A revision of the cis-andean species of the genus Brycon Müller & Troschel (Characiformes: Characidae)
Author
Lima, Flávio C. T.
text
Zootaxa
2017
4222
1
1
189
journal article
37268
10.5281/zenodo.257769
9ea59a17-588e-4af1-8c0d-ebcd50ad0395
1175-5326
257769
F0EC0A87-B1EE-4B5C-8F53-77A7EEA75F3A
Brycon dulcis
Lima & Vieira new species
(
Figs. 30–31
)
Brycon
sp.:
Travassos & Freitas, 1948
: 628
(Espírito Santo, Lagoa de Juparanã, Brazil: parasites).
Brycon devillei
(not Castelnau): Amaral-Campos, 1950: 140 (Rio Doce, Espírito Santo, Brazil); Vieira
et al.
, 2008: 47–48 (part: rio Doce system, Espírito Santo and Minas Gerais, Brazil; ecology, distribution, conservation);
Travenzoli
et al.
, 2015
: 6, 8–11, 13–14 (Lagoa Carioca, rio Doce, Minas Gerais; cytogenetics, phylogenetics relationships, molecular taxonomy).
Brycon
cf.
devillei
: Sunaga & Verani, 1991: 360
, 361 (abundance, Lago Dom Helvécio, Lagoa Carioca, rio Doce system, Minas Gerais, Brazil);
Godinho & Formagio, 1992
: 95, 98 (Lago Dom Helvécio, Rio Doce State Park, Minas Gerais);
Godinho
et al.
, 1994
: 80, 82 (Lago Dom Helvécio, Lago Ferrugem and Lagoa Carioca, Rio Doce State Park, Minas Gerais); Gomes
et al.
, 2007: 1–2 (Lagoa Carioca, rio rio Doce system, Minas Gerais, Brazil).
Diagnosis.
Brycon dulcis
can be diagnosed from all remaining cis-andean
Brycon
species, with the exception of
B. stolzmanni
,
B. coxeyi
,
B. coquenani
,
B. vermelha
,
B. insignis
,
B. howesi
,
B. ferox
,
B. vonoi
,
B. opalinus
, and
B. nattereri
by possessing a color pattern consisting in a humeral blotch and a caudal peduncle blotch, without body stripes or other obvious color markings on caudal and anal-fins (vs. body stripes and caudal/anal fin color markings present; see
Fig. 5
).
Brycon dulcis
can be distinguished from
B. stolzmanni
,
B. coxeyi
,
B. vonoi
,
B. opalinus
, and
B. nattereri
by possessing a distinctly acute head profile (vs. a roughly rounded to slightly acute head profile).
Brycon dulcis
can be additionally distinguished from
B. stolzmanni
and
B. coxeyi
by the absence of a patch of dark pigmentation on the opercle (vs. dark patch of pigmentation present on opercle).
Brycon dulcis
can be distinguished from
B. insignis
,
B. howesi
,
B. coquenani
, and
B. vermelha
by possessing a fifth infraorbital bone about as wide as high (vs. fifth infraorbital bone wider than high; see
Fig. 6
).
Brycon dulcis
can be distinguished from
B. ferox
by presenting an approximately isognathous mouth, with premaxillary and dentary mostly overlapping, leaving only part of outer premaxillary series exposed in ventral view in some specimens (vs. mouth distinctly anisognathous, premaxillary pointed, extending beyond dentary, leaving outer series, and often also the second, series of premaxillary teeth exposed in ventral view).
Brycon dulcis
can be additionally diagnosed from other
Brycon
species occurring in the rio Doce basin,
B. opalinus
, by possessing a higher number of anal-fin rays (22–27, modally 25, vs. 17–23, modally 20). See “Remarks”, for additional notes on the recognition of the species.
Description.
Morphometric data are presented in
Table 9
. Large-sized species, largest examined specimen 372.0 mm SL. Body slender to moderately slender. Largest body height slightly ahead of dorsal-fin origin. Dorsal body profile slightly convex from upper lip margin to vertical through anterior naris, slightly concave to straight from latter point to basis of supraoccipital process, moderately convex from latter point to dorsal-fin origin, straight along dorsal-fin basis, and straight to slightly convex from dorsal-fin basis to adipose-fin origin. Dorsal profile of caudal peduncle slightly concave. Ventral profile slightly convex from lower lip to pelvic-fin insertion, straight to slightly convex from this point to anal-fin origin and approximately straight along anal-fin base. Ventral profile of caudal peduncle slightly concave.
Head profile moderately acute anteriorly, considerably acute in specimens <
140 mm
SL, mouth terminal. Jaws isognathous to slightly anisognathous, premaxillary projecting slightly relative to dentary in some specimens, leaving outer row of premaxillary teeth exposed when mouth is closed. Maxillary long, extending posteriorly to slightly posterior to middle of pupil. Adipose eyelid well developed. Premaxillary teeth in three rows; teeth of third row largest. Six (3), 7 (5), 8 (8), 9 (8), 10 (11), 11 (7), or 12 (3) tricuspidate teeth in outer series. Three (3), 4 (21), 5 (13), or 6 (4) tetra- to pentacuspidate teeth in second, inner premaxillary row, plus 1 (1), 2 (17), 3 (18), or 4 (2) tricuspidate teeth between the first and third rows. Two teeth in third premaxillary row, medial teeth largest, symphyseal teeth smaller, slightly tilted towards each other, both hexacuspidate. Maxillary with distal portion distinctly expanded and rounded in profile. Eighteen to 29 maxillary teeth, slightly smaller than teeth of first premaxillary row, anterior teeth tricuspidate, posterior teeth unicuspidate. Dentary with 9 (1), 10 (5), 11 (2), 12 (7), 13 (7), 14 (5), 15 (5), 16 (5), or 17 (2) teeth in main series. Anterior four dentary teeth assymetrical, considerably larger and bulkier than remaining teeth, pentacuspidate, each with central cusp distinctly larger than remaining cusps. Remaining dentary teeth progressivelly smaller, penta-, tetra-, tri- to unicuspidate. Inner (lingual) series consisting of a small, single unicuspid symphyseal tooth, situated immediately posterior to symphyseal dentary teeth of main series, plus row of 37–44 small, unicuspidate teeth, originating on lingual crest of dentary replacement trench slightly after the symphysis. Inner symphyseal teeth present in all specimens with relatively intact symphyseal dentaries area, except for MZUSP 42627 (2, 127.0–
138.7 mm
SL), where symphyseal teeth are missing.
Central
cusp distinctly larger and pointed, and teeth presenting diastemas in specimens <
140 mm
SL.
TABLE 9.
Morphometric data of
Brycon dulcis
(A: holotype, MZUSP 58911).
A |
n |
Range |
Mean |
Standard length (SL) |
267.0 |
45 |
74.6–372.0 |
- |
Percentages of standard length |
Depth at dorsal-fin origin |
31.6 |
37 |
25.5–32.7 |
28.6 |
Snout to dorsal-fin origin |
51.3 |
45 |
48.8–56.8 |
51.6 |
Dorsal-fin base length |
11.5 |
45 |
9.6–13.2 |
11.5 |
Posterior terminus of dorsal fin to adipose fin |
25.3 |
45 |
18.6–26.5 |
22.8 |
Posterior terminus of dorsal fin to hypural joint |
37.8 |
45 |
31.6–40.1 |
35.2 |
Snout to pelvic-fin insertion |
46.1 |
40 |
44.0–52.5 |
46.5 |
Snout to anal-fin origin |
65.2 |
45 |
56.8–73.1 |
65.0 |
Anal-fin base length |
26.4 |
45 |
22.7–28.0 |
24.4 |
Caudal peduncle length |
15.8 |
45 |
11.0–16.2 |
13.4 |
Dorsal-fin height |
19.6 |
45 |
16.9–21.8 |
19.6 |
Pectoral-fin length |
21.0 |
45 |
17.0–21.7 |
18.8 |
Pelvic-fin length |
15.6 |
45 |
13.4–16.9 |
15.1 |
Caudal peduncle depth |
9.5 |
45 |
7.6–9.7 |
8.5 |
Head length |
24.2 |
45 |
23.3–33.6 |
26.4 |
Percentages of head length |
Head height |
81.6 |
45 |
62.4–85.5 |
72.4 |
Snout length |
27.2 |
44 |
27.2–33.4 |
29.5 |
Upper jaw length |
44.0 |
45 |
44.0–53.3 |
47.6 |
Horizontal eye diameter |
25.5 |
45 |
16.9–28.5 |
22.8 |
Post-orbital length |
49.7 |
45 |
42.3–54.7 |
47.0 |
Least interorbital width |
36.7 |
45 |
24.7–40.8 |
32.4 |
Scales cycloid. Lateral line complete, from supracleithrum to caudal-fin base. Forty-eight (3), 49 (1), 50 (1), 51 (9), 52 (6), 53 (8), 54 (6), 55 (5), 56 (1), 57 (3), 58 (1), 60 (2), or 64 (1) scales in lateral line series. Laterosensory tube simple in small (<
180 mm
SL) specimens, deflected upwards in the first 5–6 scales, downwards in the remaining lateral-line scales. Larger specimens (>
180 mm
SL) with branched tubules, mostly bifurcated in middle-sized specimens but some scales with 3–5 tubules. Horizontal scale rows between dorsal-fin origin and lateral line 8 (3), 9 (27), 10 (14), or 11 (3). Horizontal scale rows between lateral line and pelvic-fin 4 (12), 5 (25), or 6 (10). Circumpeduncular scales 16 (2), 17 (7), 18 (17), 19 (11), 20 (5), 22 (1), or 23 (2).
Dorsal-fin rays ii, 9. Dorsal fin origin slightly ahead of middle of SL. First dorsal-fin pterygiophore inserting behind neural spine of 13th (1), 14th (1), or 15th (1) vertebrae. Anal-fin rays iii (not including first, small unbranched ray only visible in the cs specimen), 22 (4), 23 (7), 24 (11), 25 (19), 26 (4), or 27 (2). First anal-fin pterygiophore inserting behind haemal spine of 23th (1) or 24th (2) vertebrae. Last unbranched and anterior 3–4 branched anal-fin rays longer, remaining rays progressively shorter towards anal-fin end. Anal fin displaying numerous (c. 20–30 per fin-ray main branch) middle-sized hooks on last unbranched and posterior main branch of branched rays 13–22, associated with dense, gelatinous tissue in 10 specimens (MZUSP 17070, 265.1–
270.1 mm
SL; MZUSP 36663, 225.7–
237.9 mm
SL; MZUSP 58911, 267.0 mm SL; MZUSP 106770, 285.0–323.0 mm SL; MCZ
20031, 236.5 mm
SL; NMW 62947, 337.0–372.0 mm SL). Some fin rays with hooks also arranged along anterior ray margin. A single hook per ray segment, except when hooks also present on anterior ray margin. Sheath of scales covering basis of anal-fin rays composed of two scale rows, lower scale row formed by 20–22 rectangular scales. Pectoral-fin rays i, 11 (2), 12 (8), 13 (24), 14 (9), or 15 (2). Pelvic-fin rays i,7. Main caudal-fin rays 10/9. Caudal fin forked, lobes pointed.
Four branchiostegal rays, three on anterior ceratohyal and one on posterior ceratohyal. First branchial arch with 10 (1), 11 (2), 12 (2), 13 (5), 14 (6), or 15 (3) lower, 1 at angle, and 9 (1), 10 (1), 11 (9), 12 (5), 13 (3), or 14 (1) upper gill rakers. Vertebrae 44 (3). Supraneurals 9 (1) or 10 (2).
FIGURE 30.
Brycon dulcis
,
new species
, holotype, MZUSP 58911, 267.0 mm SL: Brazil, Minas Gerais, Lago Dom Helvécio, rio Doce basin.
FIGURE 31.
Brycon dulcis
,
new species
, paratype, ZUEC 6776, 156.0 mm SL: Brazil, Minas Gerais, rio Corrente Grande.
Coloration in alcohol.
Overall body coloration clear to dark. Top of head, snout, supraorbital, sixth infraorbital, and dorsal portion of body light to dark-brown. Second, third, and fourth infraorbitals, and opercle light brown, with a silvery hue in specimens retaining guanine (e.g., MZUSP 58911, NMW 62947, ZUEC 6363). Dentary, maxillary, gular area and lower portion of body cream to light brown. Lateral portion of body lightbrown, silvery in specimens retaining a considerable amount of guanine (e.g., MZUSP 58911, NMW 62947, ZUEC 6363). Humeral blotch present, little conspicuous, approximately rounded in shape, situated immediately above lateral line, its anterior margin at level of second to third, extending longitudinally to posterior margin of fourth to fifth lateral line scales, and vertically less than one scale high. Large, moderately conspicuous, oval-shaped caudal peduncle blotch, extending along 8–10 last lateral-line scales. All rayed fins with a considerable amount of dark pigmentation at the interradial membranes, imparting an overall dark coloration to these fins. Caudal-fin with a poorly discernible, roughly V-shaped blotch, formed by dark pigmentantion situated on outer caudal-fin rays. Adipose fin light-grey to light-brown. Specimens collected in lakes surrounded by forest, as those from the lakes of the middle and lower Rio Doce (e.g., MZUSP 36663, MZUSP 58911, MZUSP 17070), with a dark overall coloration, while specimens collected in tributaries of the rio Doce in more turbid water conditions (e.g., ZUEC 6776, ZUEC 6363) with a considerably clearer overall coloration.
Color in life.
Description based on a photo of a freshly collected specimen, collected at Rio Doce State Park by P. S. Pompeu, two specimens (ZUEC 6363, and ZUEC 6776) collected by T.C. Pessali from rio Corrente Grande, and specimens collected at a tributary of rio Santo Antônio and at the upper rio Doce, Rio Doce county (photos supplied by J. Dergam). Top of head, snout, and dorsum light-grey to dark. Dentary and maxillary clear. Opercle, infraorbital bones, and lateral portion of body silvery, with some dark pigmentation in the specimen from the Rio Doce State Park. Humeral blotch inconspicuous; caudal peduncle blotch moderately conspicuous. Series of pinkish spots situated along the four horizontal scale rows situated below scale row immediately below lateral line and anterior to insertion of pelvic fins in the specimens from Rio Doce State Park, upper Rio Doce and rio Santo Antônio. Pinkish pigmentation concentrated on the basal portion of the scales. All fins with some amount of dark pigmentation. Caudal fin with outer fin-rays dark, forming a roughly V-shaped blotch.
Sexual dimorphism.
As discussed above, anal-fin hooks are present in 10 specimens (MZUSP 17070, 265.1–
270.1 mm
SL; MZUSP 36663, 225.7–
237.9 mm
SL; MZUSP 58911, 267.0 mm SL; MZUSP 106770, 285.0–323.0 mm SL; MCZ
20031, 236.5 mm
SL; NMW 62947, 337.0–372.0 mm SL). Two of them (MZUSP
17070, 270.1 mm
SL and MZUSP
36663, 237.9 mm
SL) were dissected and are males, with poorly developed testicles. On other hand, a single dissected specimen that does not possess anal-fin hooks (MZUSP
36663, 283.1 mm
SL) proved to be a female, with moderately well-developed ovaries.
Etymology.
Dulcis
, after the Latin word for sweet (“doce” in Portuguese), in allusion to the river system from which the species is apparently endemic.
Common names.
“
Piabanha
”.
Distribution
. Only known from the middle and lower rio Doce basins, states of
Minas Gerais
and
Espírito Santo
, eastern
Brazil
(
Fig. 32
). There is a putative historical record for the species for the rio Santa Maria da Vitória, a small coastal drainage situated southward from the rio Doce basin (MZUSP 1748, CAS 11171, CAS 11173). However, there is some uncertainty about the exact locality of these lots (though actually the lots from CAS originated from the lot from MZUSP, and as such refer to a single collection event) since it was recorded in the original MZUSP book catalog as being collected in “
Porto
Cachoeiro, rio Doce”. “
Porto
Cachoeiro” is the old name of Santa Leopoldina, a small town situated at the banks of the rio Santa Maria da Vitória, rather than the rio Doce. The collector, Ernst Garbe, collected also at the rio Doce during the same field expedition (
Pinto, 1945: 17
), so the mixing of contradictory information in the label of this lot does not lend much confidence as to the exact provenance of these specimens. Consequerntly, this locality was not mapped.
Ecological notes.
Brycon dulcis
is recorded as occurring in lake systems associated with the rio Doce, as the Lagoa Juparanã and some other lakes on the lower rio Doce basin and the Lago Dom Helvécio and Lagoa Carioca on the middle rio Doce. Lakes of the middle rio Doce basin, where most the recent records of the species come from, are ancient tributaries of the rio Doce which were dammed either as a consequence of paleoclimatic flutuations or neotectonic activity during the Quaternary (
Suguio & Kohler, 1992
). It is unclear whether populations of
Brycon dulcis
occurring in these lakes are in fact isolated from populations in the rio Doce mainstream or not. Stomach contents of two specimens were examined. One specimen (MZUSP
1531, 146.6 mm
SL) ingested nine
Astyanax
sp. (
Characidae
), one
Geophagus brasiliensis
(Cichlidae)
, one
Poecilia vivipara
(Poeciliidae)
and a
Lepidoptera
larvae. The stomach of the second specimen (MZUSP
36643, 277.8 mm
SL) contained unidentified vegetal matter and crushed insects.
Conservation.
The rio Doce basin is highly disturbed environmentally, water pollution, siltation and deforestation being the main anthropogenic disturbances in this river basin.
Brycon dulcis
was apparently extirparted from the lower portion of the basin, in the Espírito Santo state, where it was relatively common until at least the decade of 1960. Recent records of the species come mostly from the natural lakes system at the middle rio Doce in the Rio Doce State Park (Vieira
et al.
, 2008; Gomes
et al.
, 2007; as
Brycon devillei
). Introduction of two predatory fishes, a species of
Cichla
and the red-bellied piranha,
Pygocentrus nattereri
, have however impacted native fish populations in these isolated lakes (
Godinho & Formagio, 1992
;
Godinho
et al.
, 1994
). The species still persists at the rio Corrente Grande, a tributary of the rio Doce, in the river stretch between Naque and Periquito, where fishermen report that about three specimens are fished per year (T.C. Pessali, pers. comm.), and was also recently recorded at the upper Rio Doce at Rio Doce county and at the mouth of rio Tanque, a tributary of rio Santo Antônio, near Ferros (J. Dergam, pers. comm.,
March 2012
). The species is considered officially threatened in Brazil (Vieira
et al.
, 2008; as
Brycon devillei
). The recent catastrophic burst of a tailings dam at the upper rio Doce basin in
November 2015
, releasing millions of tons of iron waste into the rio Doce, very likely extirpated
Brycon dulcis
from the rio Doce main channel, though the species might still persist in some tributaries and in the lakes of the middle rio Doce.
FIGURE 32.
Map of eastern Brazil, showing known localities of
Brycon dulcis
(red dots; red star indicates type locality),
B. ferox
(black dots), and
B. vonoi
(purple dots; purple star indicates type locality).
Remarks.
Brycon dulcis
was previously misidentified in the literature as
B. devillei
(e.g., Amaral-Campos, 1950; Sunaga & Verani, 1991;
Godinho
et al.
, 1994
; Vieira
et al.
, 2008;
Travenzoli
et al.
, 2015
; see item “Remarks” of
Brycon devillei
and
B. ferox
). As noticed under
Brycon devillei
, this latter name is actually a
species inquirenda
, and more similar to
B. insignis
.
Brycon dulcis
is very similar morphologically to
B. ferox
and
B. vonoi
. Although diagnostic features among these three species are slight and refer only to overall color pattern and head/ snout shape, they are consistent and no gradation among these three morphological
types
was observed. See additional notes on the item “Remarks” of
Brycon vonoi
.
Recently,
Travenzoli
et al.
(2015)
argued in favor for the usage of the name
Brycon devillei
for the
Brycon
species from the rio Doce herein described as
Brycon dulcis
. They argued that the species also occurs at the rio Mucuri, which has its mouth at the southern portion of
Bahia
state, and consequently that the name
B. devillei
could be applied to the species. We have not examined the specimen from the rio Mucuri that
Travenzoli
et al.
(2015)
considered to be conspecific with specimens from the rio Doce and consequently cannot confirm their claim. All
Brycon
specimens from the rio Mucuri examined in the present study were either
B. ferox
or
B. vermelha
(see under these species). At any rate, the name
Brycon devillei
cannot be applied to the
Brycon
species herein named
B. dulcis
because its
holotype
presents a sixth infraorbital bone wider than high (vs. a sixth infraorbital bone as high as wide in
B. dulcis
). See also the item “Remarks” of
Brycon devillei
.
Material examined
.
Holotype
:
MZUSP
58911
(1, 267.0 mm SL):
Brazil
,
Minas Gerais
,
Marliéria
,
Lago Dom Helvécio
,
Rio
Doce State Park,
rio Doce
system,
19°46’S
,
42°36’W
;
A
.L. Godinho &
P.S. Pompeu
,
Sept 1993
.
Paratypes
:
Brazil
,
Minas Gerais
,
rio Doce
basin:
MZUSP
28983
(2, 1 cs,
113.8–223.9 mm
SL):
Marliéria
,
Lagoa Carioca
,
Rio
Doce State Park,
19°45’S
,
42°37’W
;
J.R.Verani
,
27 June–14 July 1983
.
MZUSP
36663
(5, 211.3–
287.9 mm
SL):
Marliéria
,
Lago Dom Helvécio
,
Rio
Doce State Park,
19°46’S
,
42°36’W
;
J.R.Verani
,
27 Nov –13 Dec 1985
.
MZUSP
28968
(
1, 111.9 mm
SL):
Marliéria
,
Lago Dom Helvécio
,
Rio
Doce State Park,
19°46’S
,
42°36’W
;
J.R.Verani
,
25–28
Jun 1983
.
MZUSP
36643
(3, 1 skel.,
147.1–277.8 mm
SL):
Marliéria
,
Lagoa Carioca
,
Rio
Doce State Park,
19°45’S
,
42°37’W
;
J.R. Verani
,
23–25 Nov 1985
. LISDEBE uncat. (7, 245.0–260.0 mm SL):
Marliéria
,
Lago Dom Helvécio
,
Rio
Doce State Park,
19°46’S
,
42°36’W
;
J.G. da Silva
,
March 1977
.
ZUEC
6363
(1, 263.0 mm SL):
Governador Valadares
,
rio Corrente Grande
(trib.
rio Doce
),
18°59’38’’S
,
42°13’33’’W
;
T.C. Pessali
,
May 2010
.
ZUEC
6776
(1, 156.0 mm SL):
Periquito
,
rio Corrente Grande
(marginal lagoon),
18°58’20’’ S
,
42°17’13’’ W
;
T.C. Pessali
,
July 2011
.
Espírito Santo
,
rio Doce
basin:
MNRJ
11248
(4,
87.7–152.6 mm
SL):
Linhares
, lagoa
de Juparanã
, road
Linhares
/
São Mateus
,
19°16’S
,
40°8’W
;
L. Travassos
&
H. Travassos
,
8 March 1948
.
MNRJ
11250
(2, 112.8–
126.9 mm
SL):
Linhares
, lagoa
de Juparanã
,
19°16’S
,
40°8’W
;
L. Travassos
&
H. Travassos
,
29 Feb 1948
.
MZUSP
17070
(3, 265.1–
283.1 mm
SL):
Linhares
,
Lagoa Juparanã
,
19°16’S
,
40°8’W
; H.
A
.Britski & Izáurio Dias,
3–9 Feb 1965
.
MZUSP
3308
(
1, 183.3 mm
SL):
Linhares
,
rio São José
, tributary of
Lagoa Juparanã
,
19°10'S
40°12'W
;
O. Pinto
,
Sept 1942
.
Non
types
.
Minas Gerais
:
MCZ
20031 (3, 194.4–
236.5 mm
SL)
:
Minas Gerais
(?)
,
Rio
Doce (no specific locality);
C.F. Hartt
&
E. Copeland
,
Aug–Sept 1865
.
MZUSP
42627
(2, 127.0–
138.7 mm
SL):
Viçosa
, c.
20°43’S
,
42°54’W
; J. Dergam, no date. MZUSP 106770 (2, 285.0–323.0 mm SL);
MZUSP
106771
(2, 1 skel., 308.0–326.0 mm SL):
Marliéria
,
Lagoa Carioca
,
Rio
Doce State Park,
19°45’26’’S
,
42°37’6’’W
;
E.N. Fragoso
et al.
,
March 2007
–
Sept 2009
.
Espírito Santo
:
MNRJ
11249
(6, 87.0–
104.9 mm
SL):
Linhares
,
Lagoa Juparanã
,,
19°16’S
,
40°8’W
; L. Travassos, H. Travassos & J.T. de Freitas,
8 March 1948
. MNRJ 11251 (
1, 110.7 mm
SL): Linhares, Lagoa de Freitas; L. Travassos & J.T. de Freitas,
Oct 1963
.
MZUSP
1531
(5, 1 cs,
88.7–146.6 mm
SL):
Linhares
,
rio Doce
, c.
19°24’S
,
40°3’W
; E. Garbe, 1906. MCZ 60931 (2, 109.9–
167.4 mm
SL); MCZ 21102 (2, 148.6–
184.4 mm
SL);
MCZ
21101 (1, 357.0 mm SL):
Rio
Doce, between
Linhares
and Aimorés,
19°37’S
,
39°49’W
; C.F. Hartt & E. Copeland, 1865. MZUSP 1748 (7, 1 cs,
74.6–138.1 mm
SL); CAS 11171 (
1, 132.4 mm
SL);
CAS
11753 (
1, 105.1 mm
SL):
Santa Leopoldina
,
rio Santa Maria da Vitória
,
20°6’S
,
40°31’W
;
E. Garbe
, 1906.
Rio Doce, locality not specified:
NMW 69928 (3, 132.9–200.0 mm SL); NMW 62940 (4, 191.5–235.0 mm SL); NMW 62947 (2, 337.0–372.0 mm SL); NMW 62940 (4, 191.5–235.0 mm SL); NMW 62947 (2, 337.0–372.0 mm SL); “rio Doce, Steindachner Don.,
1912–1913
”.