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ABOUT FOCUS
DE Serrasalmus SPILOPLEURA Kner, 1858 AND
REHABILITATION S.! Maculatus Kner, 1858
(CHARACIDAE!: SERRASALMINAE)
by
JEGU Michel (1) & Geraldo Mendes dos Santos (2)
SUMMARY.! -! Serrasalmus spilopleura Kner, 1858 and S.! Maculatus Kner, 1858 from the basin
Guaporé are described. The authors, for the most part still considered S.! Maculatus as a synonym
S.! spilopleura, a species known basins of the Paraná-Paraguay and Amazon. Examination of
standard series of the collection of Vienna showed that both species are in fact different. In
S.! Spilopleura the bones of the infraorbital series are narrower and the bare area of the cheek is larger than
in S.! maculatus. The 2 / 3 at the base of the tail are dark and the last third hyaline, with
S.! Spilopleura, while the tail of S.! Maculatus has a terminal or subterminal black bar,
depending on the size of specimens and collection area. The specimens identified S.! Spilopleura by
recent authors are S.! maculatus. Specimens of Guaporé, with the same morphotype that
S.! Spilopleura, differentiated by the presence of a gray to black distal band on the tail.
ABSTRACT.! -! Review of Serrasalmus spilopleura Kner, 1858 and restoration of S.! Maculatus Kner,
1858.
Serrasalmus spilopleura Kner, 1858 and S.! Maculatus Kner, 1858 are Described from Guaporé
Basin. Most authors Have Always Considered S.! Maculatus as a synonym of S.! Spilopleura, a Well-Known
species from Paraná-Paraguay and Amazon basins. The examination of the type-series in Vienna shows
S. That! and S. maculatus! spilopleura are in Fact Two different species. In S.! Spilopleura the infraorbital
series bones are narrower and the naked cheek area IS in S. Broader Than! maculatus. Two Thirds at the base
caudal end of the last are dark and hyaline in S. Third! Whereas S. spilopleura! maculatus shows a final gold
subterminal black bar in the caudal end, DEPENDING on the size of the specimens and the sampling area.
S.! Spilopleura specimens Identified by recent authors Have To Be Placed Under S.! Maculatus. The specimens
from Guaporé basin show a very close to morphotype of S. That! spilopleura goal THEY Differ from this
species by a gray to black terminal band on the caudal end.
Key words.! -! Serrasalmus spilopleura - S.! Maculatus - Brazil - Gold Itenez Guaporé River - Type-specimen
revision - Distribution.
Serrasalmus maculatus and S.! Spilopleura have been described by Kner Rio Guaporé
(1858!: 166), from material brought to Vienna by Natterer and presented with more
detail in the work of Kner (1860) on the Characidae.
Kner (1858) indicates that the edge of the anal and caudal fins of S.! Maculatus
black and the sides are patches while in S.! spilopleura the edge of the caudal
clear and the sides are a black humeral. Kner (1860) indicates that according to Natterer,
(1)! Antenna IRD, Laboratory of General and Applied Ichthyology, MNHN, 43 rue Cuvier, 75231 Paris
Cedex 05, FRANCE. [Jegu@mnhn.fr]
(2)! Laboratorio de Ictiologia, INPA, CPBA, Cx P. 478, 69083-970 Manaus, A.M., BRAZIL
Cybium 2001, 25(2): 119-143.
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JEGU & DOS SANTOS
that collected the fish, common names are "! pequena Piranha!" for S.! maculatus
and "! Piranha doce" for S.! spilopleura that Heckel (in Kner, 1860) called Piranha dulcis.
Kner, 1858 does not give value for meristic and morphometric characters
of Serralmus spilopleura, unlike S.! maculatus (Tableau! I). The author reports no
of S.! spilopleura examined does a full set of teeth ectoptérygoïdiennes,
when he observed 5-6 teeth on the palate in S.! maculatus alcohol and at least 3 of the
dry specimens. Kner (1858) also indicates that the infraorbital series is notoriously more
close in S.! spilopleura than in S.! maculatus, the area under the naked infraorbital 3 is more
large in S.! spilopleura. The representation of S.! Spilopleura (Kner, 1860!: Pl.! 5, Fig.! 11),
we found about 17 rays in the dorsal and anal with 34. Finally, the author reports the
presence of black marks on the sides of the two species and the presence of a band
distal caudal black in S.! maculatus, while the caudal edge of the hyaline in
S.! Spilopleura. Black dots the size of the pupil are clearly visible on the representation
S.! maculatus (Kner, 1960!:! pl.! 4, Fig.! 10), but only a humeral spot is darker
visible on the sides of S.! spilopleura (Kner, 1960!: pl.! 5, Fig.! 11).
For either species, it is reported the number of specimens examined,
but the author indicates that specimens of S.! spilopleura come from Bogota, Rio Guaporé
in Mato Grosso. For S.! Maculatus, which also comes from the Guaporé, Kner (1858) states
Some specimens are from dry Caiçara.
Günther (1864!: 370) cites S.! Spilopleura Capim River in Brazil and resumes
description of Kner (1858) for S.! maculatus. Eigenmann (1915!:! 249) distinguishes
S.! Spilopleura with a black tail band and submarginal hyaline distal edge of
S.! Maculatus whose tail has a black margin. Eigenmann (1915!: 253)
S. said that! spilopleura is known to the Amazon and the Paraguay River Basin, where it is cited
for the first time by Boulenger (1896), while S.! maculatus would be limited to Bolivia
and the Amazon basin. Norman (1929!: Fig.! 13) represents S.! Spilopleura with a band
submarginal caudal neat and confirmed its presence in the basins of the Paraguay and
Amazon!, he (p.! 799) suggests that S.! be synonymous with S. maculatus! spilopleura. The
Table! I.! -! Features Serrasalmus maculatus and S.! Spilopleura by Kner (1860). [Serrasalmus
maculatus and S.! spilopleura characters from Kner (1860).]
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Serrasalmus spilopleura and S. maculatus
last quote of S.! maculatus is presented by La Monte (1935) for the Rio Juruá. By
Later, Fowler (1950!: 384) and Géry (1976) consider S.! maculatus as a synonym for
S.! Spilopleura.
Géry (1964) emphasizes that it is often difficult to observe the black subterminal
the tail in S.! spilopleura the region of Iquitos in Peru. According to this author (1978!: 286),
S.! Spilopleura is characterized by the presence of teeth on the palate and a band distal
hyaline at the tail. However, this species shows two specimens with the color purple
uniform (p.! 292). Santos et al. (1984!: 35), then Jégu and Santos (1988!: Pl.! 11), represent
S.! Spilopleura of Tocantins with the characteristic subterminal black band on the caudal
black spots on the sides of a humeral spot and young adults. These
latter authors report (p.! 256) that the bare area of the cheek appears wider and the eyes
smaller on the representation of S.! spilopleura of Kner (1860), but they follow Norman
(1929) and S. place! Maculatus, Tocantins cited by Ulrey (1895), as a synonym for
S.! Spilopleura. Lauzanne and Loubens (1985!: 53, fig.! 48) identify S.! Spilopleura Basin
Mamore and Itenez (Bolivia) on the same criteria for coloring. After reviewing the
bibliography on S.! spilopleura Paraná-Paraguay basin, Gery et al. (1987!: Fig.! 58,
59) represent S.! Spilopleura of Paraguay with a black subterminal tail bar and
black dots the size of the pupil on the side of a youthful 77! mm SL. Santos (1990!:
176, fig.! 178) described S.! Spilopleura Guaporé as a kind of body dark unmarked
clearly defined, without the caudal edge hyaline but with teeth ectoptérygoïdiennes.
The author (p.! 175-177) indicates the presence of two other species without teeth ectoptérygoïdiennes
in Rondonia. In one (Serrasalmus sp.! 1, fig.! 186), and cited the Guaporé
Jamari, a right tributary of the Madeira, the edge of the tail is black and in the other (S.! Aureus,
Fig.! 183), cited only the Jamari, the edge of the caudal hyaline. Castro (1994!: 64) for
Putumayo (Colombia), and Ferreira et al. (1998!: 82), to the Amazon area
Santarem (Brazil), are S.! Spilopleura with a black subterminal tail band.
Fernandez Yepez (1969) and Mago Leccia (1970) cite S.! Spilopleura for Venezuela
But Machado Allison and Fink (1996) do not retain this species in their catalog
piranhas from Venezuela. Finally, Jégu and Keith (1999a) reported the presence of
S.! Spilopleura in the swamps of the right bank of the Oyapock by an adult form the body
uniformly purple and dark, the criterion on which the color of the tail is no longer
visible. These authors therefore agree on the presence, in young S.! Spilopleura basins
the Amazon and Paraguay, with a subterminal black band on the tail and spots
black on a light background on the sides. The dress of adults would be darker, sometimes purplish and
coloration of the caudal more difficult to observe (Jégu and Santos, 1988!, Azuma, 1990).
Jégu and Keith (1999b) placed S.! Spilopleura in the Guild of Serrasalminae Varzea
in the Amazon basin, with an eastward extension to the south to the upper Tocantins
and north in the coastal marshes of Amapá (Brazil) to the Oyapock.
STUDY OF A KIND
In August 1999 we examined the collection of the Museum of Serrasalminae
Vienna and the various registers of entries. The measurements and counts on specimens
examined were carried out following the methodology proposed by Merckx et al. (2000).
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Fig., 1.! -! Side view of lectotypes. A!: Serrasalmus spilopleura Kner, 1858!: NMW 57085, 180! Mm SL!;
B!: Serrasalmus maculatus Kner, 1858!: NMW 17995, 162! Mm SL. [Lateral view of lectotype specimens.
A: Serrasalmus spilopleura Kner, 1858: NMW 57085, 180! Mm SL, B: Serrasalmus maculatus Kner, 1858:
NMW 17995, 162! Mm SL.]
Type series of Serrasalmus spilopleura Kner, 1858
We found five specimens brought to the series-type spilopleura Serrasalmus,
which one kept in alcohol (NMW 57085, 180! mm SL) and four dry (NMW 79457-9 and
NMW 16344, 145 to 210! Mm SL). The material preserved in alcohol is well preserved (Fig.! 1a),
but the dried specimens appear to have been deformed at the time of preparation (Fig.! 2a-d).
The three specimens NMW 79457-9 are several labels on the sides with the
name given to them by Natterer [Salmo (Piranha doce)], derived from the common name, and
one has assigned Kner [Serrasalmo spilopleura Kner (Heck. Pygoc. dulcis)] (Fig.! 2b).
The morphometric characters of these specimens are homogeneous (Tableau! II). all
these specimens are elongated body, the snub nose and short head. The head and interorbital space
are wide. The third infraorbital leave a bare area on the cheek with a width
ranges from 1.9 to 2.5 times in that of bone. The mouth is short and wide. we
observed in ectoptérygoïde teeth in only one specimen that has a tooth on one side
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Serrasalmus spilopleura and S. maculatus
palate and the other two. The teeth of the premaxilla and tooth are low and wide and
median is not cusped tooth (Fig.! 3a).
The meristic characters of these specimens (Tableau! III) are also homogeneous. the
number of rays varies from 31 to 35 in the anal, which from 28 to 32 branched rays, and from 15 to 16 in
backbone, which branched from 13 to 14. The number of ventral serrae varies from 32 to 37, from 23 to 27
serrae simple and always a thorn in front of the anal fin. Based on the shelves
the anal there are from 6 to 12 sets of scales. The number of scales in lateral line varies from 85 to
96.
Table! II.! -! Morphometric features, expressed in% of SL, specimens of series-type
Serrasalmus maculatus Kner, 1858 and Serrasalmus spilopleura Kner, 1858 (the underlined values correspond
with deformed characters). [Morphometric characters of Serrasalmus maculatus Kner, 1858 and
Serrasalmus spilopleura Kner, 1858-series type specimens, Expressed in percent of SL (Underlined values
corresponds to deformed features).]
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Fig.! 3.! -! Side view of teeth of the principal series of premaxillary and dental lectotypes.
A!:! Serrasalmus spilopleura NMW 57085!; B!: S.! Maculatus, NMW 17995. [Lateral view of the premaxillary
dentary teeth and hand series of lectotypes. A: Serrasalmus spilopleura NMW 57085;
B:! S.! Maculatus, NMW 17995.]
The rays of the caudal are shaded from the base up to 2 / 3 of their length when
the color is stronger. The last third of the rays hyaline. The caudal spot is therefore
not reduced to a black subdistale. All specimens have a brand humeral
triangular, more or less clear. While there are small dark spots on the flanks
and the center of the anal lobe appears slightly dimmed on the specimen preserved in alcohol,
no other brand color appears on the dry specimens.
These specimens fit the description of Kner (1860), except the number
ectoptérygoïdiennes teeth, alteration probably caused by handling.
The different labels correspond to those provided by Kner.
It is therefore of specimens of the type series of Serrasalmus spilopleura. We
propose to designate as lectotype of S.! spilopleura Kner, 1858 the specimen NMW
57085, 180! Mm SL, and paralectotypes of S.! Spilopleura, dry specimens NMW 16344,
79457-79459.
Type series of Serrasalmus maculatus Kner, 1858
Two specimens of Guaporé, collected by Natterer and preserved in alcohol (NMW
17995, 162! Mm LS and NMW 17996, 122! Mm SL), are identified as syntypes of
S.! Maculatus. A third specimen (NMW 57058, 109! Mm SL), collected by Natterer in
Barra do Rio Negro, was recently (February 10, 1978) designated syntype in the collection. The
town "! Barra do Rio Negro," is located at the mouth of the Rio Negro, downstream of Manaus. It
can not be a specimen of the type series for Kner (1860) clearly indicates
the specimens used in the description of S.! maculatus from the Rio Guaporé.
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JEGU & DOS SANTOS
In his work, Kner (1858!: 164) describes the Guaporé Mylett maculatus which it takes
also described in 1860 (p.! 26) and present a figure (pl.! 2, Fig.! 5). A sample
Table! III.! -! Meristic characters of specimens of series-Serrasalmus maculatus Kner types,
1858 and Serrasalmus spilopleura Kner, 1858. [Meristic characters of type specimens of Serrasalmus-series
maculatus Kner, 1858 and Serrasalmus spilopleura Kner, 1858.]
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Fig.! 4.! -! Serrasalmus maculatus, paralectotypes. A!: NMW 16230, 154! Mm SL!; B!: NMW 17996,
122.1! Mm SL. [Serrasalmus maculatus, paralectotypes. A: NMW 16230, 154! Mm SL, B: NMW 17996,
122.1! Mm SL.]
Mylett maculatus will be sent later to the Museum of London and incorporated into the collection under
number BMNH 1928.1.24!: 10. Eschmeyer (1998!: 994) indicates this specimen in error
as syntype of Serrasalmus maculatus, while it is probably a syntype of
Mr.! Maculatus now placed in the genus Metynnis.
Examination of the dry collection of individuals allowed us to find a specimen
under Pygocentrus melanurus from Heckel Caiçara (NMW 16230, 154! mm SL,
Caiçara, Rio Guaporé, collected by Natterer in? 1825). This is undoubtedly the specimen
Caiçara mentioned by Kner (1860!: 34), with 3 and 5 teeth and appointed ectoptérygoïdiennes
Pygocentrus melanurus by Heckel (in Kner, 1860).
We present a description of additional specimens NMW 17995
(Fig.! 1b), 17996 and 16230 (Fig.! 4a, 4b), the main meristic and morphometric characters
are given in Tables II and III.
The three individuals have homogeneous morphometric characters. The general form of
Serrasalmus spilopleura and S. maculatus 129
body is quite similar to that of S.! spilopleura, but these specimens are distinguished by
more characters. The body is a little higher (54.2 to 57.2% vs 49.3 to 53.6 for LS
S.! Spilopleura) and lay less in its posterior part (distance postdorsale post-anal-
28.6 to 29.5% vs 31.2 to 33.6 for S. LS! Spilopleura). The muzzle is short and obtuse, the head and
front are wide. The bones of the infraorbital series are wider in S.! Maculatus (infraorbital
3, 10 to 12.9% SL vs. 7.6 to 8.7 in S.! Spilopleura!, SO4 10.4 to 13.6% SL vs. 8.6 to 10.2)
(Fig.! 5a, 5c). The gap to the cheek is very narrow in S.! Maculatus (5 to 11 times the width
of infraorbital 3 vs 1.9 to 2.5 times in S.! spilopleura) (Fig.! 5b). Postoccipitale the distance, the
predorsal distance, the base of the dorsal and the base of the anal are longer in
S.! Maculatus (Tableau! II). However, the distance interdorsale and distance from the end of the
back at the end of the anal are shorter in S.! maculatus, which explains the impression
greater elongation of the posterior in S.! spilopleura (Fig.! 1).
The mouth of S.! Maculatus, short and massive, is quite close to that of
S.! Spilopleura. In S.! Maculatus, the teeth of the premaxilla and tooth are high and
sharp (Fig.! 3b). The median premaxillary tooth is sharp, with a small cusp in
form of spine to the front of the main lobe and a larger lobe to the back. We
observed 3-6 pairs of teeth ectoptérygoïdiennes the palate.
Meristic characters are homogeneous in the three specimens of S.! Maculatus
(Tableau! III). S.! Maculatus present just under the lateral-line scales
S.! Spilopleura (75-84 vs. 85-96).
The type of coloration of the two specimens of S.! Maculatus is well preserved in alcohol
preserved. The eyes have a vertical black bar, and the spots on the flanks, the size
of the pupil, are much larger and less numerous than in S.! spilopleura. We
have not distinguished as in S. humeral spot! spilopleura. The tip of the ventral
and the distal edge of anal fin are black. The base of the caudal fin is hyaline and the distal edge of the
fin has a black band of the width of the eye where the boundaries are well defined. The
tip of caudal fin rays is not colored, forming a narrow border clear. This type of
coloration is very different from that of S.! spilopleura with 2 / 3 of the base of the caudal
color, leaving a clear strip wider.
These specimens correspond to the descriptions given by Kner (1858, 1860) for
S.! Maculatus and do well in the series-type of this species. We refer to the specimen
NMW 17995 as lectotype of S.! Maculatus, specimen NMW 17996 and NMW 16230
are designated paralectotypes.
There is no doubt that S.! Maculatus and S.! Spilopleura are two different species,
and assumptions of synonymy proposed by the authors should be rejected.
Rehabilitation of S.! Maculatus Kner, 1858
Serrasalmo maculatus Kner, 1858.! -! Kner, 1858!: 166, Rio Guaporé.! -! Kner, 1860!:
41-42, pl. 4 of Fig.! 10, Rio Guaporé.! -! Günther, 1864!: 371, cit. of Kner.
Serrasalmus gibbus Castelnau, 1855. - Azuma, 1990!: 64-69, reproduction, Rio Amazonas,
Manaus.
Serrasalmo spilopleura Kner, 1858. - Günther, 1864!: 370. Capim Rio, Brazil.! -
! Eigenmann, 1915!: 252-253, pl.! 49, fig.! 5, Paraná-Paraguay Basin.! -! La Monte, 1935!: 7,
Juruá, Brazil.
Serrasalmus spilopleura Kner, 1858. - Ulrey, 1895!: Tocantins, Brazil.! -! Norman,
1929!: 798-800, fig.! 13, Rio Capim, Pará, Amazonas, Solimões, Paraná Plata.! -! Pearson
1937!: 109, Rio Beni, Mamore and Paraguay, Bolivia !;!-! Fowler, 1950!: 384-385, in part.! -! Gery,
page 10
130 JEGU & DOS SANTOS
1964!: 27, Peru.! -! Géry, 1976!: 53.! -! Géry, 1978!: 286, 290, fig.! P.! 292.! -! Rodrigues et al.
1978!: Reproduction, Brazil.! -! Ulibarri, 1982! Anatomy, Rio Paraná, Argentina.! -! Junk et al.
1983!: 407, Tab.! 3, Ilha Marchantaria, Rio Solimões, Amazonas, Brazil.! -! Santos et al. 1984!:
35, Rio Tocantins.! -! Miquelarena, 1984!: 615, fig.! 37, caudal skeleton.! -! Lauzanne and Loubens,
1985!: 53, 110, fig.! 48, Rio Mamore and Itenez (Guaporé), Bolivia.! -! Miquelarena, 1986!:
41-42, fig.! 23, Punta Lara and Corrientes, Argentina.! -! Ortega and Vari, 1986!: 9, cit., Peru.! -
! Gery et al. 1987!: 452-454, fig.! 58, 59, Rio Paraguay and Rio Parana, Argentina.! -! Sazima
Guimarães, 1987!: Ethology, Pantanal, Brazil.! -! Zamprogno and Sazima, 1985!: Ethology, Rio
Atibaia, Brazil.! -! Jégu and Santos, 1988!: 255-257, Tab.! 7, 8! Fig.! 9, 10!, Pl.! 1-5, Fig.! G!, pl.! 11,
Fig.! 1, 2, Rio Tocantins.! -! Iriart and Lopez, 1989!: Rio Salado Basin, Argentina.! -! Santos
1990!: 176, fig.! 178, Guaporé, Brazil.! -! Sazima and Machado, 1990!: 17-31, ethology, Pantanal,
Brazil.! -! Porto et al. 1991!: Cytogenetics, Rio Amazonas.! -! Thatcher, 1991!: 570, parasitology.
! -! Lauzanne et al. 1991!: Tabl.! 1, Mamore, Madre de Dios, Iten, Bolivia !;!-! Porto et al.
1992!: Tabl.! 1, cytogenetics.! -! Cestari and Galletti, 1992!: Karyotype, Paraná-Paraguay basin.! -
! Kristsky et al. 1992!: 38, parasitology, Rio Solimões, Brazil !;!-! Every and Kritsky, 1992!: 53 -
57, 62-63, Rio Solimões and Amazonas, Brazil !;!-! Thatcher, 1993!: Parasitism, Rio Guaporé,
Brazil.! -! Martins-Santos et al. 1994!: Karyotype.! -! Castro, 1994!: 64, fig.! 25, Rio Putumayo,
Colombia.! -! Hamann, 1995!: Parasitism, Rio Paraná, Argentina.! -! Biston and Haro, 1995!:
eating habits, Rio Dulce, Argentina.! -! Santamaria, 1995!: 78, Rio Paraná-Igara, Case.
Rio Putumayo, Colombia.! -! Lamas and Godinho, 1996!: 161-168, reproduction, Rio Parnaiba,
Brazil.! -! Orti et al. 1996!: Phylogeny, Rio Paraguay.! -! Kritsky et al. 1996!: Parasitology, Rio
Amazonas, Brazil.! -! Oliveira et al. 1996!: Physiology, Rio Tocantins, Brazil.! -! Cavicchioli and
al. 1997!: 97-106, larval morphology, Rio Paraná, Brazil.! -! Agostinho et al. 1997!:
ethology, Rio Paraná, Brazil.! -! Kritsky et al. 1997!: parasitology, Rio Amazonas, Brazil.! -
! Ferreira et al. 1998!: 82, fig.! 52, Rio Amazonas, Santarem.! -! Giarrizzo, 1998!: Lago do Arar
Island of Marajó, Pará, Brazil !;!-! Jégu and Keith, 1999a!: Tabl.! 1, 3, Fig.! 6c, and Rio Araguari Oiapoque,
Amapá, Brazil.! -! Jégu and Keith, 1999b!: Tabl.! 1, fig.! 2, 3, Amazon Basin.! -! Britski et al.
1999!: 66, figure, Pantanal, Brazil.! -! Nakayama et al. 2000!: Cytology, Rio Solimões and Amazonas,
Brazil.! -! Cetofante, 2000!: Cytology, Central Amazon, Brazil.
We compared the types of S.! Maculatus and S.! Spilopleura to literature data
and directly to specimens from the Rio Amazonas, Tocantins Basin (Brazil),
basin of the Mamore (Bolivia) and Guaporé (or Itenez), initially reported in S.! spilopleura.
The list of material examined appears in Appendix I.
The specimens reported in S.! Spilopleura have a wide range infraorbital area
naked very close to the cheek (Fig.! 5a-c). The type of coloration of the young, with black spots
the size of the pupil on the sides and a band tail clear black terminal to subterminal,
remember the color of the types of S.! maculatus. The position and width of this band
vary by region. In the basin of the Paraná-Paraguay, the black band is narrow,
size similar to that of the distal light zone, blackheads are larger than in the basin
Amazon (Gery et al. 1987!: fig.! 58, 59!, pers. obs.). In the Tocantins (Santos et al.
1984!: 35!; Jégu and Santos, 1988!: 256, pl.! 11, fig.! 1, 2) and the marshes of Amapá (pers. obs.!:
ZMA 107-764 and 755, Oyapock), the black band is wide and located very near the end of
radii. In Central Amazon, both in the region of Manaus (Fig.! 6a, 6b) that nearly
Santarem (Ferreira et al. 1998!: Fig.! 52), in the Bolivian Amazon (Lauzanne and Loubens,
1985!: Fig.! 48) and in the Colombian Amazon (Castro, 1994!: Fig.! 25) the black band is broad and
located closer to the end of the rays in adults than in young people. As in
types of S.! maculatus, there is a black band on the distal edge of the adipose
the ends of the rays of the anal is black and the eye has a vertical black band. The most(132)
Serrasalmus spilopleura and S. pictures maculatus 131)
Fig.! 6.! -! Serrasalmus maculatus Kner. Change in color in the Lago do Rei, Ilha do Careiro, Rio
Amazonas. A!: Young!; B!: Adult!; C!: Adult in breeding (?). [Serrasalmus maculatus Kner. variation
of the color pattern in the Lago do Rei, Ilha do Careiro, Rio Amazonas. A: Juvenile; B: Adult C:
Breeding adult (?).]
132 JEGU & DOS SANTOS
large adults are stained darker, yellow-violet (Amapá) to purple (Azuma
1990: 65 fig.! Sup., Amazonas, Manaus: Fig.! 6c), the unpaired fins becoming uniformly
gray to black, but the black to the eye persists. The purplish color is
observed only in large specimens, and Santos was able to verify that all copies with
such coloration were in breeding season. It is likely to change
related to the sexual cycle, a hypothesis to be confirmed. These observations show
clear the misunderstanding of the authors in interpreting the type of color in
S.! Spilopleura and S.! Maculatus.
The types of S.! Maculatus have the body and the infraorbital series and wide area naked
cheek very narrow features observed in the specimens reported to
S.! Spilopleura Tocantins (Jégu and Santos, 1988) and the Paraná-Paraguay (Gery et al., 1987).
Dentition specimens of Tocantins (Jégu and Santos, 1988!: Pl.! 2, Fig.! G) and Basin
Paraguay (Miquelarena, 1986!: Fig.! 23) reported S.! Spilopleura is quite comparable to
the types of S.! maculatus (Fig.! 3b).
Specimens of the basins of the Amazon and Paraná-Paraguay, mistakenly identified
as Serrasalmus spilopleura, must be appointed S.! maculatus Kner, 1858. Nakayama
et al. (2000) highlight the presence of two cytotypes assigned to
S.! Maculatus in the region of Manaus at the confluence of the Rio Negro and Solimões. The authors
recall that the other four cytotypes have been detected in S.! of maculatus
Paraná-Paraguay system, including two involving a hybrid zone in the lower Paraná (Cestari
and Galetti, 1992). S.! Maculatus would not be uniform throughout its range.
Yet the meristic characters of the specimens in the Amazon and Paraná-Paraguay are
very similar (Tableau! IV) except for the teeth ectoptérygoïdiennes. Norman
(1929!: 798) and Géry et al. (1987) do not observe more than 4 or 5 teeth in the palate
specimens of the Paraná-Paraguay, Miquelarena (1986!: 41) points out six teeth in specimens
Argentina, while we found up to 7 pairs of teeth in the palace of the specimens
Amazon Basin (Fig.! 7). The diameter of the eye would be reduced in the Amazon than in the
Paraná-Paraguay (including 4.5 times or less in length from head to Amazon vs.
up to 5.5 times after Géry et al. or Norman), but we have not found this difference
among the four specimens we examined Paraná (Tableau! V).
Finally, the taxonomic position of S.! Maculatus seems uncertain. Gery et al.
(1987) point out that S.! Maculatus (S.! Spilopleura in the text) can be easily confused
Pygocentrus nattereri with which it is distinguished in particular by the narrower head and
Table! IV.! -! Geographic variation in meristic characters Serrasalmus maculatus Kner, 1858.
[Geographical variability of meristic characters in Serrasalmus maculatus Kner, 1858.]
page 15 with board (number 133 on pdf doc)
Fig.! 7.! -! Serrasalmus maculatus Kner. Change in number of teeth depending on the ectoptérygoïdiennes
standard length in the Amazon basin and the Paraná. [Serrasalmus maculatus Kner. Variation of the
ectopterygoid teeth number versus standard length in the Amazon and Paraná Basins.]
presence of teeth on the palate. The authors also consider S.! Maculatus as a form of transition
gender Serrasalmus and Pygocentrus. Fink (1993) points out that in these spe-Pygocentrus
have a head much wider than the other piranhas. but Orti
et al. (1996) show that S.! Maculatus (S.! Spilopleura in the text) should be excluded from the genus Serrasalmus
to form the sister group of Pygocentrus.
Status of S.! Spilopleura Kner, 1858
The further description of Kner (1860), combined with examination of the types of
S.! Spilopleura allows us to identify several criteria for this species. The body, rather
lying is not rhomboidal. The head and body are broad and blunt snout. we know
Table! V.! -! Serrasalmus maculatus! Kner, 1858. Geographic variation of morphometric characters
used by Gery et al., 1987. [Serrasalmus maculatus! Kner, 1858. Geographical variability of morphometric
Used characters in Géry et al., 1987.]
page 16 (number 134 on pdf doc)
Fig.! 8.! -! Changes A!: body height, B!: predorsal distance, C!: the width of the infraorbital
3, D!: the gap in the cheek and E!: the infraorbital 4, depending on the LS, in S.! Hollandi and
P.! Eigenmann of the Bolivian Amazon and in the types of S.! Spilopleura Kner. [Variations of A: body
depth, B: predorsal length, C: third infrorbital width, D: naked area on cheek and E: fourth infrorbital
width in S.! Holland and P.! Eigenmann from the Bolivian Amazonia and in the S.! spilopleura typespecimens Kner,
Expressed versus SL.]
S. also! spilopleura has a wide gap in the cheek, even in large specimens,
and not more than five teeth on the palate. The jaws are short and the teeth to the jaws are low.
The caudal fin is black from the base up to 2 / 3 rays, but the free edge is hyaline.
Lauzanne et al. (1991) reported three species of the Mamore and Serrasalminae
the Itenez (Guaporé) with the edge of the tail is clear!: S.! maculatus (identified S.! spilopleura).
page 17 (135)
Fig.! 9.! -! Left Side View of the specimens of Rondonia, Brazil, identified S.! Aureus (A) and Serrasalmus
sp.! 1[B] by Santos (1990). [Left lateral view of specimens from Rondonia, Brasil, Identified as S.! Aureus
(A) and Serrasalmus sp.! 1 [B] by Santos (1990).]
Table! VI.! -! Variability of meristic characters of Serrasalmus hollandi Pristobrycon and Eigenmann
of the Bolivian Amazon. [Meristic characters variability in Holland and Serrasalmus Pristobrycon Eigenmann
from the Bolivian Amazonia.]
page 18 (136)
S.! Hollandi describes the Guaporé Pristobrycon and Eigenmann. Santos (1990) cites three species
Guaporé and reports of the presence of a fourth form of the Jamari, a right tributary of Madeira,
with the caudal edge of the hyaline and identified S.! aureus. These species are all different
types of S.! spilopleura.
S.! Hollandi differs from S.! Spilopleura by the more general form of the body rhomboidal,
head more slender, the dorsal inserted far forward, the infraorbital 3 and 4! broader and
bare area closer to the cheek (Fig.! 8c-e). In addition, S.! Holland, between 50 and 185! Mm SL,
This 6-10 pairs of teeth ectoptérygoïdiennes (Tableau! VI) against less than 5
S.! Spilopleura, by Kner (1858). Finally, the mark at the base of the caudal fin in S.! Hollandi
is crescent shaped and much narrower than in S.! spilopleura.
The type of coloration of the tail of P.! Eigenmann is close to that of
S.! Spilopleura, especially for large specimens in whom the distal edge of the
Brand tail is straight. The number of teeth is low in ectoptérygoïdiennes
specimens of P.! Eigenmann over 100! mm SL (Tableau! VI). In P.! Eigenmann,
S. at home! spilopleura, the muzzle is short and pug, narrow infraorbital bones (Fig.! 8c
8e) and the large gap in the cheek (Fig.! 8d). P.! Eigenmann is different types of
S.! Spilopleura by body height (Fig.! 8a) and the insertion of the dorsal farther back
(Fig.! 8b). Moreover, in P.! Eigenmann diameter of the eye is slightly larger, the
Table! VII.! -! Variability of morphometric characters of Serrasalmus and hollandi Pristobrycon
Eigenmann of the Bolivian Amazon (numbers in parentheses indicate the number of specimens
examined for these characters). [Variability of morphometric characters in Holland and Serrasalmus
Pristobrycon Eigenmann from the Bolivian Amazonia (numbers in brackets Indicate number of specimens
Examined For Each character).]
page 19 (137)
Serrasalmus spilopleura and S. maculatus 137
interdorsale distances and longer pre-anal and dorsal and pectoral more
shorter than in S.! spilopleura (Tableau! VII).
The specimen from the Rio Jamari (Rondonia, Brazil) identified S.! Aureus by Santos (1990)
presents a well-growing fairly large caudal (Fig.! 9a), but differs from types of
S.! Spilopleura the narrower interorbital (11.6% of SL vs. 12.3 to 13.8% in
S.! Spilopleura) and the much wider infraorbital 3 (11.2% vs LS less than 9%
S.! Spilopleura). The black spots on the flanks are much larger in this specimen, but
we observed such a variation between specimens of P.! Eigenmann 130 and
140! Mm SL, from the same collection station (MNHN 1989-1362).
Santos (1990!: Fig.! 186) reported Serrasalmus sp.! 1, the Guaporé and Jamari, including
general appearance of the body and morphometric characteristics are very similar to those of
S.! Spilopleura (Tableau! VII), including the level of the infraorbital series. The head and base
of the ridge are slightly longer than the types of S.! spilopleura. None of these
specimens of teeth does the palate. The black dots on the sides, the presence of a
humeral marks and gray color of the tip of the first rays of the anal is similar
that we observed on the lectotype of S.! spilopleura. These specimens differ
types of S.! spilopleura by the mere presence of a gray to black band located
the tip of the tail (Fig.! 9b), although specimens have the caudal edge of the most
clear.
These specimens can not be identified in S.! Spilopleura Kner. Examination of
additional material from the Guaporé will better appreciate the variability
type of coloration of the caudal and possibly expand the diagnosis S.! spilopleura.
S.! Spilopleura this type of color much like that of specimens
S.! Goulding Fink & Machado-Allison, preserved in alcohol (Fink and Machado-Allison, 1992!:
Fig.! 2). The two species are not more than five teeth on the palate, teeth have shapes
adjacent to the premaxilla (ibid.!: fig.! 8b). But in S.! Goulding 3 covers the infraorbital
completely plays while that of S.! spilopleura reminds Pristobrycon careospinus
(Ibid.!: Fig.! 8b).
The truncated form of the infraorbital 3 with a wide gap, the low number of
ectoptérygoïdiennes teeth and lower teeth as seen in the premaxilla
S.! Spilopleura also recall the characters of Pristobrycon Calmon, type species of Pristobrycon.
Only obtaining new specimens of Serrasalmus spilopleura Kner, 1858,
associated with an osteological study finer will better appreciate the relationship of this
species with the kind Pristobrycon.
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Accepted for publication on 03.01.2001.
page 23 (141)
Serrasalmus spilopleura and S. maculatus 141
ANNEX! I
List of specimens examined
List of acronyms
INPA!: Instituto Nacional de Pesquisas da Amazônia, Manaus!; MCL!: Museu Costa Lima, Macapá
!; MNHN!: National Museum of Natural History, Paris!; MNRJ!: Museu Nacional do Rio de Janeiro,
Rio de Janeiro!; MZUSP!: Museu da Zoology of Universidade de São Paulo, São Paulo!, NMW!: Naturhistorisches
Museum, Vienna!; USNM!: National Museum of Natural History, Washington!, UTB!: Universidad
Técnica del Beni, Trinidad!; ZMA!: Zoological Museum van Amsterdam, Amsterdam.
Serrasalmus maculatus Kner, 1858
Lectotype of S.! Maculatus Kner, 1858!, NMW 17995, 162! Mm SL, Mato Grosso, 1826, Natterrer,
alcohol. Paralectotypes of S.! Maculatus Kner, 1858!, NMW 17996, 1 ex., 122! Mm SL, Mato Grosso,
1826, Natterrer, alcohol!, NMW 16230, 1 ex., 154! Mm SL, Caiçara, Rio Guaporé, Natterer, XI (1!?) 825.
Supplementary material
Rio Mamore, Bolivia.! -! MNHN 1989-1365, 2 ex., Laguna Mocovi, Trinidad!, MNHN 1989-1366,
An ex., Laguna Coitarama, Trinidad!, MNHN 1989-1367, 1 ex., Rio Secur, Laguna Candelaria!, UTB
UNCAT., Laguna Candelaria, Rio Secure!; UTB UNCAT, 1 ex., Laguna Patajusal, Rio Secure!, USNM 310990,
Beni.
Rio Madre de Dios, Peru.! -! USNM 319654, 1 ex., Manu river drainage!, USNM 319299, 1 ex.,
Manu river drainage.
Rio Itenez (or Guaporé), Bolivia and Brazil.! -! UTB UNCAT., 14 ex., Boca do Rio Machupo!, UTB
UNCAT., 2 ex., Rio Blanco!; UTB UNCAT., 1 ex., Laguna da Lundra!; polo INPA 213, one ex., 87.6! mm SL, Rio
Guaporé, Surpresa!; Polo INPA 286, one ex., 144! Mm SL, Rio VIP Novas, Guajara Mirim!; INPA polo
225, 4 ex., 76-147! Mm SL, Rio Guaporé, Surpresa!; INPA UNCAT, ex 2, 5-69! Mm SL, Rio Guaporé in
frente to Surpresa
Rio Tocantins.! -! INPA 1006, 12 ex. Lower Tocantins!, INPA 1020, 4 ex., Lower Tocantins!; INPA
1025, two ex., Acari Pucu!, INPA 1026, 2 ex., Tucurui Market!, INPA 1027, 2 ex., Tucurui Market!; INPA
1028, 4 ex., Jatobal!, INPA 1029, 9 Ex., Lower Tocantins, downstream Itupiranga!, INPA 1030, 1 ex., Igarapé Canoal,
Breu Branco!, INPA 1031, 1 ex., Itupiranga!, INPA 1032, 1 ex., Lago Otacilio, Itupiranga!; INPA
1033, two ex., Acari Pucu!, INPA 1034, 2 ex., Lower Tocantins, downstream Itupiranga!, INPA 1035, 1 ex., Itupiranga
!, INPA 4503, 6 ex., Réservatorio Tucurui!; MZUSP UNCAT, 1 ex., Ig. Muru, Tucurui.
Rio Amazonas, Peru.! -! MNHN 1996-1004, 1 ex., Iquitos.
Rio Amazonas, Brazil.! -! INPA 3628, 16 ex., Lago do Rei, Ilha de Careiro!; INPA UNCAT., 27 ex.,
Lago Catalão, Confluencia com Rio Negro!, MNHN 1909-1979, 1 ex., Santarem!, MNHN 1909-158, an ex.,
Tefé!; INPA UNCAT, 2 ex., Ilha Marchantaria, Amazonas!; MZUSP UNCAT, 1 ex., Itacoatiara!; MZUSP UNCAT,
An ex., Rio Arar Marajo!, USNM 233486, 5 ex., Paraná do Lago Januaca!, USNM 308966, 7 ex., Ilha da
Marchantaria!, USNM 309102, 1 ex., Janauaca of Paraná, entrada do Lago do Castanho!, USNM 309082,
6 ex., Castanho do Lago, Januaca!, USNM 309083, 5 ex., Lago Murmumuru, Januacá!, USNM 309107, 5
ex., do Paraná Januaca!, USNM 309065, 11 ex., Lago Murumuru, Januaca!, USNM 233496, 1 ex., Lago
Castanho, Paraná do Januaca!, USNM 233488, one ex., Lago Januari!, USNM 233489, one ex., Entrada do
Lago Januari!, USNM 308976, one ex., Canta Galo Lago, Lago Januari!, USNM 309085, 4 ex., Lago Canta
page 24 (142)
142 JEGU & DOS SANTOS
Galo, Lago Januari!, USNM 309090, 2 ex., Lago Januari!, USNM 309081, 5 ex., Lago Castanho, Januaca
!, USNM 23499, 1 ex., Lago Castanho!, USNM 309283, 50 copies., Lago Januari.
Uatumã Rio, Brazil.! -! INPA UNCAT, 3 ex., 145-160! Mm SL, Lago Tapan, Frente a Villa Santana.
Vila Nova and Rio Anauerapucu, Amapá.! -! MNRJ 12534, 6, 90-140, Rio Vila Nova, Ig. do Lago,
afl. Margem esq do VN, a prox Babolândia (51 ° 25'54 .5 "W-00 ° 11'43 .6" N), April 1981.
Rio Macacoari, Amapá.! -! MCL 101, 4, 65-85, Braco do Rio Macacoari has jusante Carmo, Municipio
of Itaubal (50 ° 47'10 .9 "W-00 ° 34'21 .8" N), April 15, 1982!, MCL 122, 2, 60-65, same loc.!, MCL
125, 1, 85, same loc.!, MCL 102, 6, 70-75, same loc!, MCL 106, 2, 70-75, Braco Macacoari do Rio, a
jusante Carmo, Mun. of Itaubal, Rio Macacoari (50 ° 47'10 .9 "W-00 ° 34'21 .8" N), 16 jul. 1982!, 113, 2,
65-70, Braco do Rio Macacoari has jusante Carmo, Mun. of Itaubal, Rio Macacoari (50 ° 47'10 .9 "W-
00 ° 34'21 .8 "N), November 15, 1982!, MCL 123, 1, 100, Braco do Rio Macacoari has jusante Carmo, Municipio
of Itaubal, (50 ° 47'10 .9 "W-00 ° 34'21 .8" N), February 19 1981!, MCL 149, 1, 120, Braco Macacoari do Rio, a
jusante Carmo, Mun. of Itaubal, Rio Macacoari (50 ° 47'10 .9 "W-00 ° 34'21 .8" N), March 19, 1982!, MCL
952, 1, 135, Cidade de Carmo, Mun. of Itaubal, Rio Macacoari (50 ° 48'32 .7 "W-00 ° 34'63 .5" N), November 24
1982!, MCL 992, 2, 70-85, Cidade de Carmo, Mun. of Itaubal, Rio Macacoari (50 ° 48'32 .7 "W-
00 ° 34'63 .5 "N), February 15 1983!, MCL 1106, 1, 65, Cidade de Carmo, Mun. of Itaubal, Rio Macacoari
(50 ° 48'32 .7 "W-00 ° 34'63 .5" N), 15 jul. 1982!; MNRJ 12533, 2, 75-79, Braco do Rio Macacoari
(50 ° 35'27 .2 "W-00 ° 45'16 .4" N), April 1981.
Rio Araguari, Amapá.! -! MNHN 1998-267, 7, 140-170, Araguari, Lago Comprido, reserva Piratuba
(50 ° 09'16 .4 "W-01 ° 22'21 .8" N), August 15, 1992!; INPA (AP 33b). 11, 137-166, same loc.!; MNHN
1998-184, 5, 55-100. Araguari, Vista Alegre (50 ° 16'21 .8 "W-01 ° 19 '38.2" N), August 15, 1992!; INPA (AP
34), from 6.55 to 137, same loc.!; MNRJ 12541, 16, 31-40, Rio Aporema, afl. esq do Rio Araguari, Fazenda
Aporema, in April 1981!; MNRJ 12539, 3, 155-175, same loc.!; MNRJ 12540, 24, 90-150, same loc.
Rio Flexal, Amapá.! -! MCL 98, 1, 90, Lago Pracuúba, Mun. Pracuúba, drenagem do Rio Flexal
(50 ° 46'54 "W-01 ° 45'16" N), Jul 19-23. 1984!, MCL 111, 1, 55, same loc.!, MCL 982, 1, 130!, MCL 985,
1, 130, same loc.!, MCL 994, 4, 40-55, same loc.!, MCL 1104, 1, 90, same loc!, MCL 943, 2, 65-70,
Lago Pracuúba, Mun. Pracuúba, drenagem do Rio Flexal (50 ° 46'54 "W-01 ° 45'16" N), January 25 1984!, MCL
335, 1, 160, Lago Pracuúba, Mun. Pracuúba, drenagem do Rio Flexal (50 ° 46'54 "W-01 ° 45'16" N), 24-26
October 1986!, MCL 376, 1, 130, same loc.!, MCL 377, 1, 130, same loc.!, MCL 560, 1, 90, same loc.!;
MCL 661, 3, 115-145, same loc.!, MCL 866, 2, 134-140, same loc.!, MCL 872, 1, 155, same loc.!;
MCL 873, 1, 140, same loc.!, MCL 874, 1, 130, same loc.!, MCL 876, 1, 130, same loc.!, MCL 343, 1,
170, Rio Flexal, Região dos Lagos, Pracuuba, Lago Comprido (50 ° 44'10 .9 "W-01 ° 42'32 .7" N), October 27
1986!, MCL 375, 1, 160, Fazenda Sta. Tereza, Lago Pracuúba, Mun. Parcuúba, drenagem do Rio Flexal
(50 ° 46'54 "W-01 ° 45'16" N), November 2, 1986.
Fl Oyapock, Amapá.! -! MNHN 1981-586, 3, Juminan Creek (51 ° 38'06 .5 "W-04 ° 03'30 .8" N), 20
September 1977!, MNHN 1998-137, 2, 70-75, same loc.!; ZMA 107-755, 2, 110-120, same loc.!; ZMA 107 -
764, 1, 60, same loc.!, MNHN 1998-111, 5, 100-160, Taparabu Creek, Savannes Taparabu
(51 ° 41'06 .3 "W-03 ° 58'35 .9" N), 11-13 October 1997!, MNHN 1998-1991, 2, 137-140, same loc.!, MNHN 1998 -
1890, 1, 165, same loc.!, MNHN 1998-1992, 5, 135-160, same loc.!, MNHN 1998-1993, 3, 129-160, even
loc., 15-17 October 1996.
Serrasalmus spilopleura Kner, 1858
NMW 57085, lectotype of S.! Spilopleura, 160! Mm SL, Rio Guaporé, Natterer, alcohol. Paralectotypes
!: NMW 16344, 1 ex., 155! Mm SL, Rio Guaporé, Bogota, Natterer, dry!, NMW 79457, 1 ex.,
210! Mm SL, Rio Guaporé, Mato Grosso, Natterer, dry!, NMW 79458, 1 ex., 155! Mm SL, Mato Grosso,
page 25 (143)
Serrasalmus spilopleura and S. maculatus 143
Natterer, 1829!, NMW 79459, 1 ex., 145! Mm SL, Mato Grosso, Natterer, 1829.
Serrasalmus sp.! 1
Guaporé basin, Brazil.! -! Polo INPA 360, 4 ex., 160-180! Mm SL!; Polo INPA 442, one ex.,
200! Mm SL!; INPA polo 68, 1 ex., 155! Mm SL!; Polo INPA 1235, 3 ex., 162-175! Mm SL.
Serrasalmus Spix & Agassiz cf aureus, 1829
Madeira Basin, Rio Jamari, Brazil.! -! Polo INPA 1171, 1 ex. 147! Mm SL.
Serrasalmus hollandi Eigenmann, 1915
Guaporé basin, Brazil, and Iten, Bolivia.! -! UTB UNCAT., 1 ex., 64! Mm SL, Rio Blanco, affl. R.
Itenez.
Mamore Basin, Bolivia.! -! UTB UNCAT., 9 Ex., 69-92! Mm SL, Rio Mocovi!; UTB UNCAT., 11 ex.,
37-74! Mm SL, Lago San Jose, r. Tijamuchi, affl. Mamore!, UTB, UNCAT., 9 Ex., 53-67! Mm SL, Coneto,
Trinidad!; UTB UNCAT., 8 ex., 100-110! Mm SL, R. Secure, el Lago Patujusal!, UTB, UNCAT., 2 ex., 29 -
31! Mm LS, Aroyo Sao Jose, Trinidad!; UTB UNCAT., 4 ex., 130-165! Mm SL, Rio. Secure, L. Remanso!;
UTB UNCAT., 4 ex., 137-183! Mm SL, Rio Secur, L. Candelaria!; UTB UNCAT., 3 ex., 123-134, Rio Secur
El Lago Patujusal!, UTB, UNCAT., 113-177! Mm SL, Rio Mocovi, L. Mocovi!, MNHN 1989-1363, 5 ex., 70 -
165, Lago Candelaria, Rio Secur.
Pristobrycon Eigenmann (Norman, 1929)
Mamore Basin, Bolivia.! -! UTB UNCAT., 2 ex., 40-53! Mm SL, R. Tijamuchi, affl. Mamore, L.
São José!; UTB UNCAT., 1 ex., 151! Mm SL, L. Chimimita!; UTB UNCAT., 6 ex., 129-164! Mm SL, R. secure,
L. Candelaria!; UTB UNCAT., 1 ex. 63, 7! Mm SL, Boca do Machupo, R. Itenez!; UTB UNCAT., 1 ex.,
95.7! Mm SL, R. Secure, el Lago Patujusal!, UTB, UNCAT., 5 ex., 129-157! Mm SL, R. Mocovi, L. Mocovi!;
MNHN 1989-1362, 3 ex., 130-180! Mm SL.
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