Revision of the Neotropical trahiras of the Hoplias lacerdae species-group (Ostariophysi: Characiformes: Erythrinidae) with descriptions of two new species
Author
Oyakawa, Osvaldo T.
Author
Mattox, George M. T.
text
Neotropical Ichthyology
2009
2009-06-30
7
2
117
140
http://www.scielo.br/scielo.php?script=sci_arttext&pid=S1679-62252009000200001&lng=en&tlng=en
journal article
10.1590/S1679-62252009000200001
1982-0224
5420421
0041B40D-74DE-41E3-B245-0D3B36B695F2
Hoplias lacerdae
Miranda-Ribeiro, 1908
Fig. 10
Hoplias lacerdae
Miranda-Ribeiro, 1908: 3
[original description, type locality: rio Ribeira de Iguape, Iporanga,
São Paulo state
,
Brazil
;
Holotype
: MNRJ 211, dried specimen]. -
Eigenmann, 1909: 374
[reference, occurrence in the rio Ribeira de Iguape]. -
Eigenmann, 1910: 448
[reference, occurrence in the rio Ribeira de Iguape]. -
Fowler, 1950: 364
[in species list]. -
Azevedo
et al
., 1965: 103
[redescription, differences from
H
.
malabaricus
]. -
Britski, 1972: 81-82
[in part, species list, differences from
H
.
malabaricus
, occurrence in the rio Ribeira de Iguape basin]. -
Géry
et al
., 1987: 368
[identification key]. -
Bizerril, 1994: 56
[in part, occurrence in eastern river basins]. -Bizerril & Lima, 2000: 107 [occurrence in the rio Ribeira de Iguape]. -
Oyakawa, 2003: 239
[species list, distribution]. -
Oyakawa
et al
., 2006: 166
[species list with pictures, differences from
H
.
malabaricus
]. -Oyakawa & Netto-Ferreira, 2007: 64 [species list, distribution]. -
Menezes
et al
., 2007: 153
[typelocality, distribution].
Hoplias
aff.
lacerdae
: -
Malabarba, 1989: 127
[remarks on occurrence in the rio Jacuí,
Rio Grande do Sul
]. -
Zaniboni Filho
et al
., 2004: 104
[biological and ecological features, with picture].
Hoplias lacertae
: -
Géry, 1977: 102
[unavailable name,
lapsus calami
, remarks on validity of name, identification key].
Holotype
.
BRAZIL
.
São Paulo State
:
MNRJ 211
, 750.0 mm SL,
rio Ribeira de Iguape
,
Iporanga
, no date,
R
. Krone. (
Fig. 11
).
Material examined.
BRAZIL
.
São Paulo State
.
Ribeira de Iguape basin:
MNRJ
11217, 1, 443.0 mm SL, rio Juquiá, tributary to rio Ribeira de Iguape, Registro;
MZUSP
7900, 1, 625.0 mm SL;
MZUSP
7901,
1, 380.3 mm
SL, rio Ribeira de Iguape;
MZUSP
38569,
1, 157.2 mm
SL, rio Ribeira de Iguape, Registro;
MZUSP
7902, 1, 367.0 mm SL;
MZUSP
37989, 2, 279.4-
417.2 mm
SL, rio Pardo on the road Iporanga-Barra do Turvo,
14 km
from Barra do Turvo;
MZUSP
45433,
1, 234.9 mm
SL, rio Ribeira de Iguape by rio Pilões, Eldorado;
MZUSP
51104, 1, head length
244.5 mm
SL, rio Juquiá, Registro;
MZUSP
61748,
1, 150.5 mm
SL, rio André Lopes, Iporanga;
MZUSP
72953,
1, 388.6 mm
SL, rio Jacupiranga, Registro.
Santa Catarina State.
Uruguay basin:
MCP
12339, 1, 202.0 mm SL;
MCP
12365, 1,
65.3 mm
SL, rio Canoas between Vargem and São José do Cerrito, Campos Novos;
MCP
12731,
1, 295.5 mm
SL;
MCP
12732,
1, 224.53 mm
SL;
MCP
12879, 2, 130.7-245.0 mm SL, rio Canoas at Passo do Canoas, SC-458, Campos Novos;
MCP
12338,
1, 218.8 mm
SL;
MCP
12745, 2, 242-
249.3 mm
SL;
MCP
13143,
1, 152.9 mm
SL;
MCP
12878, 3, 297.2-
344.4 mm
SL, rio Canoas, on Abdon Batista-Anita Garibaldi road, Campos Novos;
MCP
12884,
1, 387.8 mm
SL;
MCP
12898, 3,
63.4-147.1 mm
SL;
MCP
41003,
1, 115.3 mm
SL, rio Canoas, on Abdon Batista-Anita Garibaldi road;
MCP
13420, 1, 180.0 mm SL;
MCP
18427,
1, 203.8 mm
SL;
MCP
18828,
1, 226.6 mm
SL;
MCP
18925, 2, 242.6-301.0 mm SL;
MCP
19151,
1, 402.1 mm
SL;
MCP
19035, 2, 243.1-
325.6 mm
SL, rio do Peixe in Volta Grande, Concórdia;
MCP
19146,
1, 411.5 mm
SL;
MCP
41004,
1, 253.3 mm
SL, rio Riacho Grande, on Piritiba-BR 153 road, Concórdia;
MCP
19147,
1, 415.1 mm
SL, rio Uruguai in Itá;
UFRGS
7530,
1, 238.6 mm
SL, rio Jacutinga near water capture, Concórdia.
Rio Grande do Sul State.
Uruguay basin:
INPA
26892,
1, 144.1 mm
SL;
MCP
12719, 3, 130.5-
185.3 mm
SL, arroio Passo Alto, São Nicolau;
MCP
10896, 3, 110.0-
207.7 mm
SL, rio Garupá, tributary to rio Quaraí, on Uruguaiana-Quaraí road;
MCP
11226,
1, 142.3 mm
SL, tributary to rio Quaraí-Mirim on Quaraí-Alegrete road, Alegrete;
MCP
11231, 4, 120.0-
179.9 mm
SL;
MNRJ
30393,
1, 145.7 mm
SL, rio Garupá on Alegrete-Quaraí border;
MCP
12089,
1, 288.8 mm
SL;
MCP
13421,
1, 301.1 mm
SL;
MNRJ
30394,
1, 302.1 mm
SL, rio Ligeiro, on Marcelino Ramos- Maximiliano de Almeida road;
MZUSP
37254,
1, 107.6 mm
SL;
MZUSP
43683,
1, 196.3 mm
SL, rio Garupá, tributary to rio Quaraí;
MCP
12674,
1, 284.3 mm
SL, arroio Canoin on Pirapó- São Nicolau road, São Nicolau, Rio Grande do Sul;
MCP
13210,
1, 285.8 mm
SL, mouth of rio Ijuí-Mirim, Pirapó;
MCP
14228, 1,
28.4 mm
SL, tributary to
rio Negro
on Bagé-Aceguá road;
MCP
14280,
1, 332.2 mm
SL, arroio Ibicuí da Faxina, Santana do Livramento;
MCP
16802, 2, 304.6-
349.2 mm
SL, rio Ijuizinho near Cermissões dam;
MCP
18875,
1, 250.3 mm
SL, rio Uruguai in Marcelino Ramos;
MCP
16818
, 276.8 mm SL, rio Buricá near CERTHIL dam, Independência;
MCP
18924,
1, 251.4 mm
SL, rio Uruguai in Marcelino Ramos;
MCP
19385, 2, 86.0-
124.6 mm
SL, rio Canoas, on Tupitinga-Celso Ramos road, Tupitinga;
MCP
19378, 2, 288.4-
309.4 mm
SL, rio Pelotas at Passo da Pedra Oveira, Esmeralda;
MCP
21650,
1, 170.8 mm
SL, rio Quaraí-Mirim, on Uruguaiana-Quaraí road;
MCP
26972, 2, 43.0-
43.7 mm
SL, rio Ibirapuitã, under bridge upstream from the Biological Reserve of Ibirapuitã, Alegrete;
MCP
27452,
1, 360.5 mm
SL, Arroio Lagoão, rio Ibicuí drainage, on Santiago-Manoel de Freitas road;
MCP
27481,
1, 144.4 mm
SL, rio Itu, tributary to rio Ibicuí, Santiago;
UFRGS
8394, 6,
68.8-94.9 mm
SL, arroio Piraí, tributary to
rio Negro
between Bagé and Dom Pedrito, Bagé.
Rio Grande do Sul State.
Jacuí basin:
MCP
30714,
1, 230.7 mm
SL, rio Jacuí-Mirim, Carazinho;
MZUSP
25026, 2, 215.9-
225.4 mm
SL, rio Jacuí at Ernestina dam near Passo Fundo.
URUGUAY
.
Artigas
.
Uruguay basin:
MCP
17616, 1,
79.2 mm
SL, arroyo Arumbeba, circa of
22 km
Southeast of
Artigas
;
UFRGS
8059,
1, 101.3 mm
SL, arroyo Mandiyú, tributary to río Uruguay,
Artigas
.
Cerro Largo
.
Uruguay basin:
MCP
9878,
1, 288.6 mm
SL;
MCP
9880,
1, 301.9 mm
SL;
MCP
9896, 1, 506.0 mm SL;
MCP
9926,
1, 399.7 mm
SL,
río Negro
at Arreria.
Tacuarembó
.
Uruguay basin:
MZUSP
52006, 4, 289.6-
321.9 mm
SL,
río Negro
, Rincón del Bonete.
Fig. 10.
Hoplias lacerdae
, MZUSP
61748, 150.5 mm SL, rio Ribeira de Iguape, São Paulo State, Brazil.
Table 7.
Morphometric data of
Hoplias lacerdae
. Standard length in mm; values 1-14 are percents of standard length and values 15-22 are percents of head length. n = number of examined specimens.
Holotype |
n |
Mean |
Range |
Standard deviation |
Standard length |
750.0 |
93 |
- |
28.4-750.0 |
- |
1. Body depth |
- |
88 |
21.4 |
16.8-25.1 |
1.8 |
2. Head length |
28.0 |
89 |
31.1 |
27.6-34.9 |
1.5 |
3. Pectoral-fin length |
- |
89 |
16.9 |
11.6-19.2 |
1.3 |
4. Pelvic-fin length |
- |
89 |
17.9 |
15.0-20.4 |
1.3 |
5. Anal-fin length |
- |
89 |
17.3 |
13.2-19.7 |
1.2 |
6. Dorsal-fin length |
- |
89 |
26.2 |
23.1-30.6 |
1.6 |
7. Dorsal-fin base |
13.9 |
89 |
15.5 |
13.6-18.0 |
0.9 |
8. Anal-fin base |
6.2 |
89 |
9.1 |
6.2-11.3 |
0.8 |
9. Pre-pectoral distance |
- |
88 |
28.5 |
24.6-34.4 |
1.7 |
10. Pre-pelvic distance |
56.7 |
88 |
56.0 |
50.6-60.8 |
2.1 |
11. Pre-dorsal distance |
50.0 |
88 |
51.3 |
47.6-56.9 |
2.0 |
12. Pre-anal distance |
82.0 |
88 |
81.4 |
75.1-87.6 |
2.6 |
13. Caudal-peduncle depth |
11.0 |
88 |
13.1 |
10.8-15.1 |
0.7 |
14. Caudal-peduncle length |
12.0 |
88 |
13.6 |
11.0-16.4 |
1.0 |
15. Head depth |
45.1 |
90 |
41.7 |
37.8-47.1 |
2.0 |
16. Snout length |
26.3 |
90 |
25.1 |
22.5-29.0 |
1.1 |
17. Snout width |
- |
90 |
18.9 |
13.6-22.9 |
1.3 |
18. Snout depth |
- |
89 |
19.1 |
16.2-21.8 |
1.3 |
19. Pre-nasal distance |
- |
90 |
15.0 |
11.4-19.1 |
1.2 |
20. Orbital diameter |
8.0 |
90 |
18.5 |
8.0-27.8 |
3.7 |
21. Interorbital width |
31.0 |
90 |
21.1 |
15.6-30.1 |
2.9 |
22. Upper jaw length |
51.4 |
90 |
47.7 |
44.9-51.8 |
1.5 |
Diagnosis.
Hoplias lacerdae
differs from the other species of
H
.
lacerdae
group in the number of pores of the laterosensory system along the ventral surface of dentary (6- 8
vs
.
4-6 in
H
.
brasiliensis
and
H
.
intermedius
, always
5 in
H
.
australis
and
4 in
H
.
curupira
). It can be further distinguished from
H
.
brasiliensis
and
H
.
curupira
, in the number of scales along lateral line (43-48
vs
. 38-43 and 34-39 respectively).
Description.
Morphometric data presented in
Table 7
. Body cylindrical, deeper than wide. Greatest body depth at vertical through fifth scale anterior to dorsal-fin origin in specimens smaller than
70 mm
SL, closer to dorsal-fin origin in larger specimens. Anterior profile of head slightly rounded in lateral view, more angular in specimens smaller than
160 mm
SL. Dorsal profile of head almost straight. Dorsal margin of orbit at horizontal through of dorsal profile of head in specimens smaller than
100 mm
SL but not reaching dorsal profile of head in larger specimens. Dorsal profile of body slightly convex from vertical through first series of scale of body to dorsal-fin origin; straight and posteroventrally inclined along dorsal-fin base; straight and less inclined to slightly concave from vertical through base of last dorsal-fin ray to origin of dorsal most procurrent caudal-fin ray. Latter portion slightly more concave in specimens smaller than
100 mm
SL. Ventral profile of lower jaw distinctly angular in region of mandibular symphysis, straight to slightly inclined from vertical through anterior nostril to posterior margin of lower jaw. Medial margins of contralateral dentaries running in parallel (
Fig. 1
ac). Ventral profile of body slightly convex to pelvic-fin origin; straight to approximately convex from latter point to anal-fin origin; straight and posterodorsally inclined along anal-fin base; straight to slightly concave from last anal-fin ray to anterior most ventral procurrent caudal-fin ray.
Upper jaw slightly shorter than lower jaw, more so in larger specimens (
ca
.
380 mm
SL). Posterior portion of maxilla dorsally enlarged and extending medially to anterior margins of second and third infraorbitals. Upper and lower lips fleshy with short skin projections covering canines externally. Anterior nostril tubular with anterior slit along its distal half. Anterior and posterior nostrils situated along horizontal through center of orbit, anterior nostril located at approximate one orbital diameter from anterior margin of orbit; posterior nostril midway between anterior nostril and anterior margin of orbit. Eye proportionately larger in smaller specimens. Infraorbital bones well developed and horizontally elongate. Preopercle bearing inumerous laterosensory system pores and partially covered by posterior margin of infraorbitals 3, 4, and posteroventral portion of infraorbital 5. Anteroventral margin of infraorbital 3 relatively straight and posteroventral margin convex. Posterior margin of infraorbital 4 relatively straight and margins of infraorbitals 5 and 6 slightly convex. Small specimens (
ca
.
70 mm
SL) with infraorbital 3 barely reaching orbital rim and proximal ends of infraorbitals 2 and
4 in
contact. Larger specimens (
ca
.
160 mm
SL) with infraorbital 4 barely reaching orbital rim and proximal ends of infraorbitals 2 and 5 almost in contact. Specimens larger than
380 mm
SL with infraorbital 5 progressively separated from rim. Proximal ends of infraorbitals 2 and 6 almost meeting in some of largest specimens examined (
ca
.
750 mm
SL).
Teeth in both jaws conical or canine. Premaxillary teeth in single row. First premaxillary tooth large canine, and fifth tooth medium sized. Sixth tooth canine and almost as large as anterior most premaxillary canine. Second to fourth, seventh and eighth premaxillary teeth small. Maxilla with single row of approximately 38 relatively small teeth, except for very well developed canine-like fourth tooth. Dentary with anterior external row of teeth and posterior internal row. External series with three anterior small teeth, followed by two well developed canines, with posterior canine larger than anterior canine and then ten conical teeth, slightly smaller than anterior most dentary canine. Internal series beginning at level of last conical tooth of external row and composed of approximately 15 very small teeth. Accessory ectopterygoid and ectopterygoid toothed. Ectopterygoid with series of small conical teeth along its ventrolateral margin and many smaller viliform teeth medially on its ventral surface. Endopterygoid edentulous.
Distal margins of all fins rounded. Total dorsal-fin rays 12-15 (n = 90; ii,10-13; mode: ii,11, n = 66). Dorsal fin located at midbody, its origin at vertical through approximately fourth scale anterior on series along pelvic-fin origin. Longest dorsalfin ray approximately three-quarters of body depth in specimens smaller than
160 mm
SL and almost half of body depth in larger specimens. Anal-fin base short. Total analfin rays 9-12 (n = 89; ii,7-10, mode: ii,8, n = 56). Tip of depressed dorsal fin reaching vertical through anal-fin origin in smaller specimens (<
160 mm
SL), but falling short of vertical through that point in larger specimens. Total pectoralfin rays 11-15 (n = 89; i,10-14; mode: i,12, n = 35). Pectoral-fin origin located at vertical through central portion of opercle. Tip of pectoral fin separated from pelvic-fin origin by five to nine scales. Pectoral and pelvic fins of similar size, slightly larger than anal fin in specimens larger than
380 mm
SL. Pelvic-fin rays i,7 (n = 86). Pelvic-fin origin situated at midbody approximately four scales posterior to vertical through dorsal-fin origin.
Tips
of pelvic fin separated from vertical through anal-fin origin by five to seven scales. Caudal-fin rays i,15,i (n = 15).
Well developed cycloid scales imbricated along body. Dorsal scales begin in series at posterior margin of parietals and overlap supraoccipital spine. Last vertical series of scales on caudal peduncle forms almost straight line on caudal-fin base in specimens smaller than
160 mm
SL, and slightly convex arch on caudal-fin in larger specimens. Anterior margin of scales with small recess and posterior margin rounded. Eight to eleven
radii
extending from center of scale to its anterior margin and around eighteen
radii
, some anastomosed, extending from center of scale to its dorsal, posterior and ventral margins. Lateral line straight and complete, extending from posteroventral margin of supracleithrum to posterior most scale in body. Lateral line with 43-48 (n = 89, 43 in
one specimen
; mode: 46, n = 28) perforated scales (
Table 2
). Lateral-line scales with single laterosensory canal. Longitudinal series of scales between lateral line and dorsal-fin origin 4.5-6.5 (n = 88; mode: 5.5, n = 61). Longitudinal series of scales between lateral line and pelvic-fin origin 4.5-6.5 (n = 86; mode: 4.5, n = 63). Longitudinal series of scales around caudal peduncle 18-22 (n = 88; mode: 20, n = 73).
Approximately 11 gill rakers on first epibranchial, most in form of small denticulated plates. Lower branch of first branchial arch with five to seven more elongate rakers and approximately 12 plate-like rakers (n = 18). Laterosensory canal along ventral surface of dentary with 6-8 pores (n = 90; mode: 7, n = 67) (
Table 3
).
Color in alcohol.
Ground coloration of head and body dark to light brown, darker dorsally and paler ventrally. Ventral region white to light yellow in most specimens. Single dark midlateral stripe along lateral line stripe more evident in specimens smaller than
100 mm
SL. Midlateral stripe crossed by five to seven dark bars, each three to five scales wide. Diagonal bars in many specimens continuing posteroventrally approximately two or three longitudinal series of scales ventral of lateral line, resulting in chevron-like pattern more conspicuous in specimens smaller than
160 mm
SL. Dorsal portions of bars more evident than ventral region. Ground coloration of dorsal surface of head, infraorbital region and opercular series dark brown, with dark reticulated pattern on dorsal surface of head in some specimens. Lips with alternating dark and light vertical bands in specimens smaller than
160 mm
SL and completely dark in larger specimens. Coloration of infraorbital region similar to dorsal surface of head. Many specimens with one or two dark stripes extending radially and posteriorly from eye across opercular series and third stripe extending posteroventrally from eye across posterior region of maxilla. Ground coloration of opercular series dark brown. Opercular membrane usually lighter than opercle.
All fins light brown (lighter than body) with dark spots on rays and interradial membranes forming pattern of irregular dark stripes. Stripes of dorsal and caudal fins wider than those of anal fin. Stripes less regular in dorsal fin. Ventral surfaces of pectoral and pelvic fins either lighter than or with same pattern as dorsal surface, but with pattern less conspicuous.
Distribution.
Known from the type-locality and its vicinity, rio Ribeira de Iguape basin, and from the rio
Uruguay
basin, including the
río Negro
in
Uruguay
(
Fig. 3
).
Remarks.
Hoplias lacerdae
has been regarded as having a wide distribution, occurring in the rio Ribeira de Iguape as well as in rivers of the Amazon, Paraná, São Francisco,
Uruguay
and other basins. Results presented herein, however, demonstrate that specimens found in the rio Ribeira are morphologically different from those of remaining cited basins with the exception of rio
Uruguay
. The distribution of
Hoplias lacerdae
is therefore restricted to rio Ribeira de Iguape and rio
Uruguay
, being in sympatry with
Hoplias australis
in the latter basin.
Although samples from the rio Ribeira and rio
Uruguay
did not differ significantly in the meristic and morphometric features examined in the present study, some geographical disparities should be pointed out. Most specimens from the rio Ribeira have 6.5 and 5.5 scales above and below lateral line, respectively, while the majority of specimens from the rio
Uruguay
have 5.5 and 4.5 scales above and below lateral line respectively. Specimens from the rio Ribeira tend to have fewer lateral line scales than do specimens from the rio
Uruguay
(modes: 44 and 46, respectively) and more pectoral-fin rays than do samples from the rio
Uruguay
(i,13-14
vs
. i,10-13, respectively). Nevertheless, these subtle differences must be qualified by the fact that they are not absolute and specimens of
H
.
lacerdae
from the rio Ribeira are rare in collections (only
ten specimens
were available for this study). More specimens from rio Ribeira and the use of cytogenetics and molecular approaches may determine whether these two populations constitute different species.
Fig. 11.
Hoplias lacerdae
, holotype, MNRJ 211, 750.0 mm SL, rio Ribeira de Iguape, São Paulo State, Brazil.
Bertolletti
et al
. (1990)
and
Pessano
et al
. (2005)
cited
H
.
lacerdae
from the rio
Uruguay
basin, the former in the area of Garabi Hydroelectric Station and the latter in Arroio Quarai- Chico. These authors, however, did not mention the basis of those identifications, nor did they illustrate their material. Therefore, it is impossible to determine whether they were dealing with
Hoplias lacerdae
or the new species,
H
.
australis
, described herein, both of which occur in the rio
Uruguay
basin.
A few specimens of
Hoplias lacerdae
from Ernestina reservoir in the rio Jacuí, a river running into the Atlantic Ocean in
Rio Grande do Sul
, southern
Brazil
, were examined.
Malabarba (1989: 127)
and Carlos Lucena (pers. comm.) mentioned that the species was introduced from rio
Uruguay
to rio Jacuí system in the 1980s. Native occurrence of
Hoplias lacerdae
is therefore considered restricted to the rio
Uruguay
and rio Ribeira de Iguape drainages.
Zaniboni Filho
et al
. (2004)
reported some biological features of this species in the rio
Uruguay
including length, weight, and growth curve. According to them, the species has fractional spawning and an economic importance in local fisheries and aquacultures.
In the rio Ribeira de Iguape, the species is regarded as rare by local fisherman and a two-year survey of its ichthyofauna failed to yield specimens of this species (
Oyakawa
et al
., 2006
). The small number of specimens of
Hoplias lacerdae
from rio Ribeira in ichthyological collections may be additional evidence that this species is uncommon, requiring further studies on its ecology and biology and special attention to its conservation status.