Received: 10 May 2019
Accepted: 6 July 2019
DOI: 10.1111/jfb.14091
FISH
REGULAR PAPER
A new species of Hemigrammus (Characiformes: Characidae)
from western Amazon River basin, Colombia
Juan G. Albornoz-Garzón1 | Alejandro Méndez-López1 | Carlos DoNascimiento1
|
2
Flávio C. T. Lima
1
Colecciones Biológicas, Instituto de
Investigación de Recursos Biológicos
Alexander von Humboldt, Boyacá, Colombia
2
Museu de Zoologia da Universidade Estadual
de Campinas “Adão José Cardoso”, São Paulo,
Brazil
Correspondence
Carlos DoNascimiento, Colecciones Biológicas,
Instituto de Investigación de Recursos
Biológicos Alexander von Humboldt, Carrera
8, No. 15-08, Villa de Leyva, Boyacá,
Colombia.
Email: cdonascimiento@humboldt.org.co
Abstract
A new species of Hemigrammus is described from the Amazon Basin near Leticia,
Departamento Amazonas, Colombia. In common with some congeners and some
Hyphessobrycon spp., the new species colour pattern lacks a humeral blotch but has a
caudal-peduncle blotch. It can be distinguished from congeners with a similar colour
pattern by having: a relatively deep body (30.2%–39.0% LS), 20–24 branched anal-fin
rays, 6–8 perforated scales of the lateral line, anal-fin base without a conspicuous
black stripe, 3–5 maxillary teeth, maxillary teeth with 1–3 cusps, a thin longitudinal
midlateral line and a well-defined oval shaped caudal blotch, extending from caudal
peduncle into the lower caudal-fin rays. Comparisons with congeners and with
Funding information
FAPESP, Grant/Award Number:
2011/51532-7 and 2013/20936-0
Hyphessobrycon species sharing the same general colour pattern are presented.
KEYWORDS
freshwater fishes, Hemigrammus tridens group, Río Amazonas, Stethaprioninae, taxonomy
1 | I N T RO D U C T I O N
Colombia. This new species is similar to some congeners assigned by
Géry (1977) to the Hemigrammus tridens group by presenting a colour
The genus Hemigrammus Gill 1858 currently includes 61 valid species
pattern that includes a conspicuous caudal peduncle blotch but lacks a
and is widely distributed throughout the major cis-Andean river basins
humeral blotch. A description of the new species is given here.
across South America (Lima et al., 2016; Ota et al., 2019; Reia & Benine, 2019). Traditionally, Hemigrammus species have been defined by
a combination of non-exclusive characters, first advanced by Durbin
(in Eigenmann, 1918): incomplete lateral line, teeth in the premaxilla
arranged in two rows, inner premaxillary tooth row composed by five
teeth, few or none maxillary teeth and scales extending to the basal
region of caudal-fin lobes.
Recent phylogenies of Characidae using either morphological
(Mirande, 2010), or molecular data (Javonillo et al., 2010; Oliveira
2 | MATERIALS AND METHODS
Counts and measurements follow Fink and Weitzman (1974), except
for the number of horizontal scale rows below the lateral line (counted
to the pelvic-fin insertion) and the number of post-anal scales (median
scales between the posterior rim of the anus and the base of the first
anal-fin ray) and the addition of three measurements: postorbital head
et al., 2011) have failed to demonstrate the monophyly of the genus.
length (measured from posterior bony orbital margin to posterior bony
More recently, Mirande (2018), using a total-evidence approach, also
margin of opercle), dorsal-fin base length and anal-fin base length
failed to recover the genus as monophyletic. There are, however,
(measured between the bases of the anterior and posteriormost rays
some propositions of monophyletic groups within the genus (Lima
of each fin). All measurements were taken with digital callipers to the
et al., 2016; Ota et al., 2019).
nearest 0.1 mm on the left side of specimens. Measurements are
Recently, a new species of Hemigrammus was identified during
expressed as percentage of standard length (LS), except subunits of
searches in the IAvH-P collection, from the western Amazon Basin in
the head, which are expressed as percentage of head length (LH).
932
© 2019 The Fisheries Society of the British Isles
wileyonlinelibrary.com/journal/jfb
J Fish Biol. 2019;95:932–939.
ALBORNOZ-GARZÓN ET AL.
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933
Counts are followed by their absolute frequency in parentheses.
Asterisks indicate counts of the holotype. Circuli and radii were counted on scales from the row immediately dorsal to the lateral line at the
vertical through the dorsal-fin origin. Counts of supra-neurals,
branchiostegal rays, gill-rakers of first branchial arch, teeth cusps,
diminutive posteriormost dentary teeth, unbranched anal-fin rays,
procurrent caudal-fin rays, uroneurals and position of pterygiophores
were taken from cleared and stained (c&s) specimens prepared
according to Taylor and Van Dyke (1985). Vertebrae of the Weberian
apparatus were counted as four elements and the compound caudal
centrum (PU1 + U1) as a single element. Precaudal vertebral counts
include the Weberian apparatus and the vertebrae lacking haemal
spines. Caudal vertebral counts include all vertebrae with haemal
spines. Catalogue numbers are followed by the total number of specimens and their LS range. The number of c&s specimens is given in
parentheses, followed by their respective LS range. Institutional abbreviations follow Sabaj (2019). Morphometric data for Hemigrammus
boesemani Géry 1959 are based on the original description by
Géry (1959).
F I G U R E 2 Hemigrammus amacayacu sp. nov. paratypes,
Colombia, Amazonas, Amazon Basin, Leticia, Parque Nacional Natural
Amacayacu, unnamed tributary of quebrada Matamatá: a IAvH-P
8285, 24.9 mm LS, b IAvH-P 8283, 24.4 mm LS
3 | RESULTS
3.1 | Hemigrammus amacayacu sp. nov. (Figures 1–3)
urn:lsid:zoobank.org:act:F7BBFFA6-312D-46B7-BD68-D7BEE1CD4083.
Hemigrammus levis (not Durbin): Arbeláez et al., 2008: 514 (abundance, quebrada Matamatá, Departamento Amazonas, Colombia).
3.1.1 | Holotype
IAvH-P 21889, 23.2 mm LS , Colombia, Amazonas, Leticia, Parque
Nacional Natural Amacayacu, unnamed tributary of quebrada
Matamatá, 03 47 0 53.3” S, 70 15’ 58” W, F. Arbeláez,
20 Mar 2006.
3.1.2 | Paratypes
All from Colombia, Amazonas, Amazon Basin, Leticia, Parque Nacional
Natural Amacayacu, unnamed tributary of quebrada Matamatá. IAvHP 8282, 12, 17.9–26.3 mm LS, collected with the holotype. IAvH-P
F I G U R E 3 Hemigrammus amacayacu sp. nov. IAvH-P 8283,
paratype, 24.5 mm LS, medial view of left jaws: a maxilla, b premaxilla,
c dentary
8280, 4, 23.1–25.2 mm LS, same locality as holotype, F. Arbeláez,
F I G U R E 1 Hemigrammus amacayacu sp. nov. holotype, IAvH-P
21889, 23.2 mm LS, Colombia, Amazonas, Amazon Basin, Leticia,
Parque Nacional Natural Amacayacu, unnamed tributary of quebrada
Matamatá
17 Mar 2006. IAvH-P 8281, 18, 21.6–26.4 mm LS (1 c&s, 24.0 mm LS),
same locality as holotype, F. Arbeláez, 16 Mar 2006. ANSP 206738,
2, 24.0–24.8 mm LS; IAvH-P 8283, 14, 12.0–24.5 mm LS (2 c&s,
934
ALBORNOZ-GARZÓN ET AL.
FISH
24.2–24.5 mm LS); ZUEC 17117, 2, 23.1–23.8 mm LS, same locality as
dorsal-fin origin. Dorsal profile of head slightly convex from tip of
holotype, F. Arbeláez, 18 Mar 2006. IAvH-P 8284, 1, 19.7 mm LS, 03
snout to vertical through anterior margin of orbit; straight from latter
480 23” S, 70 15’ 59.8” W, F. Arbeláez, 15 Mar 2006. IAvH-P 8285,
point to end of supraoccipital spine; convex from that point to dorsal-
1, 25.2 mm LS, same locality as IAvH-P 8284, F. Arbeláez,
fin origin. Dorsal-fin base slightly convex and posteroventrally
13 Mar 2006.
inclined; straight and slanted from dorsal-fin terminus to adipose-fin
origin; slightly concave to straight from that point to anteriormost
3.1.3 | Diagnosis
dorsal procurrent caudal-fin ray. Ventral profile convex from anterior
tip of dentary to pelvic-fin origin, straight and dorsally slanted from
Hemigrammus amacayacu sp. nov. differs from most congeners (except
from H. boesemani, Hemigrammus brevis Ellis 1911, Hemigrammus
mahnerti Uj & Géry 1989, Hemigrammus parana Marinho, Carvalho,
Langeani & Tatsumi 2008, Hemigrammus rodwayi Durbin 1909 and
Hemigrammus tridens Eigenmann 1907) by having a conspicuous
caudal-peduncle blotch and humeral blotch lacking (v. either humeral
blotch or blotches present, caudal-peduncle blotch present, or presence of some other colour feature as dark pigmentation on caudal-fin
that point to anal-fin origin. Ventral profile along anal-fin base straight
and posterodorsally inclined. Ventral profile of caudal peduncle
slightly concave.
Mouth terminal. Posterior tip of maxilla crossing vertical through
anterior margin of pupil. Premaxillary teeth in two rows: outer row
with 2* (31) or 3 (9), uni to tricuspid teeth; inner row with 5 (40) tri to
pentacuspid teeth. Maxilla with 3 (12), 4* (20) or 5 (7) teeth,
lobes, or a conspicuous midlateral dark stripe). Hemigrammus
amacayacu differs from H. boesemani by having a deeper body
(2.6–3.1 times in LS v. 3.1–3.6 times), anal-fin base without a conspic-
T A B L E 1 Morphometric data for holotype and paratypes of
Hemigrammus amacayacu sp. nov. (n = 37)
uous black stripe (v. anal-fin base with a conspicuous black stripe) and
Paratypes
caudal blotch roughly oval in shape (v. caudal blotch roughly rhomboiHolotype
Mean
Range
SD
23.2
23.6
19.7–26.4
1.4
Depth at dorsal-fin
origin
39.0
34.6
30.2–38.1
2.1
Snout to dorsal-fin
origin
53.4
51.5
48.3–54.9
1.8
Snout to pectoral-fin
origin
27.4
28.2
25.0–30.8
1.4
Snout to pelvic-fin
origin
46.8
46.4
43.9–48.4
1.2
extending from caudal peduncle into lower caudal-fin rays (v. roughly
Snout to anal-fin origin
62.6
59.9
56.7–63.3
1.5
triangular or rectangular, restricted to caudal fin, not extending into
Caudal-peduncle depth
11.3
10.4
7.7–12.1
1.1
lower caudal-fin rays) and largest dentary teeth tricuspid (v. pen-
Caudal-peduncle length
11.9
12.5
9.8–15.2
1.2
tacuspid). Hemigrammus amacayacu can be distinguished from
Pectoral-fin length
20.4
19.8
17.1–24.6
1.9
dal in shape). Hemigrammus amacayacu can be distinguished from
H. brevis by presenting a broad, roughly oval caudal blotch (v. caudal
Standard Length (LS, mm)
blotch as an elongated midlateral stripe) and 20–24 branched anal-fin
Percentage of LS
rays (v. 17–18 branched anal-fin rays). Hemigrammus amacayacu can
be distinguished from H. mahnerti by narrow line above anal-fin base
absent (v. present), darkened area on sides of body immediately above
urogenital opening absent (v. present), maxillary teeth uni to tricuspid
(v. pentacuspid), larger dentary teeth tricuspid (v. penta to
heptacuspid). Hemigrammus amacayacu can be distinguished from
H. parana by having a thin longitudinal midlateral line (v. relatively
broad longitudinal midlateral band) and an oval shaped caudal blotch,
H. rodwayi by a lower number of lateral-line scales (6–8 v. 9–22),
Pelvic-fin length
16.3
16.5
15.0–23.0
1.7
larger dentary teeth tricuspid (v. pentacuspid) and by having a well-
Dorsal-fin length
31.9
30.0
26.4–33.3
1.4
defined caudal-peduncle blotch, slightly asymmetrical, with pigmenta-
Dorsal-fin base length
14.4
14.3
11.7–16.3
0.8
53.8
54.9
50.8–57.7
1.4
relatively diffuse, roughly symmetrical, dark pigmentation at caudal fin
Dorsal-fin origin to
caudal-fin origin
restricted to middle caudal-fin rays). Hemigrammus amacayacu can be
Anal-fin base
30.2
29.9
25.8–33.2
1.9
distinguished from H. tridens by presenting maxillary teeth uni to tri-
Anal-fin length
21.8
20.2
17.2–22.6
1.5
37.6
37.3
35.0–39.9
1.1
fin blotch, without an extension along middle caudal-fin rays (v. caudal
Posterior margin of eye
to dorsal-fin origin
peduncle blotch with posterior portion elongated along middle
Head length (LH)
27.7
26.6
25.2–28.6
0.9
Postorbital head length
35.7
37.1
32.2–43.1
2.6
Snout length
21.3
23.4
19.9–27.9
1.8
Least interorbital width
34.4
32.6
27.7–36.6
2.3
Upper jaw length
44.2
44.7
41.3–50.8
2.1
Eye diameter
36.1
37.9
35.3–44.4
1.9
tion extending into lower caudal-fin rays (v. caudal-peduncle blotch
cuspid (v. pentacuspid), by presenting a roughly oval-shaped caudal-
caudal-fin rays) and by the higher number of branched anal-fin rays
(20–24 v. 15–17).
3.1.4 | Description
Morphometric data of holotype and paratypes are given in Table 1.
Body compressed and moderately deep; greatest body depth at
Percentage of LH
ALBORNOZ-GARZÓN ET AL.
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935
anteriormost one or two teeth tricuspid and remaining posterior teeth
darkened area present in humeral region due to thinning of hypaxial
conic, five from 15 examined specimens with all maxillary teeth conic.
musculature. A longitudinal band of chromatophores originating
Dentary with 11 (1), 12–13 (1) or 13–15 (1) teeth. Anteriormost four
immediately posterior to opercle, becoming more slender posteriorly
teeth larger, mostly tricuspid (except one specimen with first three
and with few chromatophores reaching caudal peduncle blotch. Lat-
teeth on right side and second teeth on left side tetracuspid),
eral septum with dark chromatophores forming a thin longitudinal
remaining teeth unicuspid, gradually decreasing in size posteriorly
midlateral line, beginning slightly anterior to vertical through middle of
(Figure 3). Central cusp of multicuspid teeth oriented slightly back-
pectoral-fins, becoming more darkly intense posteriorly and extending
wards. First gill arch with 1 (1) or 2 (2) gill rakers on hypobranchial,
to caudal peduncle blotch. Dark chromatophores aligned with margins
8 (2) or 9 (1) on ceratobranchial, 1 (1) on cartilage between
of myomeres, starting slightly anterior to anal-fin origin. Abdominal
ceratobranchial and epibranchial and 5 (2) or 7 (1) on epibranchial.
region light brown, without chromatophores. Caudal peduncle with
Total number of gill rakers on first gill arch 15 to 17. Branchiostegal
large, conspicuous oval shaped dark blotch, slightly asymmetrical,
rays four (3).
occupying most of posterior half of caudal peduncle and extending
Scales cycloid, with three to five strongly marked radii, diverging
from focus to posterior portion of scale; circuli well-marked anteriorly.
Lateral line incomplete, with 6 (13), 7* (8) or 8 (2) pored scales. Longitudinal series of scales including lateral-line scales 30 (1), 31* (4),
32 (5), or 33 (3). Scale rows between dorsal-fin origin and lateral line
along middle caudal-fin rays and ventrally to base of lower caudal-fin
rays. Remaining fins with dark chromatophores mainly scattered over
inter-radial membrane. Dorsal and anal fins with chromatophores concentrated along distal half. Adipose fin without chromatophores.
5 (19). Scale rows between lateral line and pelvic-fin insertion 3 (1) or
4* (20). Scale rows between lateral line and anal-fin insertion three*
(15) or four (2). Predorsal scale row with 9 (5), 10* (24) or 11 (6).
Circumpeduncular scales 10 (2), 11 (3) or 12* (13). Postanal scales 2*
(23) or 3 (5). Caudal fin with one to four scales on basal one-third of
each lobe, similar in size to adjacent body scales.
Pectoral-fin rays i,9 (1), 10* (22), 11 (13) or 12 (1). Tip of pectoral
fin reaching pelvic-fin insertion. Pelvic-fin rays i,6 (2) or 7*(35). Tip of
pelvic fin reaching anal-fin origin in some specimens. Dorsal-fin rays
3.1.6 | Distribution
Hemigrammus amacayacu sp. nov. is only known from two localities of
a short unnamed tributary of the quebrada Matamatá. The quebrada
Matamatá is a forest stream draining an upland area of the Parque
Nacional Natural Amacayacu and is a direct tributary of the main
channel of the Amazon River, departamento Amazonas, Colombia
(Figure 4).
ii,9* (37); first unbranched ray about one-half length of second
unbranched ray; second unbranched and first branched rays longest.
Dorsal-fin origin posterior to vertical through pelvic-fin origin; base of
last dorsal-fin ray slightly posterior to vertical through anal-fin origin.
Anteriormost dorsal-fin pterygiophore inserted posterior to neural
spine of 9th (3) vertebra. Anal-fin rays iv* (3), 20 (2), 21 (9), 22* (11),
23 (13) or 24 (2). Anteriormost anal-fin pterygiophore inserted posterior to neural spine of 15th (3) vertebra. Anal fin with anterior lobe
formed by last unbranched and five anteriormost branched-fin rays,
remaining rays decreasing gradually in size. Adipose fin present. Caudal fin forked, lobes similar in size. Principal caudal-fin rays i,10 + 9,i.
3.1.7 | Ecological notes
The localities where the specimens of Hemigrammus amacayacu
sp. nov. were found are streams draining pristine terra firma forest.
Some physical and chemical variables from these localities are: temperature 24.7–24.9 C, dissolved oxygen 3.7–4.0 mg O2 l−1,
pH 5.7–5.9, conductivity 13.5–15.3 μS cm−1, phosphates 0.08 mg l−1
and total bases 0.13–0.15 mg l−1 (Arbeláez et al., 2008). The gut content of one cleared and stained specimen (IAvH-P 8281, 24.0 mm LS)
was filled with insect remains, from which several ants and a small
Dorsal caudal-fin procurrent rays 8 (2); ventral caudal-fin procurrent
cockroach could be identified. Hemigrammus amacayacu was collected
rays 6 (2) or 8 (1); one pair of uroneurals. Supraneurals 4 (3). Vertebrae
with Characidium sp., Hoplias malabaricus (Bloch 1794), Serrasalmus
33 (3); precaudal vertebrae 13 (1) or 14 (2), caudal vertebrae
sp., Cyphocharax pantostictos Vari & Barriga Salazar 1990, Copella vil-
19 (2) or 20 (1).
mae Géry 1963, Nannostomus marginatus Eigenmann 1909, Carnegiella
strigata (Günther 1864), Gasteropelecus sternicla (L. 1758), Ace-
3.1.5 | Colour in alcohol
strorhynchus falcatus (Bloch 1794), Astyanax anterior Eigenmann 1908,
Chrysobrycon mojicai Vanegas-Ríos & Urbano-Bonilla, 2017, Charax
Overall body colour uniformly light brown. Small, dark chromato-
tectifer (Cope 1870), Gymnocorymbus thayeri Eigenmann 1908, Knodus
phores densely concentrated on dorsal surface of head, premaxilla,
sp., Moenkhausia comma Eigenmann 1908, Moenkhausia oligolepis
anterior portion of maxilla and dentary. Infraorbitals 3–6, opercle and
(Günther 1864), Tyttocharax cochui (Ladiges 1949) and Bryconops inpai
preopercle with scattered dark chromatophores. Ventral portion of
Knöppel, Junk & Géry 1968.
head clear, except by presence of scattered dark chromatophores
restricted to anteroventral region of dentary. Dark chromatophores
densely concentrated on predorsal area. Scales from laterodorsal por-
3.1.8 | Sexual dimorphism
tion of body with reticulated pattern formed by dark chromatophores
No secondary sexually dimorphic characters were observed in exam-
concentrated along scale borders. Humeral blotch absent, but
ined specimens of Hemigrammus amacayacu sp. nov. However, it
936
ALBORNOZ-GARZÓN ET AL.
FISH
72°
71°
70°
69°
N
3°
Putumayo River
F I G U R E 4 Map of southern
portion of Colombia showing type
locality
of Hemigrammus
amacayacu sp.nov. The symbol
represent more than one lot and
locality
4°
r
ive
va
Ya
ri R
Am
o
Ri
r
ve
Ita
qu
ai
Ri
ve
r
5°
az
s
na
50
0
50
100
150
200 km
should be noted that all specimens were collected in the same month
498–499) as lacking a humeral blotch, actually possess a relatively
(March 2006).
inconspicuous humeral blotch (Lima et al., 2013, pp. 260–263, for pictures of specimens of H. hyanuary and H. levis). A comparison with
3.1.9 | Etymology
these species is relevant given the fact that they are superficially similar to H. amacayacu and indeed, the species was mistakenly identified
The specific name amacayacu comes from the Parque Nacional Natu-
as H. levis by Arbeláez et al. (2008). In addition to the lack of a humeral
ral Amacayacu where the specimens of this species were found. The
blotch, H. amacayacu differs from H. hyanuary and H. levis by having a
word amacayacu is a quechua word meaning el río de las hamacas (i.e.,
deeper body (32.7%–39.0% LS v. 24.4%–29.6% LS and 22.9%–29.8%
river of the hammocks). The Parque Nacional Natural Amacayacu is an
LS, respectively), a higher number of branched anal-fin rays (20–24 v.
important national park because represents around 40% of the area
11–14 and 16–18, respectively). Hemigrammus amacayacu further dif-
known in Colombia as the Amazonian trapeze, conserving vast areas
fers from H. levis by a fewer number of cusps in the largest teeth of
of both terra firme and seasonally flooded forests. A noun in
inner row of premaxilla and dentary (3–5 v. 7–9) and is distinguished
apposition.
from H. micropterus by the lower number of perforated scales of the
lateral line (6–8 v. 8–12), largest tooth of inner row of premaxilla pen-
4 | DISCUSSION
tacuspid (v. heptacuspid), number of maxillary teeth and cusps (3–5
Hemigrammus amacayacu sp. nov. shares with H. boesemani, H. brevis,
blotch oval shaped, slightly asymmetrical, with pigmentation exten-
H. mahnerti, H. micropterus, H. parana, H. rodwayi and H. tridens the
ding into lower caudal-fin rays (v. caudal-peduncle blotch roughly
lack of a humeral blotch and the presence of a caudal-peduncle
rhomboidal in shape, extending into middle caudal-fin rays).
conical or tricuspid teeth v. 1 heptacuspid tooth) and caudal-peduncle
blotch. Some of these species (the ones already known at the time,
Faria et al. (2018) discussed that some Hyphessobrycon species
H. boesemani, H. brevis, H. micropterus, H. rodwayi and H. tridens) were
(namely, Hyphessobrycon diastatos Dagosta, Marinho & Camelier
included by Géry (1977, pp. 495) in his “He. tridens-group”, which was
2014, Hyphessobrycon juruna Faria, Lima & Bastos 2018, Hyphe-
defined by the combination of these characters. There is no evidence
ssobrycon negodagua Lima & Gerhard 2001 and Hyphessobrycon par-
that this group is a monophyletic unit and as other species groups pro-
vellus Ellis 1911) closely resembles He. rodwayi by sharing a similar
posed by Géry (1977) for species-rich characiform genera, was a mere
colour pattern; i.e., the lack of a humeral blotch and the presence of a
convenient tool for aiding species identifications rather than a propo-
diffuse caudal peduncle blotch. The traditional diagnosis between
sition of a possible monophyletic group. Among the species included
Hyphessobrycon Durbin 1908 and Hemigrammus is based on the
by Géry (1977) in the “He. tridens-group”, Hemigrammus hyanuary
absence of scales on the caudal peduncle (v. its presence in the latter
Durbin 1918, Hemigrammus levis Durbin 1908 and Hemigrammus
genus),
micropterus Meek 1907, although reported both by Durbin
Hemigrammus spp. possess just a few scales at the caudal-fin base,
(in Eigenmann, 1918, pp. 136–137) and Géry (1977, pp. 495,
often missing in preserved specimens. Hemigrammus amacayacu is
an
unreliable
diagnostic
character,
given
that
most
ALBORNOZ-GARZÓN ET AL.
937
FISH
distinguished from Hy. diastatos by presenting caudal peduncle blotch
possible that the occurrence of H. amacayacu may be connected with
asymmetrical, extending into lower portion of caudal peduncle and
the nutrient-richer water from streams draining the Pebas formation.
base of lower caudal-fin rays (v. caudal-peduncle blotch lying at the
Hemigrammus amacayacu is another recent discovery from the
centre of caudal peduncle and base of middle caudal-fin rays), dorsal
Leticia area, a relatively well-sampled region in the Amazon Basin for
fin not elongated and anal-fin lobe only moderately developed (v. dor-
fishes, but which yielded some recently described fish species (Lima
sal fin and anal-fin lobe considerably elongated in mature males) and a
et al., 2016; Valdesalici, 2016; Vanega-Ríos & Urbano-Bonilla, 2017;
higher number of branched anal-fin rays (20–24 v. 15–18).
DoNascimiento et al., 2019). These recent discoveries emphasizes the
Hemigrammus amacayacu can be distinguished from Hy. juruna,
need of additional, broader ichthyological surveys in the Amazonian
H. negodagua and H. parvellus by presenting a well-defined caudal
Trapeze of Colombia.
peduncle blotch (v. caudal-peduncle blotch blurred, with ill-defined
contours) and larger dentary teeth with only three cusps (v. larger
5 | ADDITIONAL SPECIMENS EXAMINED
dentary teeth with five cusps in H. negodagua and H. parvellus and
with 7–9 cusps in H. juruna). It can be additionally distinguished from
5.1 | Hemigrammus boesemani
H. negodagua by possessing 20–24 branched anal-fin rays (v. 14–17).
The area surrounding Leticia was intensely surveyed ichthyologi-
MHNG 2180.009, 10, 25.1–30.4 mm LS, no locality data. MHNG
cally by Galvis et al. (2006), who recorded seven Hemigrammus spp.
2180.010, 2, 24.0–25.4 mm LS, Suriname, Coropina Creek. MHNG
from the area; i.e., Hemigrammus analis Durbin 1909, Hemigrammus
2180.011, 6, 24.5–30.8 mm LS, Suriname, Marowijne. MHNG
bellottii (Steindachner 1882), Hemigrammus cf. erythrozonus Durbin
2180.012, 3, 33.4–36.7 mm LS, no locality data. MHNG 2180.016,
1909, Hemigrammus luelingi Géry 1964, Hemigrammus ocellifer
(Steindachner 1882), Hemigrammus pulcher Ladiges 1938 and
Hemigrammus schmardae (Steindachner 1882). However, as noticed
respectively by Lima et al. (2013), p. 233) and Lima et al. (2016,
pp. 124–125), the record of H. analis in Galvis et al. (2006) actually
refers to Bryconella pallidifrons (Fowler 1946), while the record of
H. ocellifer refers instead to Hemigrammus aguaruna Lima, Correa &
1,
27.1
mm
LS,
Suriname,
Marowijne.
MHNG
2180.015,
7, 21.4–25.5 mm LS, Suriname, Paramaribo. MHNG 2181.077,
4, 21.0–28.4 mm LS, French Guyana, Cayenne. MHNG 2181.078,
2, 27.1–28.9 mm LS, French Guyana, Maroni River. MHNG 2616.053,
7, 19.6–25.3 mm LS, French Guyana, Sinnamary River. NRM 70353,
5, 25.7–29.8 mm LS, French Guyana, Kourou. NRM 70684,
1, 26.5 mm LS, French Guyana, Sinnamary Basin.
Ota 2016. Additionally, the record of Hemigrammus cf. erythrozonus
by Galvis et al. (2006) is a misidentification, the examination of the
voucher material at ICN-MHN by us showed it to belong to an unidentified Hemigrammus species. Hemigrammus erythrozonus, the popular glowlight tetra of aquarists, is only known from some tributaries of
5.2 | Hemigrammus brevis
ZUEC 13494, 6, 19.0–26.7 mm LS, Brazil, Sergipe, Rio São Francisco
basin.
the Essequibo River in Guyana and Venezuela (F.C.T.L., unpublished
data). Among the sympatric Hemigrammus spp., H. amacayacu is more
similar to H. schmardae, with which it shares a conspicuous caudalpeduncle blotch. However, H. schmardae possess a conspicuous, narrow, vertically elongated humeral blotch, while H. amacayacu lacks
any hint of the presence of a humeral blotch and for presenting a
5.3 | Hemigrammus hyanuary
ZUEC 8879, 28, 24.1–31.0 mm LS, Brazil, Pará, Santarém, Rio Tapajós
drainage. ZUEC 13813, 141, 25.4–31.5 mm LS, Brazil, Amazonas,
Guajará, Rio Juruá drainage. ZUEC 15314, 20, 31.2–36.5 mm LS, Brazil, Amazonas, Tefé, Rio Tefé.
lower number of branched anal-fin rays (15–20 v. 20–24, respectively), among other differences.
Galvis et al. (2006) probably did not have access to any specimen
5.4 | Hemigrammus levis
of Hemigrammus amacayacu because they have examined relatively
ZUEC 7971, 30, 39.1–47.8 mm LS, Brazil, Amazonas, Barcelos, Rio
little material from the Parque Nacional Natural Amacayacu. Most of
Negro drainage. ZUEC 8310, 136, 24.6–41.3 mm LS, Brazil, Amazonas,
the material used in their study from upland streams was collected in
Iranduba, lago Janauari. ZUEC 11813, 23, 23.5–47.4 mm LS, Brazil,
the quebradas crossing the road to Taracua, to the north-west of the
Pará,
town of Leticia (Galvis et al., 2006, p. 74). The quebrada Matamatá
62, 25.0–40.3 mm LS, Brazil, Amapá, Calçoene, Rio Amapá Grande.
drains the Pebas formation, while streams to the east sampled by
ZUEC 16746, 166, 23.4–35.4 mm LS, Brazil, Amapá, Tartarugalzinho,
Galvis et al. (2006) drain the Terciario Superior Amazónico formation,
Rio Araguari drainage.
Santarém,
Rio
Arapiuns
drainage.
ZUEC
16529,
which presents more leached and coarser soils than soils from the
Pebas formation (Arbeláez et al., 2008). Arbeláez et al. (2008) reported
considerable differences in fish composition between streams draining
5.5 | Hemigrammus mahnerti
the Pebas formation v. streams draining the Terciario Superior
ZUEC 3216, 1, 17.0 mm LS; ZUEC 5476, 1, 20.4 mm LS, Brazil, Mato
Amazónico formation, with the first presenting a considerable larger
Grosso, Poconé, Rio Bento Gomes. ZUEC 6656, 48, 15.1–19.0 mm LS,
biomass and species richness when compared with the latter. It is
Brazil, Mato Grosso, Poconé, Rio Piraputanga.
938
ALBORNOZ-GARZÓN ET AL.
FISH
5.6 | Hemigrammus micropterus
FUNDING INF ORMATI ON
IAvH-P 9509, 592, 12.5–34.2 mm LS, Colombia, Casanare, Aguazul,
Financial support was given by FAPESP, project “Systematics of the
laguna El Tinije, río Meta drainage. IAvH-P 20232, 1, 33.6 mm LS,
tetras (genera Hemigrammus, Hyphessobrycon, Thayeria, Parapristella
Colombia, Vichada, Cumaribo, caño Fruta, Caño Matavén drainage,
and Bryconella), with emphasis on the species from cis-Andean South
Orinoco River basin. IAvH-P 20250, 72, 8.7–34.5 mm LS, Colombia,
America”, grants # 2011/51532-7 and 2013/20936-0.
Meta, Puerto Gaitán, laguna Mururito, río Meta drainage. ZUEC
16918, 50, 20.3–28.1 mm LS, Colombia, Meta, río Meta drainage, OriOR CID
noco River basin.
Carlos DoNascimiento
https://orcid.org/0000-0002-8680-7942
5.7 | Hemigrammus parana
ZUEC 7505, 10, 24.8–29.8 mm LS, Brazil, Minas Gerais, Conceição
das Alagoas, Rio Grande drainage.
5.8 | Hemigrammus rodwayi
ZUEC 12885, 85, 20.2–34.0 mm LS, Brazil, Pará, Primavera, Rio Primavera drainage. ZUEC 14437, 14, 26.1–40.5 mm LS, Brazil, Pará,
Igarapé
Açú,
Rio
Maracanã
drainage.
ZUEC
16062,
100, 22.2–32.3 mm LS, Brazil, Amapá, Porto Grande, Rio Araguari
drainage.
5.9 | Hemigrammus schmardae
CIACOL 2050, 13, 18.5–28.5 mm LS, Colombia, Amazonas, Amazon
River basin. IAvH-P 9024, 22, 24.7–30.1 mm LS, Colombia, Amazonas,
Leticia, quebrada Sufragio, Amazon River basin. ZUEC 15091,
38, 21.0–31.1 mm LS, Brazil, Amazonas, Tabatinga, Rio Solimões
drainage. ZUEC 15149, 61, 26.0–33.1 mm LS, Brazil, Amazonas, Tefé,
Rio Tefé drainage. ZUEC 16989, 733, 12.7–26.6 mm LS, Brazil, Amazonas, Tabatinga, Rio Solimões drainage.
5.10 | Hemigrammus tridens
ZUEC 6683, 1, 17.8 mm LS, Brazil, Mato Grosso, Poconé, Rio
Piraputanga. UMMZ 206594, 147, 15.2–28.1 mm LS, Paraguay, San
Pedro, Río Aguaray-Mi.
ACKNOWLEDGEMEN TS
We thank Jorge E. García-Melo and Eduardo G. Baena for the photographs of the paratypes (Figures 2a and 2b, respectively). The fourth
author (FCTL) is grateful to A. Acosta-Santos and E. Agudelo Cordoba
(CIACOL), J. I. Mojica, H. D. Agudelo-Zamora and G. Serrato (ICNMHN), D. W. Nelson (UMMZ), M. H. Sabaj, J. G. Lundberg and
M. Arce Hernández (ANSP), R. Covain (MHNG), S. O. Kullander,
B. Delling and E. Ahlander (NRM) for access to comparative material
used in this study.
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How to cite this article: Albornoz-Garzón JG, MéndezLópez A, DoNascimiento C, Lima FCT. A new species of
Hemigrammus (Characiformes: Characidae) from western
Amazon River basin, Colombia. J Fish Biol. 2019;95:932–939.
https://doi.org/10.1111/jfb.14091