Tropical Zoology 11: 139-148, 1998

Revision of the neotropical annual fish genus

Pituna Costa 1989 (Cyprinodontiformes Rivulidae)

WILSON J.E.M. C OSTA

Laboratório de Ictiologia Geral e Aplicada, Departamento de Zoologia, Universidade
Federal do Rio de Janeiro, Caixa Postal 68049, CEP 21944-970, Rio de Janeiro, Brasil
(E-mail: wcosta@acd.ufrj.br)

Received 8 February 1997, accepted 5 January 1998

The hypothesis that Pituna Costa 1989 is a member of the tribe Plesiolebiatini is
supported by the presence of a concave anterior border of the rostral cartilage, non-
overlapping autopalatine and entopterygoid, a concave anteromesial border of the first
epibranchial, and an elongate basihyal. Its sister group relationship with the clade
comprising the genera Maratecoara Costa 1995, Plesiokbias Costa 1989 and Stenolebias
Costa 1995 is corroborated by the presence of plesiomorphic features for character
states defining that clade: a distinct sharp ventral process on autopalatine, a short
posterior process of the quadrate, and a wide distal basihyal. Pitma is diagnosed by a
posteriorly placed dorsal fin, unique anal-fin colour pattern in males, a metallic blue
humeral blotch in males, and scattered black spots over pectoral fins in males. Pituna
poranga Costa 1989 is considered to be a synonym of P. compacta (Myers 1927), which
occurs in a vast central Brazilian region, in temporary pools of the Araguaia and
Tocantins river basins.
KEY WORDS: Pisces, systematics, phylogeny, Neotropica, rio Tocantins, rio Araguaia.

Introduction . . . . . ........... 140

Material and methods . . . . . . . . . . . 140
Pituna Costa 1989 : . . . . . . . . . . . . 140
Diagnosis and relationships . . . . . . . . . . . 140
Remarks . . . . . . . . . . . 141
Pituna compacta (Myers 1927)’ 1 . . . . . . . . . . . 141
Material examined . . . . . . . . . . . . . 142
Diagnosis . . . . . . . . . . . . . . . 142
Description . . . . . . . . . . . . . 143
Osteology . . . . . . . . . . . . . . 144
Colouration . . . . . . . . . . . . . 146
Distribution . . . . . . . . . . . . . 146
Habitat . . . . . . . . . . . . . . . 146
Remarks . . . . . . . . . . . . . . . 147
Acknowledgements . . . . . . . . . . . . . . 148
References . . . . . . . . . . . . . . . 148
140 W.J.E.M. Costa

INTRODUCTION

The genus Pituna Costa 1989 was first described to include P. porunga Costa
1989 from the rio Araguaia basin, central Brazil (COSTA 1989), and then hypothesized
to be the closely related to Pterolebias Garman 1895 (COSTA 1989, 1990). Subsequent-
ly, two species previously placed in the genus Rivulus Poey 1860 were transferred to
Pituna (COSTA 1991). The first one, R. compactus Myers 1927, from the rio Tocantins
basin, central Brazil, was then known only from three females of the type-series, and
was placed in Pituna due to the high similarity in general morphology of these
specimens with females of P. poranga. The second species, R. stellifer Thomerson &
Turner 1973, from the Rio Orinoco basin, Venezuela, was transferred to Pituna due
to an apomorphic colour pattern of the male anal fin and a similar morphology of the
premaxilla. However, inclusion of R. stellifer in Pituna is not corroborated in the
present study.
The purpose of the present study is to clarify the taxonomic status and diagnostic
features of the species included in Pituna, based on large new collections along rios
Araguaia, Tocantins and das Mortes floodplains.

MATERIAL AND METHODS

Measurements and counts follow COSTA (1988). Measurements are presented as percentages of
standard length (SL), except for subunits of the head, which are presented as percentages of head
length. Osteological studies were made on cleared and counterstained specimens (C&S) prepared
according to TAYLOR & VAN DYKE (1985). The compound caudal centrum was counted as a single
element in vertebrae numbers, which were taken only from cleared and stained material. Osteologi-
cal descriptions focus on traits of some importance for phylogenetic analyses of rivulid fishes (e.g.,
COSTA 1990), and illustrations show traits useful for diagnosis of the genus. Nomenclature for
frontal squamation follows HOEDEMAN (1956), and that for cephalic neuromasts follows GOSLINE’S
(1949) nomenclature for head sensory canals. Supraorbital neuromast counts were made in a
longitudinal row from the neuromast closest to the anterior naris to the neuromast nearest the orbit,
not including the transverse neuromasts over rostral region nor any posterior extension of this
series.
Abbreviations for institutions are: CAS, California Academy of Sciences, San Francisco;
MNHN, Museum National d’Histoire Naturelle, Paris; MNRJ, Museu Nacional do Rio de Janeiro,
Rio de Janeiro; MZUSP, Museu de Zoologia, Universidade de São Paulo, São Paulo; UFRJ,
Universidade Federal do Rio de Janeiro, Rio de Janeiro; and, UMMZ, University of Michigan
Museum of Zoology, Ann Arbor.

Pituna Costa 1989

Pituna COSTA 1989: 225 (type species Pituna poranga Costa 1989, by original designation).

Diagnosis and relationships

Pituna is similar to other genera of the Plesiolebiatini (sensu COSTA 1995a)
(Stenolebias Costa 1995, Maratecoara Costa 1995 and Plesiolebias Costa 1989), but
differs from other genera of the Rivulidae by the possession of a concave anterior
border of the rostral cartilage (vs convex to straight); non-overlapping autopalatine
and entopterygoid (vs overlapped); a concave anteromesial border of first epibranchial
Revision of Pituna 141

(vs straight); and, elongate basihyal, its longitudinal length about 54% of the length
between anterior border of basihyal and posterior ossified edge of third basibranchial
(vs about 46-52%); another synapomorphic condition shared by Pituna and other
plesiolebiatins, is not present in all members of the clade: presence of alternate black
and white or yellow spots on basal and posterior part of the anal fin of males (vs
absent; reversed in Maratecoara). Pituna is similar to other genera of the Plesiolebiatini
and all genera of Cynolebiatini (Spectrolebias Costa & Nielsen 1997, Simpsonichthys
Carvalho 1957, Cynolebias Steindachner 1876, Leptolebias Myers 1952, Cynopoecilus
Regan 1912, and Campellolebias Vaz-Ferreira & Sierra 1974) and distinguished from
other genera of the Rivulidae by a reduced ventral process of the angulo-articular (vs
not reduced) and absence of the lateral processes on the neural spine of the first
vertebra absent (vs present). Pituna is distinguished from all other members of the
Plesiolebiatini by the following autapomorphic features: dorsal fin placed posteriorly,
second proximal radial of the dorsal fin between neural spines of vertebrae 17 and 19
(vs 12-16); unique colour pattern on the anal fin in males, consisting of a brownish red
stripe on the the distal margin, bordered dorsally by a narrow pale green stripe; a
metallic blue humeral blotch in males (vs without humeral blotches; independently
acquired in Plesiolebias bitteri Costa 1989); and, scattered black spots on the pectoral
fins in males (vs no dark marks on pectoral fin). Pituna is further distinguished from
other plesiolebiatins by some plesiomorphic features: a distinct sharp ventral process
of autopalatine (vs ventral process represented by a short expansion); a short posterior
process of the quadrate, about 45% of the total longitudinal length of the quadrate
(vs about 55-60%); and, a wide distal portion of basihyal (vs narrow).

Remarks
Pituna was originally diagnosed by a concavity on the anterolateral margin of
ascending process of the premaxilla (COSTA 1989). According to the present study,
this character state is variable both in Pituna and in other rivulid genera, therefore,
not valid to define the genus. COSTA (1989) hypothesized a sister group relationship
between Pituna and Pterolebias supported by the presence of widened proximal radials
of anal fin and a reduced interarcual cartilage. Based only on this putative sister group
relationship, LAZARA (1990) placed Pituna in the synonymy of Pterolebias. However,
under a detailed comparison among all rivulid lineages, these conditions are morpholo-
gically distinct, herein interpreted as non-homologous. On the other hand, Pituna is
hypothesized to be the sister group to a clade comprising Maratecoara, Plesiolebias and
Stenolebias, supported by the synapomorphies described in the above diagnosis.
Despite several osteological features that do not support a close relationship
between Pituna and the widespread New World genus Rivulus (e.g. COSTA 1990),
HUBER (1992) considered the former to be a subgenus of the latter. The data
presented above indicate that Pituna is a member of the tribe Plesiolebiatini, which is
closely related to the tribe Cynolebiatini, whereas Rivulus is hypothesized to be
outside this assemblage.

Pituna compacta (Myers 1927) (Fig. 1)

Rivulus compactus MYERS 1927: 120 (original description, shallow lake, Donna Franciquinha [Dona
Francisquinha], Porto Nacional, Rio Tocantins, Goyas [presently Estado do Tocantinsl, Brazil).
142 W.J.E.M. Costa

Pituna poranga COSTA 1989: 226 (original description, Aruanã, Goiás, Brazil), n. syn.
Pterolebias poranga; LAZARA 1990: 176.
Pituna compacta; COSTA 1991: 332.
Rivulus poranga; HUBER 1992: 367.

Material examined
A total of 186 specimens, 14 C&S, listed below (all localities are in central Brazil).
Lectotype (designated by HUBER 1992): CAS 40707, female, 25.5 mm SL; Estado do
Tocantins, shallow lake, Dona Francisquinha, Porto Nacional rio Tocantins basin; C. Ternetz,
16X.1924.
Holotype of P. poranga MZUSP 38511, male, 28.1 mm SL; Estado de Goiás, Aruanã; S.V.
Silva, 1986.
Paratypes of P. poranga: MZUSP 38510, a male (C&S), about 30 mm SL; MZUSP 38509, a
female, 29.6 mm SL (C&S); MNRJ 11388, a female, 29.0 mm SL; all collected with holotype.
Other specimens (non-types): Estado do Pará: UFRJ 3934, 9 ex.; UFRJ 3935, 3 ex. (C&S);
pool 50 m from rio Araguaia, Vila de Santa Cruz, São Geraldo do Araguaia. Estado do Tocantins:
UFRJ 2099, 29 ex.; UFRJ 2115, 2 ex. (C&S); MNHN uncat., 6 ex.; temporary lagoon close to rio
Tocantins, Brejinho de Nazarè; W.J.E.M. Costa, G.C. Brasil & C. Campinha, 15-16.11.1994. UFRJ
2101, 11 ex.; temporary lagoon close to rio Dona Francisquinha, Porto Nacional, rio Tocantins
basin; W.J.E.M. Costa, G.C. Brasil & C. Campinha, 16.11.1994. UFRJ 3563, 27 ex.; UFRJ 3564, 4
ex. (C&S); UMMZ uncat., 6 ex.; temporary lagoon close to Barreira do Piqui, rio Araguaia basin;
W.J.E.M. Costa, G.C. Brasil, M.I.P.F. Landim & C.L.R. Moreira, 15.11.1996. MZUSP 45222, 28
ex.; rio Agua Fria, Praia Alta 2 Farm, road Araguaçu-Barreira do Piqui, 27 km N from Araguaçu, rio
Araguaia basin; F.C.T. Lima. UFRJ 3797, 2 ex.; 10 km from Salvação Alvorada, rio Tocantins
basin; D.T.B. Nielsen, A. Carletto & A. Lucca, 05.IV.1996. Estado de Goiás: UFRJ 3540, 13 ex.;
temporary pool 4 km E of rio Araguaia, road to Peixe; W.J.E.M. Costa, G.C. Brasil, M.I.P.F.
Landim & C.L.R. Moreira, 16.II.1996. Estado de Mato Grosso: UFRJ 3543, 16 ex.; UFRJ 3545, 3
ex. (C&S); temporary pool 15 km W of rio das Mortes, road to Cocalinho; W.J.E.M. Costa, G.C.
Brasil, M.I.P.F. Landim & C.L.R. Moreira 16X.1996. UFRJ 247, 21 ex.; UFRJ 294, 1 ex. (C&S);
temporary pool in the road between Agua Boa and Cocalinho, 16 km W of rio das Mortes; G.C.
Brasil, M.T.C. Lacerda & P. Araújo, I. 1988.

Diagnosis
As for the genus.

Fig. 1. - Pituna compacta, male, specimen collected near Aruanã rio Araguaia basin, about 35 mm SL,
not preserved.
Revision of Pituna 143

Description
Morphometric data are presented in Table 1. Maximum adult size recorded in
the present study 37.5 mm SL for males, 29.2 mm SL for females. Dorsal profile of
body approximately straight on head, convex from nape to end of dorsal-fin base,
gently concave on caudal peduncle. Ventral profile weakly convex on head, nearly
straight between head and base of last anal-fin ray, slightly concave on caudal
peduncle. Anterior profile of snout rounded.
Dorsal and anal fins pointed, sometimes with short filamentous rays on tip of
male fins, reaching a line drawn vertically through the middle of the caudal fin.
Dorsal-fin origin at a point on a vertical line drawn between the bases of anal-fin rays
9-11. Caudal fin rounded. Pectoral fin elliptical, its posterior margin reaching a point
on a vertical drawn between the pelvic-fin base and the urogenital papilla. Pelvic-fin
bases almost continuous, separated by small interspace. Tip of pelvic fin reaching
between urogenital papilla and second anal-fin ray. Dorsal-fin rays 8-10 (ii-iii, 5-7,
O-i); anal-fin rays 13-16 (ii-iii, 10-14, O-2); caudal-fin rays 25-30 (v-vii, 14-17,
v-viii); pectoral-fin rays 14-16 (i, 11-14, i-iii); pelvic-fin rays 7-8 (i, 6-7, i).
Frontal squamation usually F-patterned, sometimes E-patterned; E-scales not
overlapped; frontal scales arranged circularly (Fig. 2). Supraorbital series of neuro-
masts 5-8 + 3-4; vertical section of preopercular series of neuromasts with two well-
developed neuromasts followed ventrally by two or three smaller neuromasts. Two
neuromasts arranged transversely on caudal-fin base. Longitudinal series of scales
28-30; transverse series of scales 8; scale rows around caudal peduncle 16. Caudal-fin
squamation on about 40% of the length between posterior border of hypural plate
and posterior edge of the fin.
Swimbladder short, posterior tip not reaching a vertical through anal-fin origin.
Urogenital papilla of males cylindrical, very short and not close to anal fin; urogenital
papilla of females flattened, pouch-like, not close to anal fin. Opercular membrane
short, not extending on pectoral-fin base.
Table 1.
Morphometric data of Pituna compacta.

Males Females

UFRJ UFRJ UFRJ UFRJ UFRJ UFRJ UFRJ UFRJ UFRJ

247 3543 3543 3934 2089 3543 2089 3543 3543
SL [mm] 37.5 37.1 30.8 29.4 27.1 29.2 26.3 26.0 23.4
As percentages of standard length
Body depth 25.8 24.1 25.2 24.0 25.8 24.3 23.5 23.7 24.7
Depth of caudal peduncle 17.9 18.7 18.1 17.9 17.2 16.6 15.9 15.9 17.1
Predorsal length 76.6 75.2 78.1 78.3 79.0 78.5 76.1 77.9 77.5
Prepelvic length 53.5 50.8 51.6 53.2 53.9 54.8 52.2 53.7 54.8
Length of dorsal-fin base 10.6 11.7 10.1 11.0 10.0 10.1 11.2 9.7 10.4
Length of anal-fin base 23.6 23.0 21.4 22.8 23.4 21.5 22.9 21.0 19.9
Head length 30.8 29.5 30.3 30.1 32.6 30.4 30.7 30.5 30.7
As percentages of head length
Head depth 75.0 75.6 71.8 69.4 74.1 70.3 66.8 68.6 68.9
Head width 74.4 75.3 74.1 77.7 74.5 76.2 72.3 75.9 74.1
Eve diameter 27.2 30.3 34.1 32.0 30.7 34.3 30.0 33.7 33.9
144 W.J.E.M. Costa

Fig. 2. - (Left) Diagrammatic representation of dorsal view of head in Pituna compacta showing frontal
squamation and neuromast patterns.

Fig. 3. - (Right) Lateral view of left jaw suspensorium of Pituna compacta, UFRJ 3564. Abbreviations: a,
autopalatine; e, entopterygoid; h, hyomandibula; mt, metapterygoid; p, preopercle; q, quadrate; S,
sympletic. Dots indicate bone, circles cartilage.

Osteology
Neurocranium and infraorbital series: posterior rim of autopterotic rounded.
Dorsal process of supraoccipital short, slender, directed posteriorly. Posterior process
of vomer narrow, not elongate. One or two vomerian teeth. Lacrimal almost cylindri-
cal and twisted. Dermosphenotic reduced.
Suspensorium and opercular apparatus (Fig. 3): autopalatine not contacting
quadrate; autopalatine and entopterygoid not overlapped. Ventral portion of autopa-
latine with a distinct sharp process. Entopterygoid moderate, about as long as
autopalatine, posterior tip not contacting metapterygoid. Posterior process of qua-
drate about 45% of the total longitudinal length of quadrate. Sympletic somewhat
slender. Preopercle rounded, sensory canal absent.
Jaws (Fig. 4): Anterior border of distal portion of alveolar arm of premaxilla
slightly projected anteriorly. Ascending process of premaxilla narrow and somewhat
elongate. Rostral cartilage narrow and moderate in length, smaller than premaxillary
ascending process; anterior border concave. Ventral process of angulo-articular re-
duced. Retro-articular short.
Hyal and branchial arches (Figs 5 and 6): basihyal almost triangular, elongate, its
longitudinal length about 54% of the length between anterior border of basihyal and
posterior ossified edge of third basibranchial. Basihyal cartilage moderate, its longitu-
dinal length about 40% of the total length of basihyal. Lateral border of basihyal
cartilage approximately straight. Six branchiostegal rays. Interhyal cartilaginous.
Urohyal moderately deep. Anterior border of first epibranchial concave. Interarcual
cartilage well-developed. Teeth on fourth ceratobranchial. Gill-rakers on first gill arch
2 + 9.
Pectoral girdle: ventral process of posttemporal short. Supracleithrum not elon-
gate, clearly shorter than posttemporal. Pectoral radials cubical and well-developed.
Revision of Pituna 145

Fig. 4. - (Left) Jaws of Pituna compacta, UFRJ 3564. A, dorsal view of left upper jaw; B, lateroventral
view of left lower jaw. Abbreviations: a, angulo-articular; d, dentary; m, maxilla; p, premaxilla; r, retro-
articular; rc, rostral cartilage. Dots indicate bone, circles cartilage.

Fig. 5. - (Right) Dorsal view of median and left portion of ventral gill arches and basihyal of Pituna
compacta, UFRJ 3564 (ceratobranchials not represented). Abbreviations: b, basibranchials l-3; bh,
basihyal; h, hypobranchials 1-3. Dots indicate bone, circles cartilage.

Fig. 6. - Ventral view of left dorsal gill arch of

Pituna compacta, UFRJ 3564. Abbreviations: e,
epibranchials l-4; i, interarcual cartilage; p, pha-
ryngobranchials 2-3; t, tooth plate pharyngobran-
chial 4. Dots indicate bone, circles cartilage.

Vertebrae, unpaired fin position and supports: no anterolateral processes on first

vertebrae. Neural prexygapophyses of caudal vertebrae reduced. Total number of
vertebrae 28-29, 12-14 precaudal and 15-16 caudal. Second proximal radial of dorsal
fin between neural spines of vertebrae 17 and 19. Second anal proximal radial just
posterior to pleural rib of vertebra 12. Anterior proximal radials of anal fin widened,
146 W.J.E.M. Costa

very close among themselves, forming a structure similar to a plate. Hypural elements
composing two plates.

Colouration
Males. Sides of body dark brown with oblique rows of small greenish yellow
spots, usually arranged in chevron-pattern with tip directed posteriorly; large, metallic
dark blue blotch on humeral region. Body sides of aquarium specimens sometimes
dark purple, a colouration never observed in wild specimens. Body sides of juveniles
pale brown with irregular oblique rows of dark brown spots. Anterior portion of head
sides and dorsum brown with melanophores concentrated on jaws and forming narrow
transverse stripe posterior to mouth cleft, and forming a spot on chin. Transverse
black bar on suborbital region. Opercular region metallic green, scale borders dark
brown. Iris brownish yellow; dark brown bar crossing eye. Dorsal fin dark brown,
with a row of yellow spots along fin base and posterior margin; a bluish green blotch
on anterior margin; sometimes red pigmentation on fin tip. Basal half of anal fin black
with a row of yellow spots along fin base and posterior margin; usually a brownish red
stripe on distal margin, bordered dorsally by narrow pale green stripe; sometimes
distal half dark brown, without longitudinal stripes. Caudal fin dark brown. Pectoral
fin pale yellowish white, with four or five transverse rows of black spots. Pelvic fin
brown, anterior and posterior borders pale yellow, tip reddish brown. Colour photos
are presented in COSTA (1992) and HUBER (1992).
Females. Sides of body light gray, with dark brown spots, variably arranged,
usually in irregular oblique, chevron-patterned rows, sometimes arranged longitudin-
ally forming three stripes. Sides of head light gray, with dark chromatophores
concentrated on jaws, small dark spots on opercular region, a small spot on chin, and a
black suborbital transverse bar. Iris yellow, a dark brown bar crossing the eye.
Unpaired fins hyaline, with faint rows of dark brown spots transverse to fin rays.
Paired fins hyaline.

Distribution
Araguaia and Tocantins river
Araguaia, near the confluence of
Araguaia floodplains (about 15° S),
das Mortes floodplains (about 14°
(about 13° S) (Fig. 7).
basins, herein recorded from São Geraldo do
both rivers (about 6 ° S) South to Aruanã, in
swamp between Agua Boa and Cocalinho, in rio
S) and Alvorada, in the upper Tocantins area

Habitat
Pituna compacta occurs in temporary lagoons of the “Cerrado” area (Savannah) of
central Brazil. Sometimes it is the only annual fish found in those lagoons, but usually
it is sympatric with annual fishes of the genera Trigonectes Myers 1925, Simpson-
ichthys, Plesiolebias, Maratecoara and Spectrolebias. However, in contrast with all
sympatric annual fishes, P. compacta is only found close to marginal areas, in the
shallowest parts (about 5 cm deep) of the temporary lagoons. This habitat preference
is probably related to its distinctive cylindrical and slender body, that is similar to the
body shape of species of the genus Rivulus, which also live in shallow biotopes (COSTA
1995b).
Revision of Pituna

Fig. 7. - Geographic distribution of Pituna compacta. Arrow indicates the type-locality.

Remarks
No differences in morphometric and meristic data, and osteology were found
among the populations of the Araguaia and Tocantins river basins examined. Colour
pattern of males is very similar in all populations, but the female colour pattern is
rather variable among different populations, diverging even in very close geographic
areas of the same basin, thus considered non-informative for recognizing distinct
taxa. Therefore, P. poranga is considered as a synonym of P. compacta.
Rivulus stellifer does not present either of the characters herein used to define
Pituna, nor the synapomorphies described for Plesiolebiatini or a clade comprising
Plesiolebiatini plus Cynoleboatini. Hence, inclusion of this species in Pituna is not
supported. On the other hand, the presence of a scaled caudal fin, an elongate
anterior portion of the urohyal and the apomorphic suborbital colour pattern suggest
that R. stellifer is closely related to the annual fish genera Austrofundulus Myers 1932
and Rachouia Myers 1927.
148 W.J.E.M. Costa

ACKNOWLEDGEMENTS

I am grateful to G.C. Brasil, H.A. Britski, C. Campinha, J.L. Figueiredo, M.T.C. Lacerda,
M.I. Landim, N.A. Menezes, C.L.R. Moreira, D. Nielsen, and 0. Oyakawa for the gift or loan of
material, help during fieldwork, and hospitality during visits to their institutions. The manuscript
benefited from the comments of M.J. Ghedotti. This study was supported by CNPq (Conselho
Nacional de Desenvolvimento Cientffico e Tecnológico - Brazilian Federal Government) and
FAPERJ (Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro).

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