Description of a new species of Moenkhausia (Characiformes: Characidae) from the upper Paraguay basin, Central Brazil, with comments on its phylogenetic relationships
Author
Britzke, Ricardo
Author
Troy, Waldo P.
Author
Oliveira, Claudio
Author
Benine, Ricardo C.
text
Neotropical Ichthyology
2018
2018-07-16
16
2
1
11
journal article
10.1590/1982-0224-20170086
086501d9-a2f5-418d-8b5e-368bf4c45b3f
1982-0224
3709537
Moenkhausia flava
,
new species
Figs. 1-7
;
Tab. 1
urn:lsid:zoobank.org:act:
FC435ADB-E0BC-4C44-B707- 5B5F4852FC79
Holotype
.
MZUSP 123719
, 34.0 mm SL,
Brazil
:
Mato Grosso State
,
Tangará da Serra
,
Córrego São Jorge
, tributary of
rio Sepotuba
, rio
Paraguay
basin,
14°27’24.9” S
57°34’32.7” W
,
12 Nov 2009
,
R
.
Britzke
,
T
.
S. Zanini
&
W. P. Troy.
Paratypes
.
LBP 9031
,
17
(7 cs),
20.9-36.8 mm
SL, collected with holotype
.
LBP 18414
,
15
,
29-38.3 mm
SL, same locality as holotype,
2 Apr 2013
, W. P. Troy
.
MZUSP 123720
,
10
,
24.9-35.2 mm
SL, same locality as holotype,
2 Apr 2013
, W. P. Troy
.
MZUEL 8139
,
3
,
19.9-23.5 mm
SL,
Brazil
:
Mato Grosso State
,
Tangará da Serra
,
Ribeirão do Sapo
, tributary of
rio Sepotuba
, rio
Paraguay
basin,
14°33’24.6” S
57°48’45.8” W
,
29 Aug 2013
,
J. L. O. Birindelli
,
A. Claro-García
,
F. Assega
&
E. Santana
.
Fig. 1.
Moenkhausia flava
, holotype, MZUSP 123719, 34.0 mm SL, Brazil, Mato Grosso, small stream tributary of rio Sepotuba.
Fig. 2.
Moenkhausia flava
, paratype, LBP 18414, 34.1 mm SL. Overall coloration darker, yellowish tan, where most of dark pigments are preserved.
Non types.
Brazil
:
Mato Grosso State
,
Tangará
da Serra
: LBP 8418, 2,
34.9-39.4 mm
SL, Córrego São Jorge, tributary of
rio Sepotuba
, rio
Paraguay
basin,
14°27’26.3” S
57°34’34” W
.
LBP 8406, 1,
23.4 mm
SL, Riacho Águas Claras, tributary of
rio Sepotuba
, rio
Paraguay
basin,
14°21’03.2” S
57°33’07.2” W
. Non-types also include dissected, alcoholfixed, and poorly preserved specimens
.
Diagnosis.
Moenkhausia flava
is distinguished from all congeners and members of closely related genera, except
Hemigrammus barrigonae
Eigenmann & Henn, 1914
;
Hemigrammus lunatus
Durbin, 1918
;
Hemigrammus machadoi
Ota, Lima & Pavanelli, 2014
;
Hemigrammus ulreyi
(Boulenger, 1895)
;
Moenkhausia collettii
(Steindachner, 1882)
;
Moenkhausia conspicua
Soares & Bührnheim, 2016
;
Moenkhausia copei
(Steindachner, 1882)
and
Moenkhausia venerei
Petrolli, Azevedo-Santos & Benine, 2016
, by the presence of a well-defined dark line at the base of the anal fin.
Moenkhausia flava
can be easily distinguished from
H. barrigonae
,
H. ulreyi
,
M. conspicua
and
M. venerei
by the presence of a midlateral dark thin stripe little evident on posterior half of the body (
vs
. conspicuous dark and well-defined longitudinal midlateral dark stripe from humeral region until caudal peduncle).
Moenkhausia flava
is quite similar with
H. barrigonae
and
M. conspicua
by sharing overall body shape and similar color pattern. In addition, some specimens of
H. barrigonae
also present complete lateral line (see
Géry, 1977:503
;
Soares, Bührnheim, 2016: 398
), similar to the new species and
M. conspicua
.
Moenkhausia flava
differs from
H. barrigonae
and
M. conspicua
by presenting inner having premaxillary and dentary teeth pentacuspid (
vs
. inner premaxillary and dentary teeth heptacuspid in both species). Additionally, differs from
H. barrigonae
by larger body depth (31.0-42.6% SL, mean = 35.8% SL
vs
. 30.4- 34.8% SL, mean = 32% SL); and differs of
M. conspicua
by the dorsal-fin length (24.3-31.3%
vs
. 31.2-36.8% SL, respectively), anal-fin length (14.8-22.1%
vs
. 22.6-27.1% SL, respectively) and orbital diameter (34.8-43.4%
vs
. 44.9-56.0% HL, respectively).
Moenkhausia flava
can be easily distinguished from
H. machadoi
and
H. lunatus
by an inconspicuous oval-shaped vertically elongated humeral blotch, extending horizontally from third through five lateralline scales, and vertically from third row above lateral line to first row below it (
vs
. a conspicuous vertically elongated dark humeral blotch, extending horizontally from second through sixth lateral-line scales, and vertically from third row above lateral line to first row below it in
H. machadoi
; and a small roundish humeral blotch in
H. lunatus
). The new species differs from
M. collettii
and
M. copei
by having six longitudinal rows of scales above lateral line (
vs
. five longitudinal rows of scales). Additionally, it is distinguished from
M. copei
by a greater number of branched analfin rays (20-23
vs
.
15-17 in
M. copei
) and distinguished from
H. machadoi
,
H. lunatus
and
M. collettii
by a threescale deep band of sparse, scattered dark chromatophores extending along midlateral body (
vs
. a line of concentrated chromatophores along midlateral body).
Tab. 1.
Morphometrics data of holotype and paratypes of
Moenkhausia
flava
. Standard length (SL) is expressed in mm, all other measurements are expressed as percentages of SL, except for subunits of head that are expressed as percentages of head length (HL). N=46.
| Holotype |
Paratypes Range Mean |
| Standard length (mm) |
34.0 |
19.9 - 38.3 |
27.8 |
| Percentage of standard length |
| Greatest depth |
38.7 |
31.0 - 42.6 |
35.8 |
| Snout to dorsal-fin origin |
55.5 |
48.3 - 56.3 |
54.2 |
| Snout to pectoral-fin origin |
29.9 |
28.8 - 33.7 |
30.7 |
| Snout to pelvic-fin origin |
49.7 |
45.0 - 51.8 |
49.3 |
| Snout to anal-fin origin |
65.6 |
58.9 - 69.2 |
64.4 |
| Caudal peduncle depth |
10.7 |
06.7 - 10.7 |
9.2 |
| Caudal peduncle length |
07.5 |
05.9 - 10.0 |
7.5 |
| Pectoral-fin length |
19.9 |
15.3 - 20.8 |
18.7 |
| Pelvic-fin length |
17.4 |
14.6 - 18.0 |
16.6 |
| Dorsal-fin length |
29.7 |
24.3 - 31.3 |
28.9 |
| Dorsal-fin base |
13.7 |
10.7 - 14.6 |
13.1 |
| Anal-fin length |
19.1 |
14.8 - 22.1 |
19.2 |
| Anal-fin base |
32.6 |
25.2 - 33.7 |
30.9 |
| Eye to dorsal-fin origin |
41.1 |
35.8 - 43.3 |
38.7 |
| Dorsal-fin origin to caudal-fin origin |
39.3 |
33.0 - 40.8 |
39.4 |
| Head length |
26.3 |
25.3 - 30.1 |
27.2 |
| Head depth |
19.4 |
19.3 - 22.9 |
20.7 |
| Percentage of head length |
| Snout length |
24.6 |
20.3 - 28.4 |
24.2 |
| Upper jaw length |
36.6 |
31.7 - 47.0 |
37.2 |
| Horizontal orbital diameter |
37.8 |
34.8 - 43.4 |
39.4 |
| Least interorbital width |
34.3 |
30.0 - 45.6 |
35.1 |
Description.
Morphometric data for
Moenkhausia flava
summarized in
Tab. 1
. Body relatively compressed and elongated, moderately high. Greatest body depth just before the origin of dorsal fin. Snout profile convex. Dorsal profile of head straight or slightly convex. Dorsal profile of body convex from posterior tip of supraoccipital spine to dorsal-fin origin, slightly convex and posteroventrally slanted along dorsal-fin base, slightly convex from posterior terminus of dorsal-fin base to end of adipose-fin origin, and concave along caudal peduncle. Ventral body profile convex from tip of lower jaw to caudal-peduncle origin, slightly concave along caudal peduncle. Prepelvic region transversally flattened, more so proximal to pelvicfin insertion. Postpelvic region transversally flattened proximal to pelvic-fin insertion, becoming somewhat obtuse toward anal-fin origin. Mouth terminal. Maxilla slightly beyond vertical through anterior margin of orbit. Premaxillary teeth in two rows; outer teeth row with 3(19), 4*(21) tricuspid teeth, midcentral cusps longer than remaining cusps; inner teeth row with 4(2), 5*(38) tetra- to pentacuspid teeth, midcentral cusps longer than remaining cusps. Maxilla with 2(21), 3*(19) tricuspid teeth. Dentary with 4(5), 5*(27), 6(8) pentacuspid teeth, followed by a series of small teeth, with 1-3 cusps (
Fig. 6
). Dorsal profile of head convex from upper lip to vertical through anterior nostril. Suparaoccipital process short, its tip not reaching vertical through posterior margin of opercle. Dorsal-fin rays ii,9(30). Pectoral-fin rays i,10(30), i,11*(10). Tip of pectoral fin not reaching vertical through pelvic-fin insertion. Adipose fin well developed. Pelvic-fin rays i,7(30), when adpressed, its tip reaching first anal-fin ray in a few specimens. Anal-fin rays iv, 20(3), 21*(10), 22(12), 23(3). Caudal fin forked. Principal caudal-fin rays i,17,i. Scales cycloid, with few radii along posterior border. Lateral-line completely pored. Lateral line scales 33(4), 34(17), 35*(11), 36(1). Scale rows between dorsal-fin origin and lateral-line 6; scale rows between lateral-line and pelvic-fin origin 4. Circumpeduncular scale rows 14*(32). Predorsal scales 10*(30). Scale sheath along anal-fin base in a single series of 4 scales, extending from the first to fifth branched anal-fin ray. First gill arch with 9*(18) gill rakers on ventral limb and 5*(18) on dorsal limb.
Sexual dimorphism.
Adult males with small hooks on the last unbranched and anterior four branched anal-fin rays. Also, found on the first and second pelvic-fin rays branched. In both fins, there are four to seven hooks per fin ray, located on the distal segments.
Color in alcohol.
Humeral region with an inconspicuous oval-shaped vertically elongated humeral blotch located on second to fourth lateral line scales, extending from 3 horizontal series of scales above lateral to the series of scales immediately below the lateral line. Dark chromatophores scattered on infraorbitals and opercle, longitudinal dark lighter stripe along on the eye (
Fig. 2
). A dark thin stripe extending along horizontal septum, from humeral region to caudal peduncle, more evident on posterior half of the body. A three-scale deep band of sparse, scattered dark chromatophores extending along midlateral body.
Dorsal fin rays with few dispersed chromatophores, more concentrated on anterior half. Anal-fin rays with few dispersed chromatophores, more concentrated along its proximal and distal extension, resulting in a lighter medial area. Tip of anterior anal-fin rays densely pigmented by dark chromatophores resulting in a dark dash in this area. Paired fins hyaline with scattered dark pigmentation, more concentrated on unbranched rays. Caudal fin with a narrow field of dark chromatophores on its distal margin. Conspicuous dark line at anal-fin base. All fins hyaline with few dispersed chromatophores. Anterior rays of the anal fin and base of caudal-fin lobe present orange pigmentation in freshly preserved specimens.
Color in life.
Body general color pattern pale yellowish. Dorsal region olive. Abdomen whitish to light yellow. Pelvic fin and adipose fin with yellow-orangish coloration. Dorsal fin and caudal fin with orange-reddish coloration. Anal fin with first rays with orange-reddish coloration and the remaining rays with hyaline coloration. A conspicuous dark line at the base of the anal fin. Pectoral fin yellowish (
Fig. 4
).
Fig. 3.
Type-locality of
Moenkhausia flava
, upper rio Sepotuba basin, Mato Grosso, Brazil.
Ecological notes.
All specimens of
Moenkhausia flava
were collected along the margin in semi-lentic stretches of a shallow river with clear water, with sand and scattered small stones on the bottom, aquatic vegetation (
Echinodorus
sp.,
Eleocharis
sp.,
Hygrophila
sp. and
Mayaca fluviatilis
), and riparian vegetation (
Fig. 3
). Species collected syntopically were
Hyphessobrycon hebertaxelrodi
Géry, 1961
,
Hyphessobrycon vilmae
,
Pyrrhulina australis
Eigenmann & Kennedy, 1903
,
Characidium
aff. z
ebra
Eigenmann, 1909,
Corydoras aeneus
(Gill, 1858)
,
Hypostomus
sp.,
Brachyhypopomus
sp.,
Gymnotus inaequilabiatus
(Valenciennes, 1839)
, and
Aequidens rondoni
(
Miranda
Ribeiro, 1918).
Geographic distribution.
Only known from tributaries of the upper rio Sepotuba, above the waterfalls Salto das Nuvens and Salto Maciel, rio
Paraguay
basin,
Mato Grosso State
,
Brazil
(
Fig. 5
).
Etymology.
The name of species is derived from Latin
flavus
, meaning yellow. The name refers to the yellowish color pattern of the body in live specimens. An adjective.
Fig. 4.
Live specimen of
Moenkhausia flava
, paratype, MZUEL 8139, 28.5 mm SL.
Fig. 5.
Map showing the localities of
Moenkhausia flava
. Red star represents the type locality. Black square represents the Salto das Nuvens fall and the white square represents Salto Maciel fall.
Phylogenetic analysis.
Partial sequences of two mitochondrial (16SrRNA and Cytb) and three nuclear genes (Myh6, Rag1 and Rag2) were obtained for 2 species of
Hemigrammus
and 29 species of
Moenkhausia
from the final matrix by
Mariguela
et al.
(2013)
deposited in TreeBase (treebase.org) under access number 13922, plus the species
Hemigrammus barrigonae
,
H. unilineatus
and
Moenkhausia flava
. The combined sequence data resulted in a matrix with 4,640 base pairs (bp), out of which 1,911 were conserved, 2,726 were variable and 2,196 were information parsimony. The estimated index of substitution saturation (Iss) performed in DAMBE 5.2.31 (
Xia, Xie, 2001
) showed that the data were not saturated (
i.e
., Iss.c value greater than Iss). In our analysis,
Moenkhausia flava
is more closely related to
Moenkhausia collettii
,
M. copei
,
Hemigrammus barrigonae
and
H. ulreyi
(
Fig. 7
). The new species was recovered within the clade 2 (
sensu
Mariguela
et al.
, 2013
), also composed of
Aphyodite grammica
Eigenmann, 1912
;
Bryconella pallidifrons
(Fowler, 1946)
;
Hemigrammus barrigonae
,
H. ulreyi
,
H. unilineatus
(Gill, 1858)
;
Moenkhausia comma
Eigenmann, 1908
;
M. copei
,
M
. aff.
copei
,
M
.
collettii
,
M. hemigrammoides
Géry, 1965
;
M. margitae
Zarske & Géry, 2001
;
Pristella maxilaris
(Ulrey, 1894)
, plus
Moenkhausia flava
. In this analysis, the groups proposed by
Géry (1977)
, reinforce that overall similarities are not sufficient to distinguish groups in
Moenkhausia
. However, few features seem to be still useful to designate natural groups within
Moenkhausia
and
Hemigrammus
(see discussion).
Fig. 6.
Moenkhausia flava
. LBP 9031, 31.9 mm SL, paratype: right premaxilla (top), maxilla (middle), and dentary (bottom). Scale bar = 0.5 mm.
Fig. 7.
Maximum likelihood phylogenetic tree based on Partial sequences of two mitochondrial genes and three nuclear genes. The series of three numbers (e.g. 100/94/1) at each of the main nodes represents the percentage of bootstrap support obtained by Maximum Likelihood (ML), the percentage of bootstrap support obtained by the Maximum Parsimony (MP) analysis and the posterior probability for that split obtained in Bayesian analysis (B),respectively (1000 bootstrap replicates). Dashes represent values <50% (ML, MP) or <0.5 (B).
Conservation status.
Moenkhausia flava
is known exclusively from tributaries of the upper rio Sepotuba, above the waterfalls Salto das Nuvens and Salto Maciel, rio
Paraguay
basin. Because no specific threats have been detected, the species can be categorized as Least Concern (LC) according to IUCN criteria (IUCN, 2016), however, we emphasize the apparently narrowly distribution of the species, and strongly encourage the preservation of those environments.