Fish , Creagrutus melasma ( Ostariophysi , Characiformes , Characidae )
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Fish , Creagrutus melasma ( Ostariophysi , Characiformes , Characidae )
Check List 5(1): 019–023, 2009.
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
19
Fish, Creagrutus melasma (Ostariophysi, Characiformes, Characidae):
New Venezuelan distribution records
Henry D. Agudelo-Zamora
1
Douglas Rodríguez-Olarte
2
Donald C. Taphorn
3
1
Universidad de Antioquia. Instituto de Ciencias. Laboratorio de Ictiología. Grupo de Ictiología de la Universidad de
Antioquia (GIUA). Bloque 7 - 309. Medellin. Colombia.
E-mail: hdagudelo@gmail.com
2
Universidad Centroccidental Lisandro Alvarado. Departamento de Ciencias Biológicas. Laboratorio de Ecología.
Tarabana, Cabudare, Estado Lara, Venezuela.
3
Universidad Nacional Experimental de los Llanos Occidentales “Ezequiel Zamora” (UNELLEZ). Museo de Zoologia,
Colección de Peces.
Mesa de Cavaca, Guanare, Estado Portuguesa. Venezuela.
Species of the genus Creagrutus Günther, 1864
inhabit a wide variety of Neotropical river
basins from lower Mesoamerica (Panamá)
to southern South America (Paraguay). The
diversity of species in this genus is high in
geomorphologic transition zones, such as
piedmont hill regions between the Andes
mountains and the plains, but they are also quite
diverse in lowland alluvial plains (Vari and
Harold 2001; Rodríguez-Olarte et al. 2007). This
genus includes 66 species (Harold and Vari 1994;
Vari and Harold 2001; Ribeiro et al. 2004; Torres-
Mejía and Vari 2005), and at least fifteen of them
have been reported from Venezuela (Vari and
Harold 2001).
Creagrutus melasma (Vari et al. 1994) was
described from northern Venezuela, from
the Andean drainages of Táchira state, in the
west, to the Caribbean drainages of Sucre state,
in the east. It was also reported from many
headwaters and Andean piedmont streams of
the Orinoco basin as well as in the Tuy and
Neverí rivers, which are part of the Caribbean
slope. It is abundant in the Lake Valencia
drainage from the Buey River (Burro Island)
and around Manaure (Carabobo State). It has
also been found from Guárico State in the
Orituco River, Aragua state in the Cagua River,
Carabobo state in the Guacara River and
Portuguesa state in the Boconó, Guanare,
and Las Marías rivers (Eigenmann 1920;
Taphorn 1992; Vari et al. 1994; Lopez-Rojas and
Bonilla-Ribero 2000; Mago-Leccia and Marín
2004).
We examined specimens from collections of
the Museo de Ciencias Naturales Guanare,
Universidad Nacional Experimental de los
Llanos Occidentales “Ezequiel Zamora”, Guanare,
Venezuela (MCNG); Colección Regional
de Peces, Universidad Centroccidental “Lisandro
Alvarado”, Barquisimeto, Venezuela (CPUCLA);
and Museo de Ciencias Naturales Federico
Carlos Lehmann – INCIVA, Cali, Colombia
(IMCN). Identifications were made using Géry
(1977), Taphorn (1992), Vari et al. (1994),
Vari and Harold (2001), and Agudelo-Zamora
et al. (2008). Measurements were taken
with a digital caliper with precision of 0.1 mm.
Scale and fin-ray counts were done with a
stereoscope using methods of Harold and Vari
(1994).
Creagrutus melasma specimens deposited at
CPUCLA were collected during a series of
expeditions to Falcón State between October 2005
and September 2006. Habitat was described using
a modification of the methodology proposed by
Barbour et al. (1999), and characterization of the
aquatic environments was done following Bain
(1999).
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
19
Fish, Creagrutus melasma (Ostariophysi, Characiformes, Characidae):
New Venezuelan distribution records
Henry D. Agudelo-Zamora
1
Douglas Rodríguez-Olarte
2
Donald C. Taphorn
3
1
Universidad de Antioquia. Instituto de Ciencias. Laboratorio de Ictiología. Grupo de Ictiología de la Universidad de
Antioquia (GIUA). Bloque 7 - 309. Medellin. Colombia.
E-mail: hdagudelo@gmail.com
2
Universidad Centroccidental Lisandro Alvarado. Departamento de Ciencias Biológicas. Laboratorio de Ecología.
Tarabana, Cabudare, Estado Lara, Venezuela.
3
Universidad Nacional Experimental de los Llanos Occidentales “Ezequiel Zamora” (UNELLEZ). Museo de Zoologia,
Colección de Peces.
Mesa de Cavaca, Guanare, Estado Portuguesa. Venezuela.
Species of the genus Creagrutus Günther, 1864
inhabit a wide variety of Neotropical river
basins from lower Mesoamerica (Panamá)
to southern South America (Paraguay). The
diversity of species in this genus is high in
geomorphologic transition zones, such as
piedmont hill regions between the Andes
mountains and the plains, but they are also quite
diverse in lowland alluvial plains (Vari and
Harold 2001; Rodríguez-Olarte et al. 2007). This
genus includes 66 species (Harold and Vari 1994;
Vari and Harold 2001; Ribeiro et al. 2004; Torres-
Mejía and Vari 2005), and at least fifteen of them
have been reported from Venezuela (Vari and
Harold 2001).
Creagrutus melasma (Vari et al. 1994) was
described from northern Venezuela, from
the Andean drainages of Táchira state, in the
west, to the Caribbean drainages of Sucre state,
in the east. It was also reported from many
headwaters and Andean piedmont streams of
the Orinoco basin as well as in the Tuy and
Neverí rivers, which are part of the Caribbean
slope. It is abundant in the Lake Valencia
drainage from the Buey River (Burro Island)
and around Manaure (Carabobo State). It has
also been found from Guárico State in the
Orituco River, Aragua state in the Cagua River,
Carabobo state in the Guacara River and
Portuguesa state in the Boconó, Guanare,
and Las Marías rivers (Eigenmann 1920;
Taphorn 1992; Vari et al. 1994; Lopez-Rojas and
Bonilla-Ribero 2000; Mago-Leccia and Marín
2004).
We examined specimens from collections of
the Museo de Ciencias Naturales Guanare,
Universidad Nacional Experimental de los
Llanos Occidentales “Ezequiel Zamora”, Guanare,
Venezuela (MCNG); Colección Regional
de Peces, Universidad Centroccidental “Lisandro
Alvarado”, Barquisimeto, Venezuela (CPUCLA);
and Museo de Ciencias Naturales Federico
Carlos Lehmann – INCIVA, Cali, Colombia
(IMCN). Identifications were made using Géry
(1977), Taphorn (1992), Vari et al. (1994),
Vari and Harold (2001), and Agudelo-Zamora
et al. (2008). Measurements were taken
with a digital caliper with precision of 0.1 mm.
Scale and fin-ray counts were done with a
stereoscope using methods of Harold and Vari
(1994).
Creagrutus melasma specimens deposited at
CPUCLA were collected during a series of
expeditions to Falcón State between October 2005
and September 2006. Habitat was described using
a modification of the methodology proposed by
Barbour et al. (1999), and characterization of the
aquatic environments was done following Bain
(1999).
Page 2
Check List 5(1): 019–023, 2009.
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
20
All from Venezuela: MCNG 24622, 34 paratypes,
Carabobo, Río Cúpira (sector La Cumaca), new
bridge north of San Diego (10°16'15" N,
67°56'30" W); non-types: MCNG 15281, 5,
Chirgua River, tributary of Pao River (10°8'30.1"
N, 68°11’30.1" W); MCNG 6484, 4, Táchira,
Tributary of Quinimarí River in the Junín
bridge between Veracruz and El Corozo
(7°40'0.1" N, 72°14'49.9" W); MCNG 11661,
21, Táchira, Torondoy, culvert pipe under bridge,
San Cristóbal highway (7°34'19.9" N,
72°14'49.9" W); IMCN 211, 4, Táchira,
Torondoy, culvert pipe under bridge, San
Cristóbal highway (7°34'19.9" N, 72°14'49.9" W);
MCNG 43235, 2, Quirimarí river before its
confluence with Torbes River, (approx. 8° N, 71°
E); MCNG 43264, 2, Quirimarí River before its
confluence with Torbes River (7°40'59.2" N,
72°15'0" W); MCNG 46215, 6, Anzoátegui,
Neverí River, mouth of Quebrada Vega Grande.
Vega Grande.
The range extension for Creagrutus melasma now
includes coastal drainages from the central
western region of Venezuela (Appendix I). The
new localities are: i) Tocuyo river basin (Lara
State, between 520 and 680 meters above sea
level), ii) Hueque river basin, and iii) Ricoa
river basin (at Falcón State, very close to the
“Cueva de la Quebrada del Toro” and “Juan
Crisóstomo Falcón” National Parks in the Sierra
de San Luis, between 127 and 295 m). New sites
of capture belong to the Venezuelan Caribbean
slope (Figure 1).
Figure 1. Area of distribution of Creagrutus melasma. Circles: distribution according to Vari et al. (1994) and
Vari and Harold (2001). Black stars: new records. TL = type locality. H = Hueque River. R = Ricoa River.
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
20
All from Venezuela: MCNG 24622, 34 paratypes,
Carabobo, Río Cúpira (sector La Cumaca), new
bridge north of San Diego (10°16'15" N,
67°56'30" W); non-types: MCNG 15281, 5,
Chirgua River, tributary of Pao River (10°8'30.1"
N, 68°11’30.1" W); MCNG 6484, 4, Táchira,
Tributary of Quinimarí River in the Junín
bridge between Veracruz and El Corozo
(7°40'0.1" N, 72°14'49.9" W); MCNG 11661,
21, Táchira, Torondoy, culvert pipe under bridge,
San Cristóbal highway (7°34'19.9" N,
72°14'49.9" W); IMCN 211, 4, Táchira,
Torondoy, culvert pipe under bridge, San
Cristóbal highway (7°34'19.9" N, 72°14'49.9" W);
MCNG 43235, 2, Quirimarí river before its
confluence with Torbes River, (approx. 8° N, 71°
E); MCNG 43264, 2, Quirimarí River before its
confluence with Torbes River (7°40'59.2" N,
72°15'0" W); MCNG 46215, 6, Anzoátegui,
Neverí River, mouth of Quebrada Vega Grande.
Vega Grande.
The range extension for Creagrutus melasma now
includes coastal drainages from the central
western region of Venezuela (Appendix I). The
new localities are: i) Tocuyo river basin (Lara
State, between 520 and 680 meters above sea
level), ii) Hueque river basin, and iii) Ricoa
river basin (at Falcón State, very close to the
“Cueva de la Quebrada del Toro” and “Juan
Crisóstomo Falcón” National Parks in the Sierra
de San Luis, between 127 and 295 m). New sites
of capture belong to the Venezuelan Caribbean
slope (Figure 1).
Figure 1. Area of distribution of Creagrutus melasma. Circles: distribution according to Vari et al. (1994) and
Vari and Harold (2001). Black stars: new records. TL = type locality. H = Hueque River. R = Ricoa River.
Page 3
Check List 5(1): 019–023, 2009.
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
21
The Falcón State localities, specifically those
from the Ricoa river, are the western limit of the
range of Creagrutus melasma from the Caribbean
dominion. The Caribbean dominion includes three
provinces: Western, Central and Atlantic
(Rodríguez-Olarte et al. 2009). The Western
Caribbean province borders the eastern edge of
Falcón province (Magdalena dominion). Samples
from the Coro and Mitare river drainages revealed
the presence of Creagrutus hildebrandi and C.
maracaiboensis, species that are part of the
Maracaibo basin fauna. We also found Creagrutus
lepidus and C. lassoi from southeast Falcón State,
in the Aroa and Tocuyo river basins.
Physical and chemical data from the sites where
C. melasma was captured are show in Table 1.
Riverbanks at collection sites usually had been
partially deforested, with bands of native
vegetation 6-12 m wide, and some showing signs
of erosion. Although drainage patterns have not
suffered changes in most places sampled, human
impacts of agriculture as well as dredging for
channelization are present (Rodríguez et al. 2005).
Figure 2. Morphologic and chromatic variation in
different populations of Creagrutus melasma from
Venezuela: a) Paratype MCNG 24622, 29.8 mm SL,
Caño la Comarca, Carabobo State; b) CPUCLA
1986, 33 mm SL, Hueque river sector Colombia,
Falcón State; c) CPUCLA 1243, 27.4 mm SL, Ricoa
river sector San Pablo, Falcón State.
The specimens of C. melasma captured from new
localities show differences in color intensity
when compared with specimens from previously
known localities (dorsal and anal fins more
intensely red; pectoral and pelvic fins more
intensely yellow). Figures 2B and 2C, show
differences from the illustration of Román
(1992:169), and also differs from specimens
deposited at MCNG. The snout is more elongate
in specimens from new records (Figure 2B), a
feature more evident in specimens from near the
“Cueva de la Quebrada del Toro”. These
specimens also have more elongate pelvic fins
(extending beyond the first rays of the anal fin)
than specimens from the llanos of the Orinoco in
which the pelvic fins do not surpass the first rays
of the anal fin (according to Taphorn 1992 and the
original description of Vari et al. 1994).
Specimens from Cercado creek (Guarico river
drainage, Lara State) are larger than those
(CPUCLA 609, 5 individuals, mean SL = 43 mm)
reported in the original description (32.4 mm SL
in holotype, 22.8 – 40.7 mm SL in paratypes),
although Taphorn (1992) noted that this species
can reach 50 mm SL.
Table 1. Physical features of collecting stations
from two different drainages in Falcón State, A:
Hueque river, B: Ricoa river ( ± SD), temperature
is a punctual observation, altitude is given as the
mean of the sampled stations, in Falcón State
Venezuela, date: 18-March-06.
Parameter A B
Altitude m.a.s.l 250 - 373 127 - 295
Depth cm 19.3 ± 4.04 21.9 ± 6.41
Velocity m/s 0.20 ± 0.08 0.35 ± 0.37
Substrate Mud (%) 3.66 ± 4.09 3.34 ± 1.96
Sand (%) 5.11 ± 5.10 0.70 ± 1.19
Gravel (%) 13.3 ± 11.2 14.23 ± 10.8
——————————
The population of Creagrutus melasma from
Quebrada del Toro is relatively isolated from
others and shows some unusual color and
morphological variations that require further
study. Perhaps these variations are due to the
isolation of these populations at the northwestern
limit of their distribution. Quebrada del Toro
is a small creek flowing into the lower Tocuyo
river. Vicariance events associated with
the orogenesis of the coastal mountain
range and the Andes, and the resultant shifting of
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
21
The Falcón State localities, specifically those
from the Ricoa river, are the western limit of the
range of Creagrutus melasma from the Caribbean
dominion. The Caribbean dominion includes three
provinces: Western, Central and Atlantic
(Rodríguez-Olarte et al. 2009). The Western
Caribbean province borders the eastern edge of
Falcón province (Magdalena dominion). Samples
from the Coro and Mitare river drainages revealed
the presence of Creagrutus hildebrandi and C.
maracaiboensis, species that are part of the
Maracaibo basin fauna. We also found Creagrutus
lepidus and C. lassoi from southeast Falcón State,
in the Aroa and Tocuyo river basins.
Physical and chemical data from the sites where
C. melasma was captured are show in Table 1.
Riverbanks at collection sites usually had been
partially deforested, with bands of native
vegetation 6-12 m wide, and some showing signs
of erosion. Although drainage patterns have not
suffered changes in most places sampled, human
impacts of agriculture as well as dredging for
channelization are present (Rodríguez et al. 2005).
Figure 2. Morphologic and chromatic variation in
different populations of Creagrutus melasma from
Venezuela: a) Paratype MCNG 24622, 29.8 mm SL,
Caño la Comarca, Carabobo State; b) CPUCLA
1986, 33 mm SL, Hueque river sector Colombia,
Falcón State; c) CPUCLA 1243, 27.4 mm SL, Ricoa
river sector San Pablo, Falcón State.
The specimens of C. melasma captured from new
localities show differences in color intensity
when compared with specimens from previously
known localities (dorsal and anal fins more
intensely red; pectoral and pelvic fins more
intensely yellow). Figures 2B and 2C, show
differences from the illustration of Román
(1992:169), and also differs from specimens
deposited at MCNG. The snout is more elongate
in specimens from new records (Figure 2B), a
feature more evident in specimens from near the
“Cueva de la Quebrada del Toro”. These
specimens also have more elongate pelvic fins
(extending beyond the first rays of the anal fin)
than specimens from the llanos of the Orinoco in
which the pelvic fins do not surpass the first rays
of the anal fin (according to Taphorn 1992 and the
original description of Vari et al. 1994).
Specimens from Cercado creek (Guarico river
drainage, Lara State) are larger than those
(CPUCLA 609, 5 individuals, mean SL = 43 mm)
reported in the original description (32.4 mm SL
in holotype, 22.8 – 40.7 mm SL in paratypes),
although Taphorn (1992) noted that this species
can reach 50 mm SL.
Table 1. Physical features of collecting stations
from two different drainages in Falcón State, A:
Hueque river, B: Ricoa river ( ± SD), temperature
is a punctual observation, altitude is given as the
mean of the sampled stations, in Falcón State
Venezuela, date: 18-March-06.
Parameter A B
Altitude m.a.s.l 250 - 373 127 - 295
Depth cm 19.3 ± 4.04 21.9 ± 6.41
Velocity m/s 0.20 ± 0.08 0.35 ± 0.37
Substrate Mud (%) 3.66 ± 4.09 3.34 ± 1.96
Sand (%) 5.11 ± 5.10 0.70 ± 1.19
Gravel (%) 13.3 ± 11.2 14.23 ± 10.8
——————————
The population of Creagrutus melasma from
Quebrada del Toro is relatively isolated from
others and shows some unusual color and
morphological variations that require further
study. Perhaps these variations are due to the
isolation of these populations at the northwestern
limit of their distribution. Quebrada del Toro
is a small creek flowing into the lower Tocuyo
river. Vicariance events associated with
the orogenesis of the coastal mountain
range and the Andes, and the resultant shifting of
Page 4
Check List 5(1): 019–023, 2009.
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
22
the proto-Orinoco river delta from east to
west (Albert et al. 2006; Hoorn et al. 1995),
isolated these coastal drainages approximately
11–15 m.y. in the Miocene (Lundberg et al.
1998). This was sufficient time to permit
speciation in many groups, as is evidenced by
the presence of many endemic species in this
region: Creagrutus crenatus Vari and Harold,
2001; Hyphessobrycon fernandezi Fernández-
Yépez, 1972; Hypostomus pagei Armbruster,
2003; Austrofundulus leohoignei Hrbek et al.,
2005. Although preliminary analysis of
morphometric characters did not provide
sufficient evidence to permit taxonomic
recognition of the Quebrada del Toro population,
some minor differences were found. DNA
sequence analysis might detect additional
differences.
————————————————
Acknowledgments
HDAZ thanks Hernán Martinez and Betty Zamora for economic help for post graduate studies and the Perez Calle
family for support in Colombia. This work is part of the results of project 001-DAG-2005 of the CDCHT to DRO, and
was carried out under the auspices of the Unidad de Investigación de Ciencias Biológicas de la UCLA. The UCLA’s
collections (CPUCLA) and UNELLEZ (MCNG) provided locality information and specimens for this study.
INAPESCA (MPC-Venezuela) provided scientific collection permits. We also give special thanks to Ahyran Amaro,
Jorge Coronel and Hector Rivera who participated in the field work and two anonymous reviewers for their helpful
suggestions on the manuscript.
————————————————
Literature Cited
Agudelo-Zamora, H. D., D. C. Taphorn, and D.
Rodríguez-Olarte. 2008. Clave para los peces del
género Creagrutus Günther (Teleostei: Ostariophysi:
Characiformes) del Centroccidente de Venezuela.
Boletín Científico Museo de Historia Natural,
Universidad de Caldas 12: 111-116.
Albert, J. S., N. R. Lovejoy, and W. G. R. Crampton.
2006. Miocene tectonism and the separation of cis-
and trans-Andean river basins: Evidence from
Neotropical fishes. Journal of South American Earth
Sciences 21: 14-27.
Armbruser, J. 2003. The species of the Hypostomus
cochliodon group (Siluriformes: Loricariidae).
Zootaxa 249: 1- 60 p.
Bain, M. B. 1999. Substrate; p. 95-100 In M. B. Bain
and N. J. Stevenson (ed.). Aquatic Habitat
assessment: common methods. American Fisheries
Society, Bethesda, Maryland.Barbour, M. T; J.
Gerritsen, D. D. Snyder, and J. B. Stribling. 1999.
Rapid Bioassessment Protocols for Use in Streams
and Wadeable Rivers; Periphyton, Benthic
Macroinvertebrates and Fish, Second Edition. EPA
841-B-99-002. U.S. Environmental Protection Agency;
Office of Water; Washington, D.C.
Eigenmann, C. 1920. The Fishes of Lake Valencia,
Caracas, and of the Rio Tuy at El Concejo,
Venezuela. Study No 44. Indiana University Studies.
Vol VII. 18 p.
Fernández-Yépez, A. 1972. Análisis Ictiológico del
Complejo Hidrográfico Río Yaracuy. Dirección de
Obras Hidráulicas. Ministerio de Obras Públicas.
Republica de Venezuela. 67 p.
Géry, J. 1977. Characoids of the world. T.F.H. Publ.
Neptune City, New Jersey, 672 p.
Harold, A. S. and R. P. Vari. 1994. Systematics of the
trans Andean species of Creagrutus (Ostariophysi,
Characiformes, Characidae). Smithsonian.
Contribution. Zoology 551:1-31.
Hoorn, M. C., J. Guerrero, G. A. Sarmiento, and M. A.
Lorente. 1995. Andean tectonics as a cause for
changing drainage patterns in Miocene northern
South America. Geology 23: 237-240.
Hrbek, T., D. C. Taphorn and J. E. Thomerson. 2005.
Molecular phylogeny of Austrofundulus Myers
(Cyprinodontiformes: Rivulidae), with revision of the
genus and the description of four new species.
Zootaxa 825: 1 -39
Mago-Leccia F. and G. Marín. 2004. Contribución al
estudio de los peces del río Neverí, Edo. Anzoátegui-
Venezuela. BioLlania 14:59-77.
Lundberg, John G. 1998. The temporal context for the
diversification of neotropical fishes In L. R.
Malabarba, R. E. Reis, R. P. Vari, Z. M. Lucena and
C. A. S. Lucena (ed.). Phylogeny and Classification
of Neotropical Fishes. Epipucrs, Porto Alegre, 603p.
Ribeiro, A. C., R. C. Benine, and C. A. Figuereido.
2004. A new species of Creagrutus Günther
(Teleostei: Ostariophysi: Characiformes), from the
upper Rio Paraná basin, central Brazil. Journal of
Fish Biology 64:(3), 597-611.
Rodríguez, J. P., R. Lazo, L. A. Solórzano y F. Rojas-
Suárez (ed.). 2005. Venezuela Digital. Electronic
Database accessible at http://ecosig.ivi.ve. Centro
Internacional de Ecología Tropical (CIET), Instituto
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
22
the proto-Orinoco river delta from east to
west (Albert et al. 2006; Hoorn et al. 1995),
isolated these coastal drainages approximately
11–15 m.y. in the Miocene (Lundberg et al.
1998). This was sufficient time to permit
speciation in many groups, as is evidenced by
the presence of many endemic species in this
region: Creagrutus crenatus Vari and Harold,
2001; Hyphessobrycon fernandezi Fernández-
Yépez, 1972; Hypostomus pagei Armbruster,
2003; Austrofundulus leohoignei Hrbek et al.,
2005. Although preliminary analysis of
morphometric characters did not provide
sufficient evidence to permit taxonomic
recognition of the Quebrada del Toro population,
some minor differences were found. DNA
sequence analysis might detect additional
differences.
————————————————
Acknowledgments
HDAZ thanks Hernán Martinez and Betty Zamora for economic help for post graduate studies and the Perez Calle
family for support in Colombia. This work is part of the results of project 001-DAG-2005 of the CDCHT to DRO, and
was carried out under the auspices of the Unidad de Investigación de Ciencias Biológicas de la UCLA. The UCLA’s
collections (CPUCLA) and UNELLEZ (MCNG) provided locality information and specimens for this study.
INAPESCA (MPC-Venezuela) provided scientific collection permits. We also give special thanks to Ahyran Amaro,
Jorge Coronel and Hector Rivera who participated in the field work and two anonymous reviewers for their helpful
suggestions on the manuscript.
————————————————
Literature Cited
Agudelo-Zamora, H. D., D. C. Taphorn, and D.
Rodríguez-Olarte. 2008. Clave para los peces del
género Creagrutus Günther (Teleostei: Ostariophysi:
Characiformes) del Centroccidente de Venezuela.
Boletín Científico Museo de Historia Natural,
Universidad de Caldas 12: 111-116.
Albert, J. S., N. R. Lovejoy, and W. G. R. Crampton.
2006. Miocene tectonism and the separation of cis-
and trans-Andean river basins: Evidence from
Neotropical fishes. Journal of South American Earth
Sciences 21: 14-27.
Armbruser, J. 2003. The species of the Hypostomus
cochliodon group (Siluriformes: Loricariidae).
Zootaxa 249: 1- 60 p.
Bain, M. B. 1999. Substrate; p. 95-100 In M. B. Bain
and N. J. Stevenson (ed.). Aquatic Habitat
assessment: common methods. American Fisheries
Society, Bethesda, Maryland.Barbour, M. T; J.
Gerritsen, D. D. Snyder, and J. B. Stribling. 1999.
Rapid Bioassessment Protocols for Use in Streams
and Wadeable Rivers; Periphyton, Benthic
Macroinvertebrates and Fish, Second Edition. EPA
841-B-99-002. U.S. Environmental Protection Agency;
Office of Water; Washington, D.C.
Eigenmann, C. 1920. The Fishes of Lake Valencia,
Caracas, and of the Rio Tuy at El Concejo,
Venezuela. Study No 44. Indiana University Studies.
Vol VII. 18 p.
Fernández-Yépez, A. 1972. Análisis Ictiológico del
Complejo Hidrográfico Río Yaracuy. Dirección de
Obras Hidráulicas. Ministerio de Obras Públicas.
Republica de Venezuela. 67 p.
Géry, J. 1977. Characoids of the world. T.F.H. Publ.
Neptune City, New Jersey, 672 p.
Harold, A. S. and R. P. Vari. 1994. Systematics of the
trans Andean species of Creagrutus (Ostariophysi,
Characiformes, Characidae). Smithsonian.
Contribution. Zoology 551:1-31.
Hoorn, M. C., J. Guerrero, G. A. Sarmiento, and M. A.
Lorente. 1995. Andean tectonics as a cause for
changing drainage patterns in Miocene northern
South America. Geology 23: 237-240.
Hrbek, T., D. C. Taphorn and J. E. Thomerson. 2005.
Molecular phylogeny of Austrofundulus Myers
(Cyprinodontiformes: Rivulidae), with revision of the
genus and the description of four new species.
Zootaxa 825: 1 -39
Mago-Leccia F. and G. Marín. 2004. Contribución al
estudio de los peces del río Neverí, Edo. Anzoátegui-
Venezuela. BioLlania 14:59-77.
Lundberg, John G. 1998. The temporal context for the
diversification of neotropical fishes In L. R.
Malabarba, R. E. Reis, R. P. Vari, Z. M. Lucena and
C. A. S. Lucena (ed.). Phylogeny and Classification
of Neotropical Fishes. Epipucrs, Porto Alegre, 603p.
Ribeiro, A. C., R. C. Benine, and C. A. Figuereido.
2004. A new species of Creagrutus Günther
(Teleostei: Ostariophysi: Characiformes), from the
upper Rio Paraná basin, central Brazil. Journal of
Fish Biology 64:(3), 597-611.
Rodríguez, J. P., R. Lazo, L. A. Solórzano y F. Rojas-
Suárez (ed.). 2005. Venezuela Digital. Electronic
Database accessible at http://ecosig.ivi.ve. Centro
Internacional de Ecología Tropical (CIET), Instituto
Page 5
Check List 5(1): 019–023, 2009.
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
23
Venezolano de Investigaciones Científicas (IVIC),
Conservación Internacional Venezuela y UNESCO,
Venezuela. Captured on 22 March 2007.
Rodríguez-Olarte D., D. C. Taphorn, and J. Lobon-
Cervia. 2009. Patterns of freshwater fishes of the
Caribbean versant of Venezuela. International
Review of Hydrobiology 94: (1) 67-90.
Rodríguez-Olarte, D., J. Coronel, D. C. Taphorn, y A.
Amaro. 2007. Los peces y su conservación en el río
Tocuyo, la cuenca andina de la vertiente Caribe en
Venezuela. Memoria de la Fundación La Salle de
Ciencias Naturales 165:45-72.
Román, B. 1992. Peces de agua dulce de Venezuela I.
Editorial Biosfera. Caracas, Venezuela. 191p.
Taphorn, D. C. 1992. The Characiform fishes of the
Apure river drainage, Venezuela. Revista. BioLlania.
Edición Especial N° 4. 537 pp.
Torres-Mejía, M. and R. Vari. 2005. New Species of
Creagrutus (Teleostei: Characiformes: Characidae)
from the Río Magdalena Basin, Colombia. Copeia
4:812-817.
Vari, R.; A. Harold and Taphorn, D. 1994. Creagrutus
melasma, A new species of characid fish (Teleostei:
Characiformes) from upland streams of Northern
Venezuela. Proceedings of the Biological Society of
Washington 107(1) 90-96.
Vari, R. and A. Harold. 2001. Phylogenetic study of
Neotropical fish genera Creagrutus Günther and
Piabina Reinhardt (Teleostei: Ostariophysi:
Characiformes), with a revision of the cis-Andean
species. Smithsonian Contributions to Zoology.
Number 613. Washington, D.C. 131 p.
Received February 2008
Accepted December 2008
Published online January 2009
————————————————
Appendix 1: New distribution records of Creagrutus melasma: (All from Venezuela). Lara State: CPUCLA 607, 18
specimens, El Cercado stream, Tributary of Guárico river (09°41'58.5"N 69°47'57"W), 04 March 2003, D. Rodríguez-
Olarte, J. Coronel; CPUCLA 609, 5, El Cercado stream, Tributary of Guárico river (09°41'58.5"N 69°47'57"W), 04
March 2003, D. Rodríguez-Olarte, J. Coronel; CPUCLA 830, 4, Tocuyo river, between Humocaro Bajo and Humocaro
Alto (09°47'59.3"N 69°59'1.6"W), 16 October 2002, D. Rodríguez-Olarte, J. Coronel. Falcón State : CPUCLA 342,
109, National Park Cueva de la Quebrada del Toro (10°49'16.6"N 69°08'10"W) , 08 June 2002, D. Rodríguez-Olarte, J
Coronel, H. Rivera, C. López; CPUCLA 346, 104, National Park Cueva de la Quebrada del Toro (10°49'16.6"N
69°08'10"W) , 08 June 2002, D. Rodríguez-Olarte, J. Coronel, H. Rivera, C. López; CPUCLA 352, 158, National Park
Cueva de la Quebrada del Toro (10°49'16.6"N 69°08'10"W), 08 June 2002, D. Rodríguez-Olarte, J. Coronel, H. Rivera,
C. López; CPUCLA 1243, 3, Río Ricoa, sector San Pablo (11°12'17.8"N 69°29'43.5"W), 19 October 2005, D.
Rodríguez-Olarte, D. Taphorn, J. Coronel, A. Amaro, H. Rivera; CPUCLA 1627, 1, Macoruca stream, Tributary of
Ricoa river (11°17'49.7"N 69°24'55.5"W), 18 March 2006, D. Rodríguez-Olarte, A. Amaro, H. Agudelo-Z., H. Rivera;
CPUCLA 1259, 3, Hueque river, sector Quebracho (11°08'24.6"N 69°33'13.5"W), 19 October 2005, D. Rodríguez-
Olarte, D. Taphorn, J. Coronel, A. Amaro, H. Rivera; CPUCLA 1661, 6, Hueque river, sector Quebracho
(11°08'24.6"N 69°33'13.5"W), 25 April 2006, D. Rodríguez-Olarte, A. Amaro, H. Agudelo-Z., H. Rivera.
ISSN: 1809-127X
NOTES ON GEOGRAPHIC DISTRIBUTION
23
Venezolano de Investigaciones Científicas (IVIC),
Conservación Internacional Venezuela y UNESCO,
Venezuela. Captured on 22 March 2007.
Rodríguez-Olarte D., D. C. Taphorn, and J. Lobon-
Cervia. 2009. Patterns of freshwater fishes of the
Caribbean versant of Venezuela. International
Review of Hydrobiology 94: (1) 67-90.
Rodríguez-Olarte, D., J. Coronel, D. C. Taphorn, y A.
Amaro. 2007. Los peces y su conservación en el río
Tocuyo, la cuenca andina de la vertiente Caribe en
Venezuela. Memoria de la Fundación La Salle de
Ciencias Naturales 165:45-72.
Román, B. 1992. Peces de agua dulce de Venezuela I.
Editorial Biosfera. Caracas, Venezuela. 191p.
Taphorn, D. C. 1992. The Characiform fishes of the
Apure river drainage, Venezuela. Revista. BioLlania.
Edición Especial N° 4. 537 pp.
Torres-Mejía, M. and R. Vari. 2005. New Species of
Creagrutus (Teleostei: Characiformes: Characidae)
from the Río Magdalena Basin, Colombia. Copeia
4:812-817.
Vari, R.; A. Harold and Taphorn, D. 1994. Creagrutus
melasma, A new species of characid fish (Teleostei:
Characiformes) from upland streams of Northern
Venezuela. Proceedings of the Biological Society of
Washington 107(1) 90-96.
Vari, R. and A. Harold. 2001. Phylogenetic study of
Neotropical fish genera Creagrutus Günther and
Piabina Reinhardt (Teleostei: Ostariophysi:
Characiformes), with a revision of the cis-Andean
species. Smithsonian Contributions to Zoology.
Number 613. Washington, D.C. 131 p.
Received February 2008
Accepted December 2008
Published online January 2009
————————————————
Appendix 1: New distribution records of Creagrutus melasma: (All from Venezuela). Lara State: CPUCLA 607, 18
specimens, El Cercado stream, Tributary of Guárico river (09°41'58.5"N 69°47'57"W), 04 March 2003, D. Rodríguez-
Olarte, J. Coronel; CPUCLA 609, 5, El Cercado stream, Tributary of Guárico river (09°41'58.5"N 69°47'57"W), 04
March 2003, D. Rodríguez-Olarte, J. Coronel; CPUCLA 830, 4, Tocuyo river, between Humocaro Bajo and Humocaro
Alto (09°47'59.3"N 69°59'1.6"W), 16 October 2002, D. Rodríguez-Olarte, J. Coronel. Falcón State : CPUCLA 342,
109, National Park Cueva de la Quebrada del Toro (10°49'16.6"N 69°08'10"W) , 08 June 2002, D. Rodríguez-Olarte, J
Coronel, H. Rivera, C. López; CPUCLA 346, 104, National Park Cueva de la Quebrada del Toro (10°49'16.6"N
69°08'10"W) , 08 June 2002, D. Rodríguez-Olarte, J. Coronel, H. Rivera, C. López; CPUCLA 352, 158, National Park
Cueva de la Quebrada del Toro (10°49'16.6"N 69°08'10"W), 08 June 2002, D. Rodríguez-Olarte, J. Coronel, H. Rivera,
C. López; CPUCLA 1243, 3, Río Ricoa, sector San Pablo (11°12'17.8"N 69°29'43.5"W), 19 October 2005, D.
Rodríguez-Olarte, D. Taphorn, J. Coronel, A. Amaro, H. Rivera; CPUCLA 1627, 1, Macoruca stream, Tributary of
Ricoa river (11°17'49.7"N 69°24'55.5"W), 18 March 2006, D. Rodríguez-Olarte, A. Amaro, H. Agudelo-Z., H. Rivera;
CPUCLA 1259, 3, Hueque river, sector Quebracho (11°08'24.6"N 69°33'13.5"W), 19 October 2005, D. Rodríguez-
Olarte, D. Taphorn, J. Coronel, A. Amaro, H. Rivera; CPUCLA 1661, 6, Hueque river, sector Quebracho
(11°08'24.6"N 69°33'13.5"W), 25 April 2006, D. Rodríguez-Olarte, A. Amaro, H. Agudelo-Z., H. Rivera.
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