);
Pituna compacta: UFRJ 3563, 33; UFRJ 3564, 4 (c&s); Brazil: Brazil: Mato Grosso do Sul: Campo Grande. R. rubrolineatus:
Tocantins: Barreira do Piqui. Prorivulus auriferus: UFRJ 5932, MZUSP 26371, 3; MUSM 1465, 6; Peru: Loreto: Jenaro Her-
H; UFRJ 5933, 4P; UFRJ 5934, 3P (c&s); Brazil: Bahia: rera. R. rutilicaudus: UFRJ 5965, H; UFRJ 5966, 12P; UFRJ
Valença. Rachovia maculipinnis: MCNG 35565, 2; Venezuela: 5967, 3P (c&s); Brazil: Goiás: Serranópolis. R. santensis: UFRJ
Portuguesa: Sabaneta. UFRJ 6118, 7; UFRJ 6119, 4 (c&s); 5441, 11; UFRJ 6294, 5 (c&s); Brazil: São Paulo: Boracéia. R.
Venezuela: Portuguesa: Papelón. R. stellifer: MCNG 25828, 9; simplicis: UFRJ 5940, H; UFRJ 5976, 3P; UFRJ 5942, 5P
Venezuela: Portuguesa: La Capilla. UFRJ 245, 5 (c&s); (c&s); Brazil: Rio de Janeiro: Parati. R. scalaris: UFRJ 5968, H;
Venezuela: Cojedes: 1 km N of Caño Benito. Renova oscari: UFRJ 5969, 19P; UFRJ 5970, 4P (c&s); Brazil: Mato Grosso do
MCNG 35926, 2; Venezuela: Amazonas: Isla Ratón. UFRJ Sul: Costa Rica. R. stagnatus: UFRJ 3890, 3; UFRJ 4605, 4
4606, 3 (c&s): aquarium material. Rivulus amphoreus: UFRJ (c&s); Suriname: Wageningen. R. tenuis: UFRJ 4600, 4; UFRJ
3889, 2, UFRJ 4606, 3 (c&s); Surinam: Tafelberg. R. apiamici: 4601, 2 (c&s); Guatemala: Alta Verapaz: Sebol. R. vittatus:
MZUSP 39976, H; MZUSP 39977, 3P (c&s); UFRJ 5971, 4; MZUSP 39981, H; MZUSP 39981, 6P; UFRJ 2206, 15; UFRJ
UFRJ 5972, 3 (c&s); Brazil: Mato Grosso do Sul; Bataguaçu. R. 5962, 4 (c&s); Brazil: Goiás: Cachoeira Alta. R. xiphidius: UFRJ
bahianus: UFRJ 3167, 44; UFRJ 4602, 2 (c&); UFRJ 277, 1 4608, 3 (c&s); Guyana (specimens born in aquarium).
(c&s); Brazil: Bahia: Busca-Vida. R. brunneus: MZUSP uncata- Trigonectes rubromarginatus: UFRJ 3553, 13; UFRJ 3554, 3
logued, 10 (2 c&s); Panama: Isla de Barro Colorado. R. chucu- (c&s); Brazil: Tocantins: Barreira do Piqui.
naque: USNM 293487, 68 (1c&s); Panama. R. cladophorus:
UFRJ 643, 4P (2c&s); Guyana: Fourgassier. R. cryptocallus: Appendix 2
UFRJ 359, 3; UFRJ 2126, 1 (c&s); Martinique: Ravine Vilaine. Characters (in brackets) and character states (in parentheses)
R. cyanopterus: UFRJ 5911, H; UFRJ 5913, 36P; UFRJ 5914, used to erect the phylogenetic hypothesis among species of Rivu-
5P (c&s); Brazil: Mato Grosso: Jaciara. R. cylindraceus: MHNC lus and other rivulids are listed below, with the respective refer-
uncat., 6; Cuba: Zapata. USNM uncat., 1 (c&s); Cuba: La ence to papers where the character is first described or discussed.
Habana. R. dapazi: UFRJ 5915, H; UFRJ 5916, 3P; UFRJ Distribution of character states among terminal taxa is presented
5917, 2P (c&s); Brazil: Mato Grosso do Sul: Sonora. R. decora- in the data matrix in Appendix 3.
tus: MZUSP 39982, H; MZUSP 39983, 4P; UFRJ 2135, 3
(c&s); Brazil: Bahia: Ibiraba. R. derhami: UFRJ 392, 2 (c&s); Superficial dermal bones and neurocranium
Peru: Tingo Maria. R. depressus: UFPB 2213, H; UFPB 1749, 17 [1] Lachrymal (Parenti 1981; Costa 1998a, 1998b) (CI: 1.00;
(2 c&s); UFRJ 2118, 1 (c&s); Brazil: Bahia: Porto Seguro. R. RI: 1.00): (0) flat, posterior rim wide; (1) slightly twisted, poste-
elongatus: MUSM 336, 6; MZUSP 26211, 3; Peru: Ucayali: rior rim reduced, bone formed mainly by canal; (2) very twisted
Pucallpa. R. egens: UFRJ 5973, H; UFRJ 5974, 17P; UFRJ and narrow, slender, canal vestigial.
5975, 3P (c&s); Brazil: Mato Grosso do Sul: Camapuã. R. erberi: [2] Ventral portion of lachrymal (Costa 1998a) (CI: 0.50; RI:
UFRJ 358, 3; Ecuador: Napo: Coca. R. fuscolineatus: USNM 0.66): (0) short; (1) expanded.
219778, 10P; Costa Rica: Guanacaste: Tilaran. R. geayi: [3] Dermosphenotic (Costa 2004d): (CI: 1.00; RI: 1.00): (0)
MZUSP uncatalogued Brazil: Amapa: Serra do Navio. R. har- present; (1) absent.

171 aqua vol. 11 no. 4 - 2006

 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin

[4] Vomerine teeth (Costa 1998a) (CI: 0.40; RI: 0.80): (0) usu- [24] Preopercle (Costa 1990) (CI: 1.00; RI: 1.00): (0) robust,
ally 1-4, sometimes 5-6; (1) 6-12; (2) teeth absent [not ordered]. L-shaped, with a well-developed anteromedian rim; (1) thin, C-
[5] Anterior retrorse process of lateral ethmoid (Costa 1990) shaped, with a reduced anteromedian rim.
(CI: 1.00; RI: 1.00): (0) short; (1) moderate to elongate. [25] Dorsal arm of preopercle (Costa 1990) (CI: 0.50; RI:
[6] Lateral process of sphenotic (Costa 2005b) (CI: 0.50; RI: 0.85): (0) broad; (1) narrow and pointed.
0.50): (0) narrow; (1) wide. Hyoid and branchial arches
[7] Anterolateral process of parasphenoid (Costa 1998a) (CI: [26] Anterior process of urohyal (Costa 1998a) (CI: 1.00; RI:
0.16; RI: 0.78): (0) short, free; (1) long, attached to pterosphe- 1.00): (0) short; (1) elongate.
noid. [27] Dorsal process of urohyal length (CI: 1.00; RI: 1.00): (0)
[8] Posterior portion of parasphenoid (Costa 2005b) (CI: 0.50; present; (1) absent; (?) very short to absent.
RI: 0): (0) wide; (1) narrow. [28] Dorsal process of urohyal length (Costa 1998a) (CI: 0.25;
[9] Lateral border of frontal (Costa 1998a) (CI: 1.00; RI: RI: 0.85): (0) short; (1) elongate; (?) process absent.
1.00): (0) well-ossified, approximately straight; (1) poorly ossi- [29] Basihyal (Costa 1998a) (CI: 0.50; RI: 0): (0) shorter than
fied, concave. space occupied by basibranchials; (1) longer than space occupied
Jaws, jaw suspensorium and opercular apparatus by basibranchials.
[10] General shape of the premaxilla and dentary (Costa [30] Basihyal cartilage extent, as percentage of basihyal length
1998a) (CI: 0.25; RI: 0): (0) elongate, snout profile sharply (modified from Costa 1998a) (CI: 0.40; RI: 0.62): (0) 50-70 %;
pointed; (1) short, snout profile blunt. (1) 20-40 %; (2) 10-15 %.
[11] General shape of ascending process of premaxilla (modi- [31] Interhyal (Parenti 1981) (CI: 1.00; RI: 1.00): (0) ossified;
fied from Parenti, 1981, Costa 1998b) (CI: 1.00; RI: 1.00): (0) (1) cartilaginous.
curved, posterior portion medially directed; (1) approximately [32] Interhyal (Costa 2005b) (CI: 1.00; RI: 1.00): (0) large; (1)
straight. minute.
[12] Ascending process of premaxilla (Costa 2005b) (CI: 0.50; [33] Number of branchiostegal rays (Costa 2004e) (CI: 0.33;
RI: 0): (0) wide; (1) narrow. RI: 0): (0) 6; (1) 5.
[13] Ventral process of maxilla (Parenti 1981, Costa 1998a) [34] Main axis of first epibranchial (Costa 1998a) (CI: 1.00;
(CI: 1.00; RI: 1.00): (0) slightly curved, anterior margin RI: 1.00): (0) approximately straight; (1) curved.
rounded; (1) bent, anterior margin triangular. [35] Subdistal process of second epibranchial (Costa 2004a)
[14] Maxilla (modified from Parenti 1981) (CI: 0.40; RI: (CI: 1.00; RI: 1.00): (0) present; (1) absent.
0.72): (0) approximately straight; (1) slightly twisted; (2) greatly [36] Uncinate process of third epibranchial (modified from
twisted. Costa 1998a, b) (CI: 1.00; RI: 1.00): (0) long; (1) moderate; (2)
[15] Rostral cartilage (Costa 1998a) (CI: 0.25; RI: 0.40): (0) short.
approximately rounded; state 1: longitudinal length longer than [37] Interarcual cartilage (modified from Parenti 1981) (CI:
transversal length. 1.00; RI: 1.00): (0) not reduced; (1) reduced (CI: 1.00; RI:
[16] Posterior portion of rostral cartilage (Costa 2004a) (CI: 1.00).
0.20; RI: 0.69): (0) not distinctively narrowed; (1) distinctively [38] Number and arrangement of second pharyngobranchial
narrowed. teeth (Costa 2004a) (CI: 0.50; RI: 0.88): (0) numerous teeth
[17] Coronoid process of dentary (Costa 1998a) (CI: 1.00; RI: arranged in two rows; (1) few teeth arranged in single row; (2)
1.00): (0) broad; (1) narrow. teeth absent [not ordered].
[18] Ventral process of angulo-articular (modified from Costa [39] Proximal edge of first hypobranchial (Costa 1998a) (CI:
1990, 1998a) (CI: 0.66, RI: 0.81): (0) large and broad; (1) large, 0.33; RI: 0.33): (0) plain, terminating in single cartilage united
somewhat narrowed; (2) moderate, narrow and pointed; (3) ves- to second basibranchial; (1) bifid, terminating in cartilage united
tigial; (4) long and somewhat narrowed [not ordered]. to second basibranchial and another smaller cartilage united to
[19] Curvature of ventral process of angulo-articular (Costa first basibranchial.
1998a) (CI: 1.00, RI: 1.00): (0) straight; (1) curved. [40] Distal edge of first hypobranchial (Costa 2004a) (CI:
[20] External medial teeth of premaxilla and dentary (Costa 1.00; RI: 1.00): (0) articular face restricted to cartilaginous
1998a) (CI: 0.33; RI: 0.50): (0) approximately directed as head of first ceratobranchial; (1) articular face anteriorly
other teeth; (1) laterally displaced, strongly contrasting to expanded.
other teeth. [41] Orientation of anterior tip of fifth ceratobranchial (Costa
[21] Ventral portion of palatine (Costa 1998a) (CI: 1.00; RI: 2005b) (CI: 0.50; RI: 0.50): (0) anterior; (1) anterolateral.
1.00): (0) long, overlapping dorsal portion of quadrate; state 1: Vertebrae and caudal skeleton
short, not or slightly contacting quadrate. [42] Pointed, anteriorly directed process on first vertebra
[22] Metapterygoid (modified from Costa 1998a) (CI: 0.66; (modified from Costa 1990) (CI: 1.00; RI: 1.00): (0) absent;
RI: 0.90): (0) about rectangular, dorsal and ventral portions wide (1) present.
and approximately equal in width; (1) dorsal portion slightly [43] Epipleural ribs (Parenti 1981, Costa 1998a) (CI: 0.50;
constricted; (2) about triangular, dorsal portion strongly con- RI: 0.66): (0) rod-like; (1) bifid.
stricted. [44] Neural prezygapophyses (Costa 1990) (CI: 0.33; RI:
[23] Posterior process of quadrate (Costa 1998a) (CI: 0.50; RI: 0.84): (0) short; (1) long.
0.50): (0) short, about 50% of quadrate length; (1) long, about [45] Hypurals (modified from Costa 1998a) (CI: 0.42; RI:
70% of quadrate length. 0.33): (0) two dorsal plates and one ventral plate separated by

aqua vol. 11 no. 4 - 2006 172

 Wilson J. E. M. Costa

gap; (1) two plates separated by wide gap; (2) two plates in close [67] Skin fold on corner of preopercular region (Costa
proximity, sometimes ankylosed; (3) single plate. 2005b) (CI: 1.00; RI: 1.00): (0) present; (1) absent.
[46] Proximal end of parahypural (Costa 1998b) (CI: 1.00; External morphology of fins
RI: 1.00): (0) robust with paired dorsal processes overlapping [68] Pectoral fin (Costa 1990) (CI: 0.50; RI: 0.75): (0)
preural centrum; (1) shortened and laminar, without dorsal rounded; (1) pointed.
paired process, not contacting preural centrum. [69] Pectoral-fin length (modified from Parenti 1981) (CI:
[47] Hemal spine of preural centrum two (Costa 1998b) (CI: 0.33; RI: 0.75): (0) 17.0–23.8 % SL; (1) 24.1–31.1 % SL.
0.50; RI: 0): (0) distinctively wider than hemal spines anterior [70] Extent of pelvic fin in males (Costa 1998a) (CI: 0.25; RI:
to it; (1) slightly wider or equal in width to hemal spines ante- 0.25): (0) short, tip not surpassing anterior portion of anal fin;
rior to it. (1) long, its tip reaching the central or the posterior portion of
[48] Number of vertebrae (modified from Costa 1990) (CI: the anal fin.
0.18; RI: 0.52): (0) 29-32; (1): 33-35; (2) 36-38. [71] Pelvic-fin (Costa 2005b) (CI: 1.00; RI: 1.00): (0) bases
[49] Number of caudal-fin rays (modified from Costa 1990) separated or in contact; (1) bases united; (2) pelvic fins united
(CI: 0.18; RI: 0.59): (0) 26-31; (1) 32-36; (2) 24-25 [not along proximal portion of medial margin.
ordered]. [72] Dorsal fin (modified from Costa 1998a) (CI: 0.33; RI:
Dorsal- and anal-fin skeleton 0.84): (0) short, tip rounded; (1) somewhat elongated, tip
[50] First dorsal-fin ray (modified from Parenti, 1981) (CI: pointed; (2) long, tip sharply pointed.
1.00; RI: 1.00): (0) single long first ray connected to two prox- [73] Anal fin (modified from Costa 1998a) (CI: 0.22; RI:
imal radials; (1) long fin ray connected to two proximal radials, 0.75): (0) short, tip rounded; (1) somewhat elongated, tip
preceded by one or two short fin rays. pointed; (2) long, tip sharply pointed.
[51] Anterior proximal radials of dorsal and anal fins (Costa [74] Filaments on tip of dorsal and anal fins in males (Costa
2005b) (CI: 1.00; RI: 1.00): (0) slender; (1): wide. 1998a) (CI: 0.50; RI: 0.66): (0) absent; (1) present.
[52] Number of anal-fin rays (modified from Costa 1998a) [75] Filaments on posterior border of caudal fin in males
(CI: 0.28; RI: 0.44): (0) 9-15; (1) 16-20; (2) 25-26. (Costa 1998a) (CI: 0.33; RI: 0.60): (0) absent; (1) present.
[53] Orientation of anterior proximal radials of anal fin [76] Caudal-fin length in males (Costa 1990) (CI: 0.40; RI:
(modified from Costa 1998a) (CI: 1.00; RI: 1.00): (0) anteri- 0.70): (0) 29.0-41.5; (1) 42.0-49.0; (2) 52.5-81.0.
orly or dorsally directed; (1) posteriorly directed. [77] Caudal-fin shape in males (CI: 0.33; RI: 0.40): (0)
Shoulder and pelvic girdle rounded; (1) subtruncate; (2) truncate; (3) acuminate [not
[54] Pectoral-fin insertion (Costa 1998b) (CI: 1.00; RI: ordered].
1.00): (0) lateral; (1) ventrolateral. [78] Dorsal and ventral extensions on caudal fin in males
[55] Supracleithrum and posttemporal (modified from Par- (Costa 1998a) (CI: 0.50; RI: 0): (0) absent; (1) present.
enti 1981, Costa 1998b) (CI: 0.66; RI: 0.50): (0): separated; Squamation
(1) co-ossified, limits almost inconspicuous; (2) fused to form [79] General arrangement of frontal scales (Hoedeman 1958)
a single structure. (CI: 0.50; RI: 0.85): (0) transverse; (1) circular; (2) irregular
[56] Keel along supracleithrum-posttemporal (Costa 2005b) [not ordered].
(CI: 1.00; RI: 1.00): (0) absent; (1) present. [80] Predominant frontal squamation-pattern (Hoedeman
[57] Ventral process of posttemporal (Parenti 1981, Costa 1958) (CI: 0.28; RI: 0.52): (0) G; (1) E; (2) D; (3) F; (4) S [not
1998a, b) (CI: 0.20; RI: 0.81): (0) present; (1) absent. ordered].
[58] Posterior flange of cleithrum (Costa 1998a) (CI: 1.00; [81] Arrangement of E-scales (Costa 1990) (CI: 1.00; RI:
RI: 1.00): (0) present; (1) absent. 1.00): (0) overlapped; (1) not overlapped.
[59] First postcleithrum (Parenti 1981) (CI: 1.00; RI: 1.00): [82] Caudal-fin squamation in older males (Costa 1990,
(0) present; (1) absent. 1998a) (CI: 0.25; RI: 0.57): (0) approximately on anterior
[60] Fourth pectoral radial (Costa 1998a) (CI: 0.33; RI: 10–30 % of fin; (1) approximately on 40 % of fin; (2) approx-
0.75): (0) not expanded; (1) ventrally expanded. imately on 50-80 % of fin.
[61] Ischial process of pelvic girdle (Costa 2005b) (CI: 1.00; [83] Anal-fin base squamation in males (Costa 2005b) (CI:
RI: 1.00): (0) prominent; (1) vestigial. 1.00; RI: 1.00): (0) no scales on anal-fin base; (1) 1-7 rows of
[62] Number of pectoral-fin rays (modified from Costa scales on anal-fin base.
1998a): (0) 13-15; (1) 16-17. Laterosensory system
[63] Number of pelvic-fin rays (Costa 1990) (CI: 1.00; RI: [84] Cephalic canals (modified from Parenti 1981) (CI:
1.00): (0) six; (1) seven; (2) eight. 0.75; RI: 0.92): (0) anterior and posterior infraorbital, preop-
External morphology of body and head ercular and posterior mandibular closed, supraorbital, rostral,
[64] Mouth (Costa 1998b) (CI: 1.00; RI: 1.00): (0) terminal; anterior mandibular open with skin trenches around neuro-
(1) superior. masts; (1) anterior and posterior infraorbital, and preopercu-
[65] Orbital rim (modified from Parenti 1981) (CI: 1.00; RI: lar closed, supraorbital, rostral, anterior and posterior
1.00): (0) free dorsally, attached ventrally; (1) completely mandibular open with skin trenches around neuromasts; (2)
attached. anterior and posterior infraorbital, dorsal preopercular, supra-
[66] Branchiostegal and opercular membranes (Parenti orbital, rostral, anterior and posterior mandibular open with
1981) (CI: 1.00; RI: 1.00): (0) separated by long fold; (1) skin trenches around neuromasts; (3) no vestige of canals, all
continuous. neuromasts completely exposed.

173 aqua vol. 11 no. 4 - 2006

 Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin

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

aqua vol. 11 no. 4 - 2006 174

 1–10 11–20 21–30 31–40 41–50 51–60 61–70 71-80 81-90 91-100 101-110 111-113

A. panchax 0000000000 0000000000 0000000000 0000000000 0000000000 0100201000 0000000000 0000000000 000000000? 0000000100 0000000000 00?

175
S. guignardi 1000000001 0100000000 0000000000 0000000200 0010111011 0101200000 0101101000 0000001000 0001000011 0000001000 1101000000 00?
K. brasiliensis 2000100011 0111101000 100100?002 0000010110 0001110001 0001000010 0001111000 0000000011 000200000? 1000010101 0010000000 00?
P. auriferus 2000101011 1112101000 1001000001 1110110110 0100111011 0001001010 00011110?0 0000000011 000200000? 0000000100 0000000100 00?
A. boticarioi 2100101011 1111101100 1201000101 1100120111 1100211101 0001110111 0001111110 0221111111 1002000113 0000100000 1001001010 00?
G. hoignei 2000100110 1011001101 1101000101 11001212?1 0100311201 1211110111 1011111111 2221121013 101200000? 0000100000 0000010000 00?
M. xiphophora 2000100011 1112101101 1111000111 1100120111 0100111001 0001111111 0001111110 0220013111 1002000010 0000121000 1101000001 00?
M. apurinan 200?101110 1111101100 1201000101 1100120111 1100111101 0101110111 0001111110 0221120011 1002000110 0000101000 1001000000 00?
N. paraguayensis 2000111011 1111101100 1001000101 1100120111 0100111101 0001110110 0001111010 0110000011 1102100010 1000100000 1000001000 00?
P. compacta 2100110011 1111101300 1101100101 1100120111 0100211001 0001110111 0001111010 0110000013 100320010? 0000110000 0000001000 00?
P. longipinnis 2000101011 1112101201 1211000111 1100121111 0100111001 1111110111 1011111011 1221121011 101200000? ?000100001 0000002000 00?
R. maculipinnis 2100101011 1111101100 1101010101 1100120111 0100211001 0001110111 0001111010 0110111013 120200000? 0000112000 ?100000010 00?
R. stellifer 2100101011 1111101100 1101010101 1100120111 0101211101 0101110110 0001111000 0110001011 120200000? 0000110001 0000000000 011
R. oscari 2001101011 1112101100 1111000101 1100120111 1100111011 0001110111 0001111110 0220110013 1002000010 0000120000 1001000001 011
R. amanapira 2010100011 1112101100 1001000001 1100120201 0101111201 0001111110 0001111000 0110002004 1003001110 0011000200 1100000000 011
R. amphoreus 2000101011 1112111400 1001000102 1100120111 0101111111 0101110110 0101111000 0000000011 1102001110 0000000000 0000000001 011
R. apiamici 2002100011 1112101300 1001100001 1101120211 0101111011 0001111110 0001111000 0110000011 1003001012 0000001100 0000100001 112
R. atratus 2000101011 1112011101 1001000001 1100120111 0101111021 0001111110 0001111000 0000013014 100200100? 0310000200 0000001000 00?
R. bahianus 2000101011 1112111100 1001000002 1100120111 0101111001 0001110110 0001111000 0000000011 1102001010 0010000001 1110000000 011
R. beniensis 2000100011 1112111100 1001000101 1100120111 0101111021 0001111110 0001111000 0000000011 1002001110 0010000101 0000000000 011
R. brunneus 2000101011 1112111100 1001000101 1100120111 0101111001 0001110110 0001111000 0000000012 1102001010 0000000110 1101000101 011
R. cladophorus 2002100011 1112101100 1001000001 1100120211 0111111001 0001111110 0001111000 0000001013 1002001111 1010000100 0010000001 00?
R. cryptocallus 2000101011 1112111400 1001000102 1100120111 0101111111 0001110110 0001111000 0000000011 1102001110 0110000000 1100000001 011
R. cylindraceus 2000100011 1112001100 1001000001 1100120111 0100311001 0001110110 0001111000 0000000011 1202100014 0000000100 0000000000 010
R. depressus 2000101011 1112111110 1001000001 1100120111 0101111101 0001111110 0001111000 0000000011 1002001014 0010000101 1110010000 011
R. dibaphus 2002100011 1112101100 1001001?01 1100120211 0111111001 0001111110 0001111000 0110001011 1002001111 1010000100 0010100001 00?
R. hartii 2000101011 1112101400 1001000102 1100120111 0101111111 0101110110 0101111000 0000000012 1102001110 0110000010 1100000001 011
Wilson J. E. M. Costa

R. igneus 2001101011 1112101400 1001000102 1100120111 0101111211 0101110110 0101111000 0000000012 1102001110 0100000010 0000000001 00?
R. janeiroensis 2000101011 1112111110 1001000001 1100120111 0101?11101 0001111110 0001111000 0000000011 1002001014 0010000101 1110010000 011
R. kirovskyi 2012100011 1112101100 1001000001 1100120211 0101111021 0001111110 0001111000 0000000004 1003011110 0010000200 0000000000 00?
R. litteratus 2002100011 1112101300 1001100001 1100120211 0101111011 0001111110 0001111000 0110000013 1003001014 0000001100 0000100001 112
R. luelingi 2000101011 1112111110 1001000101 1100120111 0101111001 0001111110 0001111000 0000000011 1002001114 0010000101 1110010000 012
R. micropus 2000101011 1112111400 1001000101 1100120111 0101111101 0001110110 0001111000 0000000011 1102001110 0010000010 0000000001 011
R. modestus 2002100011 1112101100 1001100001 1100120211 0101111011 0001111110 0001111000 0110000013 100300100? 0000001100 0000000000 112
R. ornatus 2000101011 1112011101 1001000001 1100120111 0101111021 0001111110 0001111000 0000013014 1002001013 0310000210 0000100000 00?
R. pictus 2002100011 1112101300 1001100001 1101120201 0101111011 0001111110 0001111000 0110000011 1003001012 0000001100 0000100001 112
R. punctatus 2002100011 1112101100 1001100001 1100120211 0101111011 0001111110 0001111000 0110000013 1003001012 0200001100 0000000001 112
R. romeri 2012100011 1112101100 1001000001 1110120211 0101111021 0001111110 0001111000 0020000004 1003011010 0010000200 0000000000 00?
R. santensis 2000101011 1112101110 1001000?01 1100120111 0101?11111 0001111110 0001111000 0000000011 1002001014 0010000101 1110010000 011
R. strigatus 2002100011 1112101100 1001001?01 1100120211 0111111001 0001111110 0001111000 0110000013 1002001111 1010000100 1010100001 00?
R. taeniatus 2000101011 1112111100 1001000102 1100120111 0101111101 0001110110 0001111000 0000000011 1002001110 0010000010 0100000101 011
R. tecminae 2012100011 1112101100 1001000001 1100120211 0101111021 0001111110 0001111001 0000002004 1003001110 0011000200 1100000000 011
R. tenuis 2000101011 1112111100 1001000101 1100120111 0101111001 0001110110 0001111000 0000000011 1102001010 00?0000110 1101000101 011
R. uakti 201?100011 1112101100 1001000001 1100120211 0101111021 0001111110 0001111001 0010000004 1003001114 0011000200 0000000000 011
R. uatuman 2012100011 1112101100 1001000001 1110120211 0101111021 0001111110 0001111000 0000000004 1003011110 0010000200 0000000000 00?
R. urophthalmus 2000101011 1112111100 1001000102 1100120111 0101111101 0001110110 0001111000 0000000011 1002001010 0110000010 1100000101 011
R. violaceus 2002100011 1112101100 1001100001 1100120211 0101111011 0001111110 0001111000 0110000013 1003001012 0000001100 1100100001 112
R. zygonectes 2002100011 1112101100 1001100001 1100120211 0101111011 0001111110 0001111000 0110000013 1003001012 0200001100 0000000001 112

aqua vol. 11 no. 4 - 2006

T. rubromarginatus 2001111010 1111101400 1001000101 1100120111 0100111211 0101110111 0001111001 0220000012 1102100010 0000100000 1000000000 00?
aqua vol. 11 no. 4 - 2006 176
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 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 ................................................................................................... 133-175

Papers appearing in this journal are indexed in: Zoological Record;

Biolis – Biologische Literatur Information Senckenberg;
www.aquapress-bleher.it; www.aquageo.com; www.Joachim-Frische.com

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. 133-181.