Rivulus amanapira Costa, 2004

Transcrição

Wilson J. E. M. Costa
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. 2003.
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, 0°2’58.7”S
66°57’48.6”W; W. J. E. M. Costa, S. Lima and L.
Silva, 30 Aug. 2003.
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 3233, gill rakers of first branchial arch 1 + 7-8, 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. 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
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.
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,
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. 2003.
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.
159
Diagnosis
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Killifish genus Rivulus (Rivulidae) from the Brazilian Amazonas river basin
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 11th and 13th 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).
Habitat notes
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.
Small shallow temporary pools in the forest,
about 0.5-2.0 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
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 confluence with río Sipapo, Orinoco basin; L. Nico & E.
Guayamare, 28 May 1989. Brazil: Estado do Amazonas: UFRJ 2085, 2; UFRJ 2095, 1 (c&s);
Inambu, right bank of upper rio Negro; D. Teixeira,
26 Aug. 1991.
Diagnosis
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.
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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, 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
160
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, 1895.
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,
161
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, 1927.
Distribution
Paraná-Paraguay-Uruguay, upper Tapajós, upper
and middle Xingu, upper and middle Tocantins,
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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. 1989.
Diagnosis
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 1 + 7-8, 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).
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Material examined
Brazil: Tocantins-Araguaia basin: Estado de Goiás:
CAS 76314, lectotype; Vereda Extrema into Cana
Brava; C. Ternetz, 14 Jan. 1924. UFRJ 1575, 3;
stream 15 km E of Aruanã; W. J. E. M. Costa et al.,
29 Aug. 1993. 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. 1993. UFRJ 1567, 5;
stream 98 km W of Jussara; W. J. E. M. Costa et al.,
30 Aug. 1993. 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. 1993. MZUSP 35416, 20; MZUSP 38334,
55; pool in Aruanã; W. J. E. M. Costa et al., 28 Jan.
1986. 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. 1994. MZUSP 38375, 2;
stream tributary to córrego Dona Francisquinha,
Porto Nacional; W. J. E. M. Costa et al., 31 Jan.
1986. 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. 1993. UFRJ 1597, 2; stream 12
km N of Sandolândia; W. J. E. M. Costa et al., 27
Aug. 1993. 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. 1976.
MZUSP 45223, 9; rio Água Fria, road AraguaçuBarreira do Piqui, 27 km N of Araguaçu; F. C. T.
Lima, 21-26 Feb. 1993. 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.
1993. UNT 555, 5; river at the road BR-153
between Filadelphia and Araguaina; E. L. Beerli, 2
Nov. 2003. UNT 556, 3; rio Brejão, Araguaina; E.
L. Beerli & L, M, Lima, 22 Nov. 2003. UNT 2045,
1; córrego Gorgulho, Porto Nacional; NEAMB, 14
Sep. 2001. UNT 2046, 1; córrego Gorgulho, Porto
Nacional; NEAMB, 14 Sep. 2001. UNT 2047, 3;
córrego Marimbondo, Tupirantins; NEAMB, 27
Oct. 2000. UNT 2048, 1; córrego Barreiro, Guaraí;
NEAMB, 20 Oct. 2000. UNT 2049, 2; córrego
Gorgulho, Porto Nacional; NEAMB, 14 Sep. 2001.
UNT 2050, 1; córrego Gorgulho, Porto Nacional;
NEAMB, 14 Sep. 2001. UNT 2051, 1; córrego
Lageado, Paranã; NEAMB, 22 Mar. 1999. UNT
2052, 9; córrego Água Suja, Tupirantins; NEAMB,
27 Oct. 2000. UFRJ 5150, 1; floodplains of left
bank of rio Tocantins, about 1200 m from the river
margin, SSE Sampaio, 5°23’00”S 47°51’41”W; G.
C. Brasil, 8 Jun. 2000. Estado do Maranhão: UFRJ
5152, 16; swamp about 500 m from the right margin of rio Tocantins, 5°12’33”S 48°27’59”W; G. C.
162
Brasil, 8 Jun. 2000. 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. 1993.
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. 1993. UFRJ 1417, 1; stream at the road BR080, 2 km E of São José do Xingu; W. J. E. M. Costa
et al., 18 Feb. 1993. 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. 1993. 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. 1993. 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.
1993. UFRJ 1357, 20; stream at the road Paranatinga-Canarana, 158 km of Paranatinga; W. J. E.
M. Costa et al., 12 Feb. 1993. 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.
1993. UFRJ 6265, 1; stream near rio Xingu,
Altamira; W. J. E. M. Costa et al., 16 Jun. 2004. Rio
Tapajós basin, Estado de Mato Grosso: MZUSP
45304, 10; upper rio Preto, road Cuiabá-Santarém;
N. Menezes et al., 24 Oct. 1992. MZUSP 45321, 2;
stream tributary to rio Preto, road to São Francisco;
N. Menezes et al., 24 Oct. 1992.
Diagnosis
origin on vertical between base of 8th and 10th 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 3335, gill rakers of first branchial arch 1 + 7-8, 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. 1989. 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. 1989. 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. 1993. UFRJ 4284, 13; idem; M. Britto, C.
Moreira & R. Cunha, 28 Oct. 1996. UFRJ 4283,
3; stream at the road Primavera do Leste-Paranatinga; M. Britto, C. Moreira & R. Cunha, 28
Oct. 1996.
Diagnosis
urogenital papilla in male, dorsal-fin origin on vertical between base of 8th and 9th 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 1 + 7-8, 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
of savanna-like region.
Rivulus litteratus Costa, 2005
Rivulus sp.: Lacerda, 1989: 25 (Ponte Branca, Rio
Araguaia, central Brazil).
aqua vol. 11 no. 4 - 2006
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, 17º33’38.5”S 53º18’33.1”W,
altitude 750 m, Estado de Mato Grosso, Brazil;
holotype: UFRJ 5956).
of first branchial arch 1 + 7-8, 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).
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. 2004.
Diagnosis
origin on vertical between base of 7th 8th 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
Habitat notes
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.
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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.
164
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
165
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
aqua vol. 11 no. 4 - 2006
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 BeniMamoré 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 ichthyoloaqua vol. 11 no. 4 - 2006
gists 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
166
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 50-150
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
167
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 02001.001660/98 and
02.022005956/02 from IBAMA (Instituto Brasileiro
do Meio Ambiente e dos Recursos Naturais Renováveis – Ministério do Meio Ambiente, dos Recursos Hídricos e da Amazônia Legal).
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Rivulus immaculatus, a new killifish from Venezuela
(Cyprinodontiformes, Rivulidae). Copeia 1991: 323-328.
THOMERSON, J. E., NICO, L. G. & TAPHORN, D. C. 1992.
Rivulus tecminae, a new killifish from Amazonas Territory,
Venezuela (Cyprinodontiformes: Rivulidae). Ichthyological
Exploration of Freshwaters 2: 289-296.
THOMERSON, J. E. & TAPHORN, D. C. 1992. Two new
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(Cyprinodontiformes: Rivulidae). Ichthyological Exploration
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WALLACE, A. R. 2002. Peixes do Rio Negro/Fishes of the Rio
Negro. EDUSP, São Paulo, 517 pp.
WEITZMAN, S. H. & WOURMS, J. P. 1967. South American
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VERMEULEN, F. B. M. & HRBEK, T. 2005 Kryptolebias sepia n.
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170
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 293487, 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
219778, 10P; Costa Rica: Guanacaste: Tilaran. R. geayi:
MZUSP uncatalogued Brazil: Amapa: Serra do Navio. R. har171
aldsiolii: 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 308411, 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.
aqua vol. 11 no. 4 - 2006
[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.
aqua vol. 11 no. 4 - 2006
[24] Preopercle (Costa 1990) (CI: 1.00; RI: 1.00): (0) robust,
L-shaped, with a well-developed anteromedian rim; (1) thin, Cshaped, 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) 50-70 %;
(1) 20-40 %; (2) 10-15 %.
[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
172
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) 36-38.
[49] Number of caudal-fin rays (modified from Costa 1990)
(CI: 0.18; RI: 0.59): (0) 26-31; (1) 32-36; (2) 24-25 [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) 25-26.
[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) 16-17.
[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.
173
[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) 17.0–23.8 % SL; (1) 24.1–31.1 % 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) 29.0-41.5; (1) 42.0-49.0; (2) 52.5-81.0.
[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
10–30 % of fin; (1) approximately on 40 % of fin; (2) approximately on 50-80 % 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.
aqua vol. 11 no. 4 - 2006
[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 poraqua vol. 11 no. 4 - 2006
tion 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.
174
A. panchax
S. guignardi
K. brasiliensis
P. auriferus
A. boticarioi
G. hoignei
M. xiphophora
M. apurinan
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
R. uatuman
R. urophthalmus
R. violaceus
R. zygonectes
T. rubromarginatus
0000000000
1000000001
2000100011
2000101011
2100101011
2000100110
2000100011
200?101110
2000111011
2100110011
2000101011
2100101011
2100101011
2001101011
2010100011
2000101011
2002100011
2000101011
2000101011
2000100011
2000101011
2002100011
2000101011
2000100011
2000101011
2002100011
2000101011
2001101011
2000101011
2012100011
2002100011
2000101011
2000101011
2002100011
2000101011
2002100011
2002100011
2012100011
2000101011
2002100011
2000101011
2012100011
2000101011
201?100011
2012100011
2000101011
2002100011
2002100011
2001111010
1–10
0000000000
0100000000
0111101000
1112101000
1111101100
1011001101
1112101101
1111101100
1111101100
1111101300
1112101201
1111101100
1111101100
1112101100
1112101100
1112111400
1112101300
1112011101
1112111100
1112111100
1112111100
1112101100
1112111400
1112001100
1112111110
1112101100
1112101400
1112101400
1112111110
1112101100
1112101300
1112111110
1112111400
1112101100
1112011101
1112101300
1112101100
1112101100
1112101110
1112101100
1112111100
1112101100
1112111100
1112101100
1112101100
1112111100
1112101100
1112101100
1111101400
11–20
0000000000
0000000000
100100?002
1001000001
1201000101
1101000101
1111000111
1201000101
1001000101
1101100101
1211000111
1101010101
1101010101
1111000101
1001000001
1001000102
1001100001
1001000001
1001000002
1001000101
1001000101
1001000001
1001000102
1001000001
1001000001
1001001?01
1001000102
1001000102
1001000001
1001000001
1001100001
1001000101
1001000101
1001100001
1001000001
1001100001
1001100001
1001000001
1001000?01
1001001?01
1001000102
1001000001
1001000101
1001000001
1001000001
1001000102
1001100001
1001100001
1001000101
21–30
0000000000
0000000200
0000010110
1110110110
1100120111
11001212?1
1100120111
1100120111
1100120111
1100120111
1100121111
1100120111
1100120111
1100120111
1100120201
1100120111
1101120211
1100120111
1100120111
1100120111
1100120111
1100120211
1100120111
1100120111
1100120111
1100120211
1100120111
1100120111
1100120111
1100120211
1100120211
1100120111
1100120111
1100120211
1100120111
1101120201
1100120211
1110120211
1100120111
1100120211
1100120111
1100120211
1100120111
1100120211
1110120211
1100120111
1100120211
1100120211
1100120111
31–40
0000000000
0010111011
0001110001
0100111011
1100211101
0100311201
0100111001
1100111101
0100111101
0100211001
0100111001
0100211001
0101211101
1100111011
0101111201
0101111111
0101111011
0101111021
0101111001
0101111021
0101111001
0111111001
0101111111
0100311001
0101111101
0111111001
0101111111
0101111211
0101?11101
0101111021
0101111011
0101111001
0101111101
0101111011
0101111021
0101111011
0101111011
0101111021
0101?11111
0111111001
0101111101
0101111021
0101111001
0101111021
0101111021
0101111101
0101111011
0101111011
0100111211
41–50
0100201000
0101200000
0001000010
0001001010
0001110111
1211110111
0001111111
0101110111
0001110110
0001110111
1111110111
0001110111
0101110110
0001110111
0001111110
0101110110
0001111110
0001111110
0001110110
0001111110
0001110110
0001111110
0001110110
0001110110
0001111110
0001111110
0101110110
0101110110
0001111110
0001111110
0001111110
0001111110
0001110110
0001111110
0001111110
0001111110
0001111110
0001111110
0001111110
0001111110
0001110110
0001111110
0001110110
0001111110
0001111110
0001110110
0001111110
0001111110
0101110111
51–60
0000000000
0101101000
0001111000
00011110?0
0001111110
1011111111
0001111110
0001111110
0001111010
0001111010
1011111011
0001111010
0001111000
0001111110
0001111000
0101111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0101111000
0101111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111000
0001111001
0001111000
0001111001
0001111000
0001111000
0001111000
0001111000
0001111001
61–70
0000000000
0000001000
0000000011
0000000011
0221111111
2221121013
0220013111
0221120011
0110000011
0110000013
1221121011
0110111013
0110001011
0220110013
0110002004
0000000011
0110000011
0000013014
0000000011
0000000011
0000000012
0000001013
0000000011
0000000011
0000000011
0110001011
0000000012
0000000012
0000000011
0000000004
0110000013
0000000011
0000000011
0110000013
0000013014
0110000011
0110000013
0020000004
0000000011
0110000013
0000000011
0000002004
0000000011
0010000004
0000000004
0000000011
0110000013
0110000013
0220000012
71-80
000000000?
0001000011
000200000?
000200000?
1002000113
101200000?
1002000010
1002000110
1102100010
100320010?
101200000?
120200000?
120200000?
1002000010
1003001110
1102001110
1003001012
100200100?
1102001010
1002001110
1102001010
1002001111
1102001110
1202100014
1002001014
1002001111
1102001110
1102001110
1002001014
1003011110
1003001014
1002001114
1102001110
100300100?
1002001013
1003001012
1003001012
1003011010
1002001014
1002001111
1002001110
1003001110
1102001010
1003001114
1003011110
1002001010
1003001012
1003001012
1102100010
81-90
0000000100
0000001000
1000010101
0000000100
0000100000
0000100000
0000121000
0000101000
1000100000
0000110000
?000100001
0000112000
0000110001
0000120000
0011000200
0000000000
0000001100
0310000200
0010000001
0010000101
0000000110
1010000100
0110000000
0000000100
0010000101
1010000100
0110000010
0100000010
0010000101
0010000200
0000001100
0010000101
0010000010
0000001100
0310000210
0000001100
0200001100
0010000200
0010000101
1010000100
0010000010
0011000200
00?0000110
0011000200
0010000200
0110000010
0000001100
0200001100
0000100000
91-100
0000000000
1101000000
0010000000
0000000100
1001001010
0000010000
1101000001
1001000000
1000001000
0000001000
0000002000
?100000010
0000000000
1001000001
1100000000
0000000001
0000100001
0000001000
1110000000
0000000000
1101000101
0010000001
1100000001
0000000000
1110010000
0010100001
1100000001
0000000001
1110010000
0000000000
0000100001
1110010000
0000000001
0000000000
0000100000
0000100001
0000000001
0000000000
1110010000
1010100001
0100000101
1100000000
1101000101
0000000000
0000000000
1100000101
1100100001
0000000001
1000000000
101-110
00?
00?
00?
00?
00?
00?
00?
00?
00?
00?
00?
00?
011
011
011
011
112
00?
011
011
011
00?
011
010
011
00?
011
00?
011
00?
112
012
011
112
00?
112
112
00?
011
00?
011
011
011
011
00?
011
112
112
00?
111-113
175
aqua vol. 11 no. 4 - 2006
aqua vol. 11 no. 4 - 2006
176
Guidelines for Authors
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Scientific names of genera, species, and subspecies
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and paratypes must be clearly identified, the institution
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acceptable as repositories for holotypes.
BLABER, S. J. M. 1980. 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 31:137-46.
DAY, J. H., BLABER, S. J. M., & WALLACE, J. H. 1981.
Estuarine fishes. In: Estuarine Ecology with Particular Reference to Southern Africa. (Ed. J.H. Day.):
197-221. A. A. Balkema, Rotterdam.
DIMMICH, W. W. 1988. Ultrastructure of North American cyprinid maxillary barbels. Copeia 1988 (1):
72-79.
TREWAVAS, E. 1983. Tilapiine Fishes of the Genera
Sarotherodon, Oreochromis and Danakilia.
British Museum (Natural History), London, 583 pp.
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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.

Rivulus amanapira Costa, 2004

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