Neotropical Ichthyology, 10(3):519-525, 2012
Copyright © 2012 Sociedade Brasileira de Ictiologia
A new species of Tyttocharax (Characiformes: Characidae: Stevardiinae)
from the Güejar River, Orinoco River Basin, Colombia
César Román-Valencia1, Carlos A. García-Alzate1,3, Raquel I. Ruiz-C.1 and Donald C.
Taphorn B.2
A new Tyttocharax species from the Güejar River system, near the Macarena Mountains in Colombia is described. This is the
first record for the genus from the Orinoco basin. The combination of the following characters distinguish Tyttocharax metae
from its congeners: presence of bony hooks on the pectoral and caudal-fin rays; bony hooks on the anal-fin rays larger than
those on the pelvic-fin rays; pectoral-fin rays i,5-6,i; presence of three unbranched dorsal-fin rays; absence of an adipose fin;
four scales rows between the anal-fin origin and the lateral line; and four scale rows between the pelvic-fin and the lateral line.
Ecological characteristics of the habitat of the new species are also presented.
Una nueva especie de Tyttocharax se describe para la cuenca del río Güejar, Serranía de La Macarena en Colombia. Tyttocharax
metae es un nuevo registro del género para la cuenca del río Orinoco. La combinación de los siguientes caracteres distingue
a Tyttocharax metae de sus congéneres: presencia de ganchos óseos en los radios de las aletas pectorales y caudal; ganchos
óseos en los radios de la aleta anal de mayor tamaño que los de las aletas pélvicas; radios de las aletas pectorales i,5-6,i; tres
radios simples en la aleta dorsal; ausencia de una aleta adiposa; cuatro escamas entre la línea lateral y el origen de la aleta anal,
y cuatro escamas entre la línea lateral y las aletas pélvicas. Se incluyen datos ecológicos del hábitat propio del nuevo taxón.
Key words: Geographic distribution, Neotropical fish, Osteology, Scales, Teeth.
Introduction
The subfamily Stevardiinae sensu Mirande (2010)
comprises several characid genera previously listed as
incertae sedis in Characidae by Lima et al. (2003), as well as
all members of the inseminating characids of the subfamilies
Stevardiinae and Glandulocaudinae sensu Weitzman et al.
(2005). Among the inseminating stevardiines, Argopleura,
Iotabrycon, Scopaeocharax, Tyttocharax, and Xenurobrycon
have been proposed to belong to a single tribe by Weitzman
& Fink (1985), the Xenurobryconini. Latter, the tribe was
expanded to include Chrysobrycon and Ptychocharax
(Weitzman & Menezes, 1998; Weitzman et al., 1994).
Fishes of the xenurobryconin are sexually dimorphic in a
variety of features (Weitzman & Ortega, 1995; Weitzman &
Menezes, 1998). The great majority of inseminating fishes,
including all known Tyttocharax species, exhibit sperm and
testis modifications that appear to be adaptations related to
this unusual reproductive habit (Burns & Weitzman, 2005;
Pecio et al., 2005).
Weitzman & Fink (1985) described five synapomorphies
that diagnose Tyttocharax: (1) in sexually mature males bony
hooks are confined to the posterior six to eight anal-fin rays.
These hooks are relatively large and arranged in a vertically
elongate cluster. (2) In sexually mature males the posterior three
to five proximal anal-fin radials are expanded in to flattened
plates that serve as the origin for very robust anal-fin erector
and depressor muscles. (3) Mature males and females have
about 25 to 40 conical teeth, arranged in four to seven diagonal
rows in each premaxilla. (4) Mature males and females have
four to eight rows of dentary teeth, all but the innermost
diagonal, with a total of 50 to 80 or more teeth on each mandible,
and with many of the teeth projecting anteriorly or laterally. (5)
The dorso-lateral lamellar process of the angulo-articular bone
is absent. Currently, three valid species are recognized in
Tyttocharax, all originally described from the Amazon River
basin: T. madeirae Fowler, 1913, T. cochui (Ladiges, 1950), and
T. tambopatensis Weitzman & Ortega, 1995.
In this paper we describe a new species of Tyttocharax,
providing morphometrics, pigmentation, and osteological
1
Universidad del Quindío, Laboratorio de Ictiología, A. A. 2639, Armenia, Quindío, Colombia. ceroman@uniquindio.edu.co,
zutana_1@yahoo.com
2
1822 N. Charles St., Belleville, IL, 62221 USA. taphorn@gmail.com
3
Universidad del Atlántico, Departamento de Biología, Barranquilla, Atlántico, Colombia. caturog@gmail.com
519
520
A new species of Tyttocharax from the Güejar River
characters for both sexes, to distinguish it from all congeners.
It is the first record of the genus from the Orinoco River basin,
thus greatly expanding the known range to include the region
around the La Macarena mountain range of the Guaviare River
drainage in Colombia.
Material and Methods
Measurements were made with digital calipers to 0.1 mm
precision, and expressed as percentages of standard length
(SL) or head length (HL). Measurements and counts were taken
on the left side, except when that side was damaged, and
recorded following Fink & Weitzman (1974: 1-2) and Weitzman
et al. (1994:48). Counts for the holotype are indicated with an
asterisk (*). We performed Principal Component Analysis
(PCA) using the Burnaby method to eliminate the influence of
size with the PAST program, version 1.81 for Windows (Hammer
et al., 2008) to test sexual dimorphism in body shape.
Observations of cartilage and bone were made on three cleared
and stained specimens (c&s) prepared following Song &
Parenti (1995). Total vertebral counts were taken from alizarin
cleared and stained specimens. These include the vertebrae of
the Weberian apparatus as well as the complex caudal
ossification, PU1 + U1 with the associated hypural bones and
“half vertebrae” all counted as one element. Bone nomenclature
follows Weitzman (1962) and Vari (1995). In the lists of
paratypes and comparative material, the catalog number is
followed by the number of individuals and the range of SL.
Examined specimens are from Auburn University Natural
History Museum, Auburn, U.S.A. (AUM); Instituto de
Investigaciones de Recursos Biológicos “Alexander von
Humboldt”, Villa de Leyva, Boyacá, Colombia (IAVH); Instituto
de Ciencias Naturales, Museo de Historia Natural-Unidad de
Ictiologìa, Universidad Nacional de Colombia, Bogotá,
Colombia (ICNMHN); Laboratorio de Ictiología, Departamento
de Biología, Universidad del Quindío, Armenia, Colombia (IUQ);
Museo Javeriano de Historia Natural-Laboratorio de Ictiología,
Departamento de Biología, Pontifica Universidad Javeriana,
Bogotá, Colombia (MPUJ); and Museo de Historia Natural,
Departamento de Ictiología, Universidad Nacional Mayor de
San Marcos, Lima, Perú (MUSM).
Tyttocharax metae, new species
Figs. 1-6
Holotype. IUQ 2581, 15.3 mm SL, adult male, Colombia, Meta,
Vista Hermosa near Palestina, río Orinoco basin, río Güejar system,
arroyo Pringamosal, tributary of arroyo Blanco 500 m below
Palestina School, 03º05’22”N 73º49’27”W, 240 m a.s.l., 9 Jan 2009,
C. Román-Valencia, C. García-Alzate & R. Ruiz-C.
Paratypes. All from Colombia: Meta State: La Macarena mountains,
río Orinoco basin, río Güejar system unless noted: AUM 50299, 2,
15.7-18.3 mm SL, IUQ 2343, 3, 14.3-18.6 mm SL, Vista Hermosa,
La Palestina, arroyo Palestina, 03º05’15”N 73º49’54”W, 282 m
a.s.l., 8 Jul 2008, C. Román-Valencia, C. García-Alzate & R. RuizC. IUQ 2344, 4, 13.3-14.0 mm SL, IUQ 2345, 3, 13.2-14.5 mm SL,
Vista Hermosa, La Palestina, creek 2 km north of Las Brisas, road to
Vista Hermosa, 03º03’00”N 73º49’05”W, 264 m a.s.l., 10 Jul 2008,
C. Román-Valencia, C. García-Alzate & R. Ruiz-C. MPUJ 6135, 2,
14.1-15.8 mm SL, Vista Hermosa, La Palestina, creek 2 km north of
Las Brisas, road to Vista Hermosa, 03º03’00”N 73º49’05”W, 264 m
a.s.l., 10 Jul 2008, C. Román-Valencia, C. García-Alzate & R. RuizC. IUQ 2346, 1, 11.4 mm SL, Granada, río Ariari system, arroyo
Mucuyita 03º27’05”N 73º47’49”W, 301 m a.s.l., 7 Jul 2008, C.
Román-Valencia, C. García-Alzate & R. Ruiz-C. IUQ 2347, 2, 12.914.1 mm SL, IUQ 2493, 1, 18.4 mm SL, collected with the holotype.
IUQ 2494, 2, 13.2-14.3 mm SL, Vista Hermosa, Puerto Lucas, arroyo
Acacias on road to Vista Hermosa, 03º05’24”N 73º45’32”W, 240 m
a.s.l., 8 Jan 2009, C. Román-Valencia, C. García-Alzate & R. RuizC. IUQ 2496, 1 c&s, 12.3 mm SL, Vista Hermosa, Puerto Lucas,
arroyo Luciana 100 m north of bridge in Puerto Lucas, 03º06’22”N
73º46’44”W, 253 m a.s.l., 8 Jul 2008, C. Román-Valencia, C. GarcíaAlzate & R. Ruiz-C. IUQ 2495, 3 c&s, 12.4-13.6 mm SL, Vista
Hermosa, La Palestina, creek 1 km north of Las Brisas, 03º02’55”N
73º49’10”W, 278 m a.s.l., 10 Jul 2008, C. Román-Valencia, C. GarcíaAlzate & R. Ruiz-C. IUQ 2755, 4, 15.4-17.1 mm SL, Vista Hermosa,
La Palestina, arroyo Pringamosal, 03º05’29”N 73º49’47”W, 273 m
a.s.l., 9 Nov 2009, M. I. González & A. M. Barrero.
Diagnosis. Tyttocharax metae can be distinguished from all
congeners by having bony hooks on the pectoral and caudalfin rays (vs. absent). Differs from T. madeirae and T. cochui
by the absence of adipose fin (vs. adipose fin present), in
having larger bony hooks on the anal fin than on the rays of
the pelvic fins (vs. bony hooks of the same size on the rays of
those fins). Differs from T. cochui in having i,5-6 pectoral-fin
rays (vs. i,7). Tyttocharax metae can be distinguished from T.
tambopatensis by absence of a sexually dimorphic color
pattern in life (vs. presence of sexually dimorphic color pattern,
males have bright orange abdomens), by the number of scales
rows between anal-fin origin and lateral line (4 vs. 6), by the
number of scale rows between pelvic-fin and lateral line (4 vs.
6), by the number of branched pectoral-fin rays (5-6 vs. 7); by
the number of unbranched dorsal-fin rays (iii vs. ii) and
distance between the dorsal and anal-fin origins (22.3 to 32.2%
SL vs. 38.0-41.0% SL).
Description. Morphometric and meristic data for holotype
and paratypes in Table 1. Body deepest between verticals
through pelvic-fin and dorsal-fin origins in females; deepest
between verticals through posterior margin of dorsal-fin base
and middle part of anal-fin base in males. Predorsal profile of
body generally convex in both sexes. Body profile slightly
elevated at dorsal-fin origin then slightly concave from dorsalfin origin to procurrent caudal-fin rays; slightly convex near
tips of depressed dorsal-fin rays in males. Origin of dorsal fin
nearer caudal-fin base than snout. Dorsal-fin origin at vertical
through anal fin in females; anterior to vertical through analfin origin in males. Tips of pelvic-fin rays reach anal-fin origin
in both sexes. Ventral profile of body convex from tip of lower
jaw to pelvic-fin insertion. Ventral body profile convex along
anal-fin base in males, concave in females. Ventral profile of
caudal peduncle convex in females, concave in sexually
dimorphic males (Figs. 1 and 2). Lower jaw prominent,
C. Román-Valencia, C. A. García-Alzate, R. I. Ruiz-C. & D. C. Taphorn B.
projecting beyond upper jaw. Jaws and lips of mature males
moderately thickened with specialized accommodation of
prominent premaxillary teeth outside of snout. Lips thin in
females. Maxilla extending anteriorly to antero-ventral border
of eye in sexually dimorphic males and females.
All teeth conical and distributed over surface of mandibles
in an apparently random pattern that makes them difficult to
count in traditional rows. Number and distribution of teeth
sexually dimorphic, with males having higher number and
more prominent teeth (Fig. 3). In males, premaxilla with four to
five sets of teeth, each with 18 to 22 conical teeth that extend
over its entire surface; fewer teeth in females, restricted to
ventral half of premaxilla, with up to three sets of conical
teeth, each with a maximum of 15 teeth; only two to three
teeth in upper row. Maxilla in males with two to three sets of
teeth that overlap dorsally; only one series of larger wellseparated teeth ventrally. Maxilla in females with only one
series of non-overlapping teeth positioned dorsally (Fig. 3).
Dentary in males with four sets of 25 to 30 unicuspid teeth
randomly distributed in front of dentary; reduced to two
series laterally where they overlap with maxillary teeth. Females
with only two to three sets of the teeth in front reducing to
just one series laterally.
Scales cycloid, small. Lateral line complete, perforated
scales 31-34 (31*, mode = 33). Scale rows between dorsal-fin
origin and lateral line five (n = 28); scale rows between lateral
line and anal-fin origin four (n = 28); scale rows between
lateral line and pelvic-fin insertion four (n = 28). Predorsal
scales 13-14, arranged in regular series (13*, mean = 13.7,
mode = 13). Dorsal-fin rays ii,5-7,i (ii,5,i*, n = 28); first
521
unbranched ray approximately one-half length of second ray,
its tip reaching proximal bifurcation of first branched ray. Analfin rays ii,13-18,i (ii,14,i*; ii,13-14,i in males and ii,14-18,i in
females, n = 26). Pectoral-fin rays i,5-6,i (i,5,i*; i,5-6,i in males
and females). Pelvic-fin rays i,3-6,i (i,3,i*; i,3-4,i in males and
i,3-6,i in females, n = 26). Pelvic-fin origin anterior to vertical
through dorsal-fin origin. Caudal-fin pouch scale of irregular
shape with approximately 26-28 radii (Fig. 6). Total number of
vertebra 40-41.
Maxilla angular, more pronounced in females in which
ventral margin extends anteriorly and teeth cover anterior
half of this structure. Supraneurals absent. Five infraorbitals,
the first vestigial, infraorbitals two to five in typical position
for genera included in Stevardiinae. The third infraorbital not
in contact with preopercular sensory canal. Anterior margin
of sphenotic extending over fifth infraorbital. Supraoccipital
spine short, not reaching anterior margin of neural complex.
Orbitosphenoid bone with an extension in anterior ventral
region with bands of cartilage in anterior and posterior margins.
Rhinosphenoid bone square, contacting orbitosphenoid and
parasphenoids by bands of cartilage. Parasphenoid not
divided, joined to ventral vomer surface by cartilage; posterior
end of parasphenoid in contact with prootic and basioccipital
by band of cartilage.
Dorsal margin of metapterygoid wide with a crest, foramen
present in medial posterior region, with band of cartilage between
it and quadrate. Ectopterygoid long and narrow, not in contact
with quadrate. Mesopterygoid with entire dorsal margin in contact
with ventral region of parasphenoid, band of cartilage over entire
antero-ventral margin that connects to ectopterygoid.
Table 1. Morphometric data of holotype (male) and paratypes (A, adult males including the holotype and B, adult females and
juveniles) of Tyttocharax metae. Values in parentheses = mean. SD = Standard deviation.
Morphometric
Standard length (mm)
Total length (mm)
Body depth
Snout-dorsal fin distance
Snout-pectoral fin distance
Snout-pelvic fin distance
Dorsal-pectoral fin distance
Snout-anal fin distance
Dorsal fin-hypural distance
Dorsal-anal fin distance
Dorsal-fin length
Pectoral-fin length
Pelvic-fin length
Anal-fin length
Caudal peduncle depth
Caudal peduncle length
Head length
Snout length
Orbital diameter
Postorbital distance
Maxilla length
Interorbital distance
Upper jaw length
A
Holotype
n = 12
SD
15.3
12.6-15.8 (13.9)
1.3
19.7
16.8-20.7 (18.0)
1.6
Percentages of standard length
26.6
20.2-30.1 (25.7)
3.2
64.0
58.7-67.0 (62.3)
1.5
29.9
21.1-33.0 (27.1)
2.4
42.6
39.3-48.2 (43.2)
2.7
46.4
40.6-46.6 (42.2)
2.0
66.6
57.9-66.7 (63.3)
2.8
41.5
41.1-47.8 (42.3)
3.2
26.4
22.3-32.2 (27.2)
1.9
21.2
17.4-26.8 (21.1)
1.8
23.0
16.0-25.3 (20.1)
2.0
23.1
18.8-32.6 (24.6)
3.1
19.4
14.7-24.7 (19.0)
2.8
12.2
11.3-15.5 (12.7)
1.7
5.8
5.7-10.6 (7.8)
1.5
29.3
22.5-29.4 (26.3)
2.2
Percentages of head length
32.2
19.0-38.4 (27.6)
4.4
37.9
37.7-49.3 (42.7)
4.4
35.8
22.1-35.8 (29.8)
4.0
16.6
15.6-34.7 (21.7)
4.6
33.9
33.4-44.4 (38.4)
4.6
23.1
21.4-33.7 (28.2)
3.0
B
n = 16
12.6-15.7 (14.0)
16.2-20.2 (18.1)
SD
1.0
1.3
20.8-27.1 (24.4)
55.5-63.3 (60.6)
23.2-29.9 (26.7)
41.0-50.0 (46.5)
36.5-45.4 (41.8)
60.3-64.4 (62.8)
39.3-44.7 (41.3)
22.1-29.3 (23.8)
15.8-24.3 (19.7)
16.3-22.5 (20.0)
10.9-22.1 (17.1)
13.3-18.1 (15.8)
7.1-12.1 (10.1)
7.5-12.7 (9.9)
21.6-26.4 (24.1)
2.5
2.7
2.7
2.9
3.0
2.9
3.1
1.2
1.7
2.7
2.8
2.8
3.0
1.8
2.6
22.7-35.6 (29.7)
41.4-54.4 (47.8)
31.1-39.9 (35.5)
15.6-28.2 (21.8)
35.7-47.1 (42.5)
26.0-32.8 (29.6)
4.2
5.3
4.9
4.3
4.9
3.7
522
A new species of Tyttocharax from the Güejar River
Fig. 1. Tyttocharax metae, Colombia, Meta. a) holotype, IUQ 2581, adult male, 15.3 mm SL; b) paratype, IUQ 2343; adult female,
15.5 mm SL. Scale bar = 1 cm.
Nasal bone present. Basihyal cartilaginous not divided.
Pharyngeal curved, an elongated plate, with thick cartilage
at dorsal and ventral ends. Sixteen gill rakers on first
branchial arch; 4-5 gill rakers on ceratobranchial and 11-12
gill rakers on epibranchial. Pectoral girdle with pointed dorsal
process on cleithrum. Cleithrum elongated with straight
posterior border; located under ventral edge of operculum.
Anterior border of scapula straight. Postcleithrum 1 and 2
absent, postcleithrum 3 elongated and curved covering more
than half of cleithrum. Proximal pterygiophore rays of dorsal
fin inserted between neural spines 10 and 16; 17th proximal
pterygiophore of anal fin inserted between hemal spines 11
and 12.
Color in alcohol. See Figs. 1 and 2 for pigment patterns in
preserved males and females. Body light yellow, dark brown
on dorsum with dark spot at base of caudal peduncle. Sides
of body with dark stripe that starts posterior to operculum
and extends to caudal spot and widens at level of vertical
through posterior tip of ventral-fin rays. Posterior margins of
scales located on upper sides anterior to dorsal fin dark.
Pectoral, pelvic and anal fins hyaline. Dorsal area of head
dark. Humeral spot visible.
Color in life. Dorsum of body, head and post ventral area
greenish yellow, with an evident absence of dark pigment.
Sides of body with blue stripe, caused by presence of
iridophores that generate an iridescent bluish aspect known
as Rayleigh scattering. The iridophores are limited on sides
Fig. 2. Tyttocharax metae, paratype, IUQ 2343, adult male,
18.4 mm SL. Caudal peduncle and fin, left lateral view.
C. Román-Valencia, C. A. García-Alzate, R. I. Ruiz-C. & D. C. Taphorn B.
523
Fig. 3. Tyttocharax metae, paratypes, IUQ 2493, 18.4 mm SL. Maxilla, premaxilla and dentary, left lateral view. a) female; b) male;
scale bar = 1 mm.
to just dorsal margin of coelomic cavity. From there they extend
posteriorly to the caudal peduncle. Lateral surface of coelomic
cavity covered with leucophores that color this part of fish
white. Bases of middle caudal-fin rays covered with narrow
band of melanophores, more concentrated on dorsal lobe of
caudal fin, and forming caudal spot. Posterior border of
opercle covered by shiny blue iridophores. Humeral spot
horizontally elongated and formed by disperse melanophores
in area along dorsal margin of coelomic cavity, overlaing
iridophores. Anal-fin rays with dispersed melanophores along
their bases. All the fins hyaline. Color pattern identical in
males and females.
Sexual dimorphism. Neither the principal component analysis
among species of Tyttocharax nor regressions comparing males
and females produced significant results. In males, teeth are
found along entire length of anterior margin of maxilla, which is
less sharply pointed on its ventral tip than in females. In both
sexes maxillary teeth diminish gradually in size (Fig. 3). Posterior
pterygiophores (last two or three including terminal piece) of
anal fin in males of Tyttocharax metae are swollen, but in females
are vestigial. Five or six penultimate anal-fin rays each have a
pair of bony hooks (Fig. 3) present at distal extremity. Fin-ray
segment dorsal to one with hook thickened in last rays with
hooks (Fig. 4).
Pelvic fin of T. metae with i,3-6,i rays as in T. tambopatensis,
but distribution of hooks is different: four hooks are present on
simple ray and last branched ray, with one pair at terminus of
each segment on their distal margins; five hooks are found on
first and penultimate branched rays (Fig. 5). Three or four bony
hooks are present on middle portion of caudal-fin rays. Principal
caudal-fin ray count 10/7. Eleventh principal caudal-fin ray (as
identified in females) is transformed in males into an accessory
structure coupled to caudal scale (Fig. 6). Anterior anal-fin lobe
larger in males (18.9% SL) than in females (15.1% SL).
Distribution. This species is known from the río Güejar system
in Meta State, La Macarena Mountains, Orinoco basin,
Colombia (Fig. 7).
Etymology. The specific epithet refers to the Meta State, in
eastern Colombia, where the new species was collected.
Ecological notes. This new species was captured in streams
characterized by relatively rapid water current, running
Fig. 4. Tyttocharax metae, paratype, IUQ 2343, adult male,
18.4 mm SL. Posterior eight anal-fin rays and supporting
pterygiophores left lateral view. Posterior ray split to its base
with anterior half bearing bony hooks. Scale bar = 1 mm.
524
A new species of Tyttocharax from the Güejar River
Fig. 5. Tyttocharax metae, paratype, IUQ 2343, adult male,
18.4 mm SL. Pelvic-fin rays illustrating distribution of pelvicfin hooks, left side, ventro-medial view. Scale bar = 1 mm.
over rocky and sandy bottoms at altitudes between 264282 m a.s.l. Water depth was from 0.5 to 3 m. and stream
width between 1 to 4 m. Riparian vegetation was grass and
trees. The transparency of the tea colored water was usually
high, dissolved oxygen was also relatively high (5.7-7.1
mg/l), pH was usually around neutral (7.1-7.6). Fish species
collected with T. metae are Aphyocharax alburnus,
Astyanax sp., Bujurquina sp., Bryconamericus
cismontanus, Charax metae, Creagrutus calai, Farlowella
vittata, Hemigrammus marginatus, Hoplias malabaricus,
Hyphessobrycon metae, Moenkhausia lepidura group, and
Pyrrhulina brevis.
Remarks. The shape of the last anal-fin pterygiophore is
reported to have an undulated margin in T. madeirae
(Weitzman & Fink, 1985) and an obvious notch in T.
tambopatensis (Weitzman & Ortega, 1995). In T. metae the
margin of the last anal-fin pterygiophore is concave and halfmoon shaped.
Fig. 7. Distribution map for species of Tyttocharax: T. cochui
(), T. madeirae (), T. tambopatensis () and T. metae ().
Modified from Weitzman & Ortega (1995).
The distal extremity of principal caudal-fin ray 11 is
leaf-shaped in T. metae with a straight but inclined dorsal
margin and arched, protruding ventral margin vs. shaped
as an elongate, thickened tubercle in T. madeirae. Three
hooks on principal caudal-fin ray 7 and two on principal
caudal-fin ray 8. The uroneural is subdivided in T. metae
vs. uroneural continuous in T. madeirae.
Four of the five synapomorphies described for
Tyttocharax (Weitzman & Fink, 1985) are present in T. metae,
except for the smaller number of dentary teeth in T. metae
(fewer than 50 teeth in each jaw, Fig. 3, instead of a total of 50
to 80 or more teeth on each mandible). It is possible that this
difference results from the difference in size of the fishes
studied by us and by Weitzman & Fink (1985).
Comparative material. Tyttocharax madeirae: all from Colombia,
Amazonas, Amazon Basin: ICNMHN 6325, 33, 10.4-13.9 mm SL,
km 95 Letícia-Tarapacá road, arroyo La Arenosa; ICNMHN 10329,
6, 14.9-16.6 mm SL, Letícia, tributary of Amazon River, km 6.5,
Tarapacá road, arroyo Tacana; ICNMHN 10045, 2, 15.8-17.8 mm
SL, Letícia, tributary of Amazon River, arroyo Tacana; IAvH 8301,
13, 12.9-17.7 mm SL, Letícia, Matamatá Creek, 03°48’23’’S
70°15’59’’W; IAvH 8302, 4 (c&s), 16.3-20.3 mm SL, Letícia, tributary
of Amazon River; IAvH 11171, 5, 13.0-18.4 mm SL; Letícia, arroyo
Sufragio, El Zafire biological station; IAvH 11170, 23, 11.6-15.9 mm
SL, Letícia, arroyo Sufragio at El Zafire biological station; IAvH 8304,
127, 15.1-20.9 mm SL, Letícia, IAvH 8304, 7 (c&s), 134, 15.4-19.8
mm SL, Letícia, río Purité, 03°41’35’’S 70°12’26’’W; IAvH 11168,
5, 16.0-20.2 mm SL, Letícia, arroyo Gravilla at El Zafire biological
station; IAvH 11169, 48, 11.6-20.3 mm SL, Letícia, IAvH 11169 , 8
(c&s), 15.9-18.5 mm SL, Letícia, arroyo Sufragio at El Zafire biological
station. MPUJ 3461, 4, 16.6-20.0 mm SL, Letícia. Tyttocharax
tambopatensis: MUSM 5087, 7 of 13 paratypes, 11.6-14.3 mm SL,
Peru, Madre de Dios, Tambopata, Tambopata Reserve, río Tambopata,
creek water stream at km 3, Tapir trail.
Acknowledgements
Fig. 6. Tyttocharax metae, paratype, IUQ 2343, adult male,
18.4 mm SL. Caudal skeleton view, left lateral view, pouch
scale. Scale bar = 1 mm.
We thank the University of Quindío, Vicerrectoria de
Investigaciones to C.R-V and C.A.G which financed this study.
We also thank the following persons and museums for loans
C. Román-Valencia, C. A. García-Alzate, R. I. Ruiz-C. & D. C. Taphorn B.
of material under their care: Jaime Aguirre C. (ICNMHN), Juan
D. Bogotá (IAvH), Saul Prada-P. (MPUJ) and Hernán Ortega
(MUSM). We are most grateful to the Juan B. Barrero (Centro
Docente “La Palestina”, Vista Hermosa, Meta) and the Marin
family (Gonzaga, Ordanet and Javier) for their help and
generous hospitality in La Macarena. We also thank Peter
Bartsch and Luiz R. Malabarba for the corrections and
suggestions to improve the paper. Alejandro Escobar Ruiz
prepared figures 1 and 2.
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Submitted July 13, 2010
Ressubmitted April 7, 2011
Accepted June 15, 2012
Published September 28, 2012