10 September 2009 NUMBER 17 checklist of freshwater fishes of the guiana shield BULLETIN of the Biological Society of Washington ISSN 0097-0298
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10 September 2009 NUMBER 17
checklist of freshwater fishes of the guiana shield
BULLETIN of the
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ISSN 0097-0298
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CHECKLIST OF THE FRESHWATER FISHES OF THE
GUIANA SHIELD
Richard P. Vari, Carl J. Ferraris, Jr., Aleksandar Radosavljevic, and Vicki A. Funk
i
Front cover illustration: Pseudolithoxus dumus, family Loricariidae (see Plate 12, Figure G).
Illustrations facing each section:
For the Introduction, montage of radiographs of fishes from the rivers of the Guiana Shield, upper left –
Rhaphiodon vulpinis, Cynodontidae; upper right – Prochilodus mariae, Prochilodontidae; center – Serrasalmus
irritans, Characidae, Serrasalminae; lower left – Corydoras filamentosus, Callichthyidae; and lower right,Sternarchorhynchus roseni, Apteronotidae, mature male with enlarged dentary dentition. Images and plate
prepared by S. Raredon.
For the Fishes of the Guiana Shield, Acnodon oligocanthus (Serrasalminae, juvenile) from Steindachner, F. 1915.
Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen
Classe, Wien 93:15–105 (date based on release of separates of main work published in 1917).
For the Guide to the Checklist, Acroronia nassa (Cichlidae) from Steindachner, F. 1875. Sitzungsberichte der
Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen Classe, Wien 71:61–137.
For the Photographic Atlas of Fishes of the Guiana Shield, Leptodoras hasemani (Doradidae) from Steindachner,
F. 1915. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftli-
chen Classe, Wien 93:15–105 (date based on release of separates of main work published in 1917).
Preferred citations:
Vari, R. P., C. J. Ferraris, Jr., A. Radosavljevic, & V. A. Funk, eds., 2009. Checklist of the freshwater fishes of the
Guiana Shield.—Bulletin of the Biological Society of Washington, no. 17.
or, e.g.,
Vari, R. P., & C. J. Ferraris, Jr. 2009. Fishes of the Guiana Shield. In Vari, R. P., C. J. Ferraris, Jr., A.
Radosavljevic, & V. A. Funk, eds., 2009. Checklist of the freshwater fishes of the Guiana Shield.—Bulletin ofthe Biological Society of Washington, no. 17.
ii
CONTENTS
CONTRIBUTORS. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iv
ABSTRACT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii
INTRODUCTION. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Vicki A. Funk and Carol L. Kelloff
FISHES OF THE GUIANA SHIELD. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Richard P. Vari and Carl J. Ferraris, Jr.
GUIDE TO THE CHECKLIST . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Aleksandar Radosavljevic
CHECKLIST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25Pristiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Mylobatiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Osteoglossiformes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Anguilliformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Clupeiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Characiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Siluriformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Gymnotiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45Cyprinodontiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Beloniformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Synbranchiformes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Perciformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Pleuronectiformes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Tetraodontiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Lepidosireniformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
PHOTOGRAPHIC ATLAS OF FISHES OF THE GUIANA SHIELD. . . . . . . . . . . . . . . . . . . . . . 53
Mark Sabaj Perez
INTRODUCTION
APPENDIX: PLATES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
INDEX TO ORDERS, FAMILIES, AND SUBFAMILIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
iii
CONTRIBUTORS
James S. Albert, Department of Biology, University of Louisiana at Lafayette, Lafayette, Louisiana 70504-2451,
U.S.A., e-mail: [email protected]
Jonathan W. Armbruster, Department of Biological Sciences, 331 Funchess, Auburn University, Alabama 36849,
U.S.A., e-mail: [email protected]
Paulo A. Buckup, Departamento de Vertebrados, Museu Nacional, Quinta da Boa Vista, 20940-040 Rio de
Janeiro, RJ, Brazil, e-mail: [email protected]
Ricardo Campos-da-Paz, Escola de Ciencias Biologicas, Universidade Federal do Estado do Rio de Janeiro, Av.
Pasteur, 458/sala 408, Urca - Rio de Janeiro, RJ, 22290-240, Brazil, e-mail: [email protected]
Marcelo R. de Carvalho, Instituto de Biociencias, Universidade de Sao Paulo, Rua do Matao, Trav. 14, no. 101,
Sao Paulo, SP, 05508-900, Brazil, e-mail: [email protected]
Lilian Cassati, Laboratorio de Ictiologia, Departamento de Zoologia e Botanica, IBILCE-UNESP, Rua Cristovao
Colombo, 2265, Sao Jose do Rio Preto, SP, 15054-000, Brazil, e-mail: [email protected]
Ning Labbish Chao, Universidade Federal do Amazonas, Caixa Postal 3695, Manaus, AM 69051-970, Brazil,
e-mail: [email protected]
Bruce B. Collette, National Marine Fisheries Service, Systematics Laboratory MRC 153, National Museum of
Natural History, Washington D.C. 20560-0153, U.S.A., e-mail: [email protected]
Wilson J. E. M. Costa, Laboratorio de Ictiologia Geral e Aplicada, Departamento de Zoologia -- UFRJ, Caixa
Postal 68049, 21944-970 Rio de Janeiro, RJ, Brazil, e-mail: [email protected]
Fabio di Dario, Universidade Federal do Rio de Janeiro, Av. Sao Jose do Barreto, Sao Jose do Barreto, Caixa
Postal 119331, Macae, RJ, 27921-550, Brazil, e-mail: [email protected]
Carl J. Ferraris, Jr., 2944 NE Couch St., Portland, OR 97232, USA; and Division of Fishes, National Museum of
Natural History, Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A., e-mail: [email protected]
John P. Friel, Museum of Vertebrates, Cornell University, E151 Corson Hall, Ithaca, New York 14853-2701,
U.S.A., e-mail: [email protected]
Vicki A. Funk, Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0166, U.S.A., e-mail: [email protected]
Michel Jegu, Antenne IRD, UR 131, Laboratoire d’Ichtyologie, M.N.H.N., 43 rue Cuvier, 75231 Paris Cedex 05,
France, e-mail: [email protected]
Carol L. Kelloff, Department of Botany, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0166, U.S.A., e-mail: [email protected]
Sven O. Kullander, Department of Vertebrate Zoology, Swedish Museum of Natural History, P. O. Box 50007, SE-
104 05 Stockholm, Sweden, e-mail: [email protected]
Francisco Langeani Neto, Laboratorio de Ictiologia, Departamento de Zoologia e Botanica, IBILCE-UNESP, Rua
Cristovao Colombo, 2265, Sao Jose do Rio Preto, SP, 15054-000, Brazil, e-mail: [email protected]
Flavio C. T. Lima, Museu de Zoologia, Universidade de Sao Paulo, Caixa Postal 42594, Sao Paulo, SP, 04299-970,
Brazil, e-mail: [email protected]
Rosana S. Lima, Universidade de Estado de Rio de Janeiro, Faculdade de Formacao de Professores, Rua Dr.
Francisco Portela, 794, Sao Goncalo, RJ, 24435-000, Brazil, e-mail: [email protected]
Carlos A. S. Lucena, Laboratory of Ichthyology, Museu de Ciencias e Tecnologia PUCRS, Caixa Postal 1429,
Porto Alegre, RS, 90619-900, Brazil, e-mail: [email protected]
iv
Paulo H. F. Lucinda, Laboratorio de Ictiologia, Universidade do Tocantins - Campus de Porto Nacional Caixa
Postal 25, Porto Nacional, TO, 77500-000, Brazil, e-mail: [email protected]
John G. Lundberg, The Academy of Natural Sciences, Department of Ichthyology, 1900 Benjamin Franklin
Parkway, Philadelphia, Pennsylvania 19103, U.S.A., e-mail: [email protected]
Luiz R. Malabarba, Departamento de Zoologia -- IB, Universidade Federal do Rio Grande do Sul, Av. Bento
Goncalves, 9500 - bloco IV - Predio 43435, Porto Alegre, RS, 91501-970, Brazil, e-mail: [email protected]
John D. McEachran, Department of Wildlife & Fishery Science, Texas A&M University, 22587 AMU, College
Station, Texas 77843-2258, U.S.A., e-mail: [email protected]
Naercio A. Menezes, Museu de Zoologia, Universidade de Sao Paulo, Caixa Postal 42594, Sao Paulo, SP, 04299-
970, Brazil, e-mail: [email protected]
Cristiano Moreira, Departamento de Ciencias Biologicas, Universidade Federal de Sao Paulo, Rua Prof. Artur
Riedel, 275, Diadema, SP, 09972-270, Brazil, e-mail: [email protected]
Carla S. Pavanelli, Fundacao Universidade Estadual de Maringa, Nupelia, Av. Colombo, 3690, Maringa, PR,
87020-900, Brazil, e-mail: [email protected]
Aleksandar Radosavljevic, The City College of New York, 526 Marshal Science Building, 160 Convent Avenue,
New York, New York 10031, U.S.A., e-mail: [email protected]
Robson T. C. Ramos, Depto de Sistematica e Ecologia, Universidade Federal da Paraıba, PB, Joao Pessoa, 58059-
900, Brazil, e-mail: [email protected]
Roberto E. Reis, Laboratory of Ichthyology, Museu de Ciencias e Tecnologia PUCRS, Caixa Postal 1429, Porto
Alegre, RS, 90619-900, Brazil, e-mail: [email protected]
Ricardo S. Rosa, Depto de Sistematica e Ecologia, Universidade Federal da Paraıba, Joao Pessoa, PB, 58059-900,
Brazil, e-mail: [email protected]
Mark Sabaj Perez, The Academy of Natural Sciences, Department of Ichthyology, 1900 Benjamin Franklin
Parkway, Philadelphia, Pennsylvania 19103, U.S.A., e-mail: [email protected]
Scott A. Schaefer, Division of Vertebrate Zoology, American Museum of Natural History, Central Park West at
79th Street, New York, New York 10024-5192, U.S.A., e-mail: [email protected]
Oscar A. Shibatta, Departamento de Biologia Animal e Vegetal, Centro de Ciencias Biologicas, Universidade
Estadual de Londrina, 86051-990 Londrina, PR, Brazil, e-mail: [email protected]
Monica Toledo-Piza, Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo, Caixa
Postal 11461, 05422-970 Sao Paulo, SP, Brazil, e-mail: [email protected]
Richard P. Vari, Division of Fishes, National Museum of Natural History, Smithsonian Institution, Washington,
D.C. 20560-0159, U.S.A., e-mail: [email protected]
Claude Weber, Department d’Herpetologie et d’Ichtyologie, Museum d’Histoire Naturelle, P. O. Box 434, CH-
1211 Geneve, Switzerland, e-mail: [email protected]
Marilyn Weitzman, Division of Fishes, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0159, U.S.A., e-mail: [email protected]
Stanley H. Weitzman, Division of Fishes, National Museum of Natural History, Smithsonian Institution,
Washington, D.C. 20560-0159, U.S.A., e-mail: [email protected]
Wolmar Wosiacki, Museu Paraense Emılio Goeldi, Laboratorio de Ictiologia, Av. Magalhaes Barata 376 Caixa
Postal Box 399, Belem, PA, 66040-170, Brazil, e-mail: [email protected]
Angela M. Zanata, Departamento de Zoologia, Universidad Federal da Bahia, Rua Barao de Geremoabo, Campus
de Ondina, Salvador, BA, 40170-290, Brazil, e-mail: [email protected]
v
Abstract.—Distributions are provided for 1168 species of fishes that live in the
freshwater drainage systems overlying the Guiana Shield in Brazil, Colombia,
French Guiana, Guyana, and Suriname. This total includes representatives of
376 genera, 49 families, and 15 orders, with five orders (Characiformes,
Siluriformes, Perciformes, Gymnotiformes, and Cyprinodontiformes) account-
ing for 96.7% of the diversity. Drainage systems on the Guiana Shield are home
to approximately 23% of the freshwater species known from Central and SouthAmerica. A summary is provided of ichthyological collecting on the Shield along
with summaries of major publications dealing with fishes of each region on the
Shield. Factors that may account for the high species level diversity in that
region are discussed. Methods for photographing fishes are detailed, and a
photographic album of 127 species of 46 families that occur on the Shield is
included.
Key Words.—Guiana Shield Fishes, Freshwaters, Brazil, Colombia, French
Guiana, Guyana, Suriname, Venezuela.
vii
INTRODUCTION
V. A. FUNK and CAROL L. KELLOFF
In the face of the growing biodiversity crisis, we must
move to document and evaluate the biota of our
natural areas. As taxonomists, it is part of our job to
name, place into groups, and keep track of the species
that we study. Often we use the distribution of the taxa
to help with our work and to provide a broader context
for the results. In doing this job, taxonomists produce
checklists, floras and faunas, and monographs. In fact,
collecting specimens and producing these documents
are essential elements in our quest to understand the
natural world and how it evolved. Checklists are the
‘‘first pass’’ in our attempts to understand the diversity
of an area. They give us the first approximation of the
known diversity of any group or groups of organisms
and they often provide the first profile of the biodiver-
sity in relatively poorly known parts of the globe. For
many groups, they may represent the most complete
information available and since basic taxonomic data
provide the essential information for studies in sys-
tematic and evolutionary biology, we continually seek
to improve and update it. Checklists have many uses:
they are aids in the identification and correct naming of
species, they serve as essential resources for biodiver-
sity estimates and biogeographic studies, and they are
starting points for more detailed studies of an area’s
biota. When reviewed by many specialists, they often
represent the most advanced state of knowledge avail-
able in the field. But they do more than that, beyond
these basics they increase our knowledge of this fasci-
nating region and act as a starting point for additional
biodiversity research because they give insight into the
species richness, endemicity, and floral and faunal
affinities for the region and provide the baseline
information for analyzing species turnover rates and
migration or invasion events and many other biological
phenomena (e.g., Funk & Richardson 2002, Funk et al.
2002, 2005; Ferrier et al. 2004, Kelloff & Funk 2004).
Checklists also increase national and regional pride by
demonstrating the diversity of the area and provide
important public outreach and fundamental informa-
tion to be used by governments and conservation
organizations in addressing the biodiversity crisis. This
Checklist of the Fishes of the Guiana Shield, when
combined with the recently published Checklist of the
Terrestrial Vertebrates of the Guiana Shield (Hollowell
& Reynolds 2005), and Checklist of the Plants of the
Guiana Shield (Funk et al. 2007), provides a sound
basis for future research and conservation efforts.
This Checklist of the Fishes of the Guiana Shield is an
excellent example of effective interaction, involving
ichthyologists from around the world who contributed
in diverse ways to its completion. It began as an
extraction from Reis et al. (2003) but quickly grew into
a larger and more involved project. As with anyendeavor of this type, insights that specialists have
gained through their experience are irreplaceable in
correcting errors and updating classifications. In order
to make future editions of this checklist as current and
accurate as possible, specialists are encouraged to
contact the Smithsonian’s Biological Diversity of the
Guiana Shield Program with additions or corrections.
The Guiana Shield
The Guiana Shield region is a biologically rich area
that includes much of northeastern South America
(Fig. 1). It is strictly defined by the underlying
geological formation known as the Guiana Shield,
and in the context of this volume the term Guiana
Shield also refers to the corresponding geographic
region. That region includes the Venezuelan states ofBolıvar and Amazonas, and a portion of Delta
Amacuro; all of Guyana, Suriname, and French
Guiana; and parts of northern Brazil. Several geolog-
ical outliers of the Guiana Shield occur west of the
Orinoco River in Colombia. In Spanish and Portu-
guese speaking countries, the region is often referred to
as the ‘‘Guayana’’; thus the terms Colombian
Guayana, Brazilian Guayana, and VenezuelanGuayana are often used. The total area of the Guiana
Shield is approximately 1,520,000 km2. Table 1 lists the
square kilometers of political divisions that occur
within the region. See Berry et al. (1995) for a review
of definitions of and terminology related to the Guiana
Shield region.
Geology
The Guiana Shield (Fig. 1) is a distinct geologic
region that underlies Guyana, Suriname, French
Guiana, parts of Venezuela, and Brazil, and a small
area in Colombia. It is roughly bounded by the
Atlantic Ocean to the north and east, the Orinoco
River to the north and west, the Rıo Negro (a major
tributary of the Amazon River) to the southwest, and
the Amazon River to the south (Gibbs & Barron 1993).The Guiana Shield is a distinct ancient geological
formation that includes the mountain systems that
form the watershed boundary between the Amazon
and Orinoco rivers. On the Shield’s western side, the
Orinoco River and Rıo Negro are connected by the
Rıo Casiquiare, making most of the region somewhat
like an island; however, there are some areas of nearby
Colombia that have remnants of the Shield formation.The southern boundary of the Guiana Shield is
difficult to define precisely, as a broad band of outwash
materials resulting from erosion occurs between
mountains on the southern boundary of the Shield
and the Amazon and Negro rivers stretching into parts
of northern Brazil. Also, much of the Venezuelan state
of Delta Amacuro occurs over thick sedimentsdeposited primarily by the Orinoco River and is really
not strictly part of the Shield; however, some moun-
tains of the Guiana Shield occur in this state’s southern
section, and a large proportion of the sediments of the
delta are derived from outwash from the highlands of
the Shield. For more detailed discussions of the
geology of the area, readers should refer to Gibbs &
Barron (1993) and Huber (1995a).
The base of the Guiana Shield is composed of
crystalline rocks of Proterozoic origin; these are mainly
granites and gneisses formed between 3.6 and 0.8
billion years ago (Mendoza 1977, Schubert & Huber
1990). Large areas of the Shield were overlain with
sediments from 1.6 to 1 billion years ago and cemented
during thermal events (Huber 1995a). These quartzite
and sandstone rocks comprise the Roraima formation
and today remnants are scattered across the central
portion of the Shield extending west from the Potaro
Plateau of the Pakaraima Mountains in central
Guyana through parts of Venezuela and Colombia
(Arbelaez & Callejas 1999) and south into northern
Brazil (Leechman 1913, Gibbs & Barron 1993). Within
this area, erosion has resulted in numerous vertical-
walled, frequently flat-topped mountains called ‘‘te-
puis,’’ among them are Chimantı-tepui (2550 m), Cerro
Duida (2358 m), and Auyan-tepui (2450 m). Pico de
Neblina (3014 m) is the Shield’s western-most tepui
and highest point, located on the southern-most
segment of the border between Venezuela and Brazil.
Mount Roraima (2810 m) is located at the juncture of
Guyana, Venezuela, and Brazil and includes the
highest point within Guyana. The eastern-most peaks
in Guyana reach approximately 2000 m elevation,
including Mt. Ayanganna (2041 m). Several of these
mesa-like formations are virtually inaccessible by foot
Figure 1. The Guiana Shield; adapted from Gibbs & Barron (1993), with the region of western outliers indicated.
Table 1.—Divisions of the Guiana Shield in approximately west to
east arrangement, with abbreviations used and estimated areas.
Division Abbr. Area (km2)
*Colombian Guayana CG 120,325
Amazonas, Venezuela VA 175,750
Bolıvar, Venenzuela BO 238,000
*Delta Amacuro, Venezuela DA 40,200
*Amazonas, Brazil BA 125,550
*Roraima, Brazil RO 173,750
*Para, Brazil PA 243,280
*Amapa, Brazil AP 98,750
Guyana GU 214,970
Suriname SU 163,270
French Guiana FG 91,000
Total (km2) 1,896,845
*5 not all parts of these politically defined areas are included in
the Guiana Shield region.
2 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
and are so unusual that they inspired a fictional
scientific expedition referred to one as ‘‘The Lost
World’’ (Doyle 1912), a term sometimes applied to all
tepuis. Notable waterfalls of the region include Angel
Falls (979 m) on Auyan-tepui in Venezuela and
Kaieteur Falls (226 m), which flows year around, on
the eastern-most edge of the Roraima formation
known as the Potaro Plateau in Guyana.
Granitic dome mountains occur on the Shield in the
southern part of the three Guianas (Guyana, Sur-
iname, French Guiana), where they are known as
‘‘inselbergs,’’ as well as in the western extreme of the
Shield in the Puerto Ayacucho region in Venezuela
where they are called ‘‘lajas.’’ Deposits of low-nutrient
white sands occur inland of the coastal plain, in belts
across the Shield, and in isolated pockets. Large areas
of savanna are found in the region, particularly the
complex of savannas that includes the Rupununi
Savanna in southwestern Guyana, the Gran Sabana
in eastern Venezuela, and the savannas of northern
Roraima, Brazil. In some of these areas the sands
overlay a clay hardpan that is resistant to penetration
by tree roots and that floods during the heavy rainy
season, resulting in limited forest growth. Tertiary and
Quaternary sediments separate the southern edge of
the Guiana Shield from the Amazon River and the
eastern edge from the Atlantic Ocean.
Climate
As a whole, the Guiana Shield region has a tropical
climate characterized by a relatively high mean annual
temperature exceeding 25uC at sea level, an annual
monthly maximum temperature range of less than 5uC,
and an average daily temperature range of approxi-
mately 6uC (Snow 1976). Because of the Guiana
Shield’s location just north of the equator, its climate
varies primarily according to elevation and effects of
the trade winds that combine to affect rainfall patterns.
The trade winds blow consistently from the east and
northeast, off of the Atlantic Ocean onto northeastern
South America, with wind speeds averaging from 3–4
m per second. Due to orographic effects, the eastern-
most escarpments of the mountains of the Guiana
Shield are generally localities of increased precipitation
where these moisture-laden winds meet the slopes
(Clarke et al. 2001). Seasonal oscillations of the
Intertropical Convergence Zone (ITCZ) also bring
variations in rainfall as the locations of low pressure
zones near the equator change (Snow 1976). Varying
primarily by latitude, one or two rainy seasons result
from shifts in the ITCZ. The heaviest rains usually
occur between May and August, whereas the rainy
season running from December to January is shorter
and less intense, with rains that do not penetrate as far
inland. Even during most dry seasons, frequent storms
provide adequate moisture to allow evergreen tropical
moist forests to persist in most low elevation parts of
the region.
Biological Diversity
The variety of landscapes of the Guiana Shield
includes sandstone tepuis, granite inselbergs, white
sands, seasonally flooded tropical savannas, lowlands
with numerous rivers, isolated mountain ranges, and
coastal swamps, each supporting a characteristic
vegetation (Huber 1995b, Huber et al. 1995). This
variety accounts for a great deal of the high diversity
and endemicity of the Shield’s biota. The highlands of
the Shield have a flora and fauna with numerous
endemic species. Some tepui endemic species occur as
low as 300 m in elevation, with increasing numbers by
1500 to 1800 m, and fully developed communities
occurring by 2000 m. Few if any plant or animal
specimens have been collected from most medium to
high elevation areas of the Guiana Shield. Many parts
of the Shield are poorly explored, including parts of
Brazil north of the Amazon River, much of eastern
Colombia, and the southern parts of Venezuela,
Guyana, Suriname, and French Guiana.
Conservation
With the exceptions of the populated localities such
as Puerto Ayacucho, Ciudad Guayana, Ciudad Bolı-
var, Boa Vista, Georgetown, Paramaribo, Cayenne, the
agricultural coastal areas, and open areas like the
Rupununi Savanna, the environment of the Guiana
Shield has benefited from limited access and low
population densities, although this same isolation has
hindered biodiversity research. Estimates vary, but
much of the vegetation is still relatively undisturbed by
human activities. Recently, however, the pace of
disturbance has greatly increased. Current threats to
the environment include large-scale logging by Asian
and local companies, large- and small-scale gold and
diamond mining, oil prospecting, bauxite mining,
hydroelectric dams, wildlife trade, and population-
related pressures such as burning, grazing, agriculture,
and the expansion of towns and villages. Taken
together, these impacts have begun to take their toll,
with vast areas vulnerable to increasing disturbance a
fact easily observed by using Google Earth and
‘‘flying’’ over the area.
The status of conservation efforts varies by country.
Throughout the Guiana Shield, many areas that are
designated as protected or otherwise restricted are
often only ‘‘paper’’ parks because of a lack of
infrastructure, funds, and will to actually protect the
areas. Over the last four decades, Venezuela has
established seven national parks, 29 natural monu-
ments, and two biosphere reserves covering about
142,280 km2, more than 30% of its share of the
NUMBER 17 3
Guiana Shield (Funk & Berry 2005). In Guyana, the
progress of conservation efforts has been slower, with
the only substantial protected area being Kaieteur
National Park, its 627 km2 comprising about 3% of
the country’s area (Kelloff 2003, Kelloff & Funk
2004), with additional reserves under consideration.
Guyana’s 3710 km2 Iwokrama forest (Clarke et al.
2001) has parts listed as reserves, but overall it is
dedicated to sustainable use; unfortunately, logging
has begun, and the section of the road from Boa Vista,
Brazil, to Georgetown, Guyana, that runs through
Iwokrama is about to be paved. Suriname’s protected
areas system includes one national park and a network
of 11 reserves, totaling almost 20,000 km2, over 12%
of its total area. This includes the recently created
16,000 km2 Central Suriname Nature Reserve, a
UNESCO World Heritage Site that joined and
expanded three existing reserves (see http://www.
stinasu.com). French Guiana has no officially desig-
nated protected areas, but 18 proposed sites total 6710
km2, about 7.5% of its area (Lindeman & Mori 1989).
The natural areas of Venezuela and Guyana are
currently under the most anthropogenic pressure,
while those of French Guiana are probably less
threatened.
The Shield encompasses part or all of six countries
with six different governments, five official languages
and many more indigenous languages. Cooperation is
sometimes hampered by border disputes, illegal cross-
border activities involving gold and wildlife, and a lack
of interest by governments that are located far away.
The implementation of conservation practices is
further complicated by many issues concerning the
indigenous peoples of the region. All of these
challenges will have to be overcome on the way to
designing and maintaining a viable reserve system for
the Guiana Shield. However, it is critical that we gain
an understanding of the flora and fauna of the Shield
area so that decisions can be made on critical areas that
have high priority for conservation and so data can be
collected from areas that might ultimately be de-
stroyed. Because it is an ancient, fairly isolated
geological area, it is rich in endemic plant and animal
taxa, with many more likely to be discovered with
additional exploration. In addition, because this area
has been long neglected by biologists, it is often an area
of ‘‘inadequate information’’ for many biodiversity
analyses.
This volume contains the fishes from the Guiana
Shield, when paired with the previously published
Checklist of the Terrestrial Vertebrates of the Guiana
Shield (Hollowell & Reynolds 2005), we can examine
the size and scope of Vertebrates, an important
monophyletic group, known to inhabit the Guiana
Shield. Table 2 lists the vertebrate groups and their
sizes. The two checklists include a total of 53 orders,
189 families, 1190 genera, and 301 species. A large
percentage of the species (38%) are contributed by the
fishes listed in this volume.
Figure 2 compares the vertebrate diversity across the
major political areas of the Guiana Shield and shows
the species turnover between different areas. Themammals and reptiles have the most similar fauna
across the Shield with a 58% and 53% overlap,
respectively, between French Guiana (the extreme east)
and Venezuela-Amazonas (the extreme west). Fish and
birds have the least (24% and 10%, respectively). With
fishes this can probably be explained by the fact that
the headwaters of the rivers in the east are widely
separated from those of the west. The rivers of theShield in the west (Venezuela) have their source in the
Venezuelan Guayana and the Andes, in the central
portion (Guyana) the Essequibo drains mainly from
the Acari Mountains which lie on the border with
Brazil as does the Corantijn River (border between
Guyana and Suriname). To the east, rivers such as the
Maroni and Oyapock drain from the Tumuk-Humak
Mountains. The bird diversity percentage of ‘turn over’was surprisingly small until one realizes that there are
very different flyways that go across the eastern and
western parts of the Shield. When the three major
avenues of vertebrate mobility are examined (land, air,
water), it seems that the land provides the most stable
species make up and the air and water provide the
least. Could this have anything to do with the resulting
high species diversity of the birds and fishes?In the wider scope of biological understanding, the
goal of checklists of this type is to understand diversity
in terms of the spatial, evolutionary, and ecological
settings of physical environments, rather than simply
by political boundaries. The assembly of these lists is a
step toward considering the fauna in terms of the
geological entity of the Guiana Shield. Future studies
will include the analyses of animal community com-position on finer landscape scales, using developing
abilities to produce customized checklists for research
and conservation with Geographic Information System
(GIS) technologies drawing upon comprehensive data-
bases that include georeferenced museum specimen
records.
Biological Diversity of the Guiana Shield (BDG)
The ‘‘Biological Diversity of the Guiana ShieldProgram’’ (BDG) is a field-oriented program of the
Table 2.—Number of vertebrate taxa at different ranks.
Orders Families Genera Species
Amphibians 2 13 59 269
Reptiles 3 22 119 295
Mammals 11 35 143 282
Birds 22 70 493 1004
Fishes 15 49 376 1168
Total 53 189 1190 3018
4 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
National Museum of Natural History that began in
1983 (federally funded since 1987). The goal of the
BDG is to ‘‘study, document and preserve the
biological diversity of the Guiana Shield.’’ Most of
the program’s field work has taken place in Guyana,
but data analyses cover the majority of the Shield. In
Guyana, the BDG operates under the auspices of the
University of Guyana (UG). The BDG program is
designed to provide specimens and data to address
questions about many groups of organisms from
locations across the Shield. Information from BDG
collections and from other herbarium collections is
used to produce checklists, vegetation maps, floristic
and faunistic studies, revisions, and monographs. The
data generated from these studies are used to ask
questions about the make up of Guiana Shield
biological diversity, such as species turnover rates,
surrogate taxa, and areas of high diversity. Finally, the
BDG is exploring practical applications of the data
that have been collected through regular collaborations
with conservation and government agencies.
In addition to collecting and research, the BDG
Program trains students and scientists in both the
U.S.A. and Guyana, assists in their research, and has
established and helped to maintain collections. Over
the years several events have been hosted in Guyana,
including two Amerindian training courses, two bird
preparation courses, two plant identification courses, a
variety of lectures at the University and public venues,
and a public scientific symposium on the biological
diversity of Guyana. We also offer training opportu-
nities; nearly every year since 1987 the Program has
hosted at least one Guyanese student or UG staff
member at the Smithsonian. Many have participated in
the Natural History Museum’s Research Training
Program or the SI/MAB training courses. BDG
worked with the University of Guyana to raise funds
from the Royal Bank of Canada to construct a new
building, the ‘‘Centre for the Study of Biological
Diversity,’’ located on the campus of UG. More
recently, we worked with UG to raise funds from
USAID to build an extension on the original building.
Figure 2. A comparison of the species lists of the political areas of the Guiana Shield Region.
NUMBER 17 5
The Centre houses collections and research space,
provides a library, and houses a Geographic Informa-tion System (GIS) facility. The goal of the Centre is to
combine research, education and conservation in the
study of biological diversity. The Centre is funded from
outside grants, but the staff is part of the University.
Currently, the plant database maintained by BDG has
161,108 specimen records and all sheets have been
barcoded. The BDG Program is working to make its
data available to the scientific community. Thecollections are being mapped using ArcMap and then
displayed on Google Earth as place marks. The
project, Georeferencing Plants of the Guiana Shield is
available on the Department of Botany public website
(http://botany.si.edu/bdg/georeferencing.cfm).
Acknowledgments
Special thanks go to the University of Guyana andthe Guyana EPA who have consistently supported our
efforts, including Mike Tamessar, Indarjit Ramdass,
and Philip da Silva, as well as past and present staff
members of the Centre for the Study of Biological
Diversity, in particular Calvin Bernard. This is number
153 in the Smithsonian’s Biological Diversity of the
Guiana Shield Program publication series.
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Arensica de la comunidad de Monochoa (Region de
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P. E. Berry, & B. K. Holst, general eds. Missouri Botanical
Garden, St. Louis, 306 pp.
Clarke, H. D., V. Funk, & T. Hollowell. 2001. Using checklists and
collections data to investigate plant diversity. I: a comparative
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Burgess, D. P. Faith, J. F. Lamoreux, G. Kier, R. J. Hijmans,
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French Guiana).—Contributions from the United States
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Shield. Oxford University Press, New York, 246 pp.
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Venezuelan Guayana. Vol. 1: Introduction. J. A. Steyermark,
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general eds. Missouri Botanical Garden, St. Louis.
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map of Guyana. Biological Diversity of the Guianas Program,
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Edipucrs, Porto Alegre, Brazil, 729 pp.
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6 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
FISHES OF THE GUIANA SHIELD
RICHARD P. VARI and CARL J. FERRARIS, JR.
History
A vast complex of wetlands, lakes, streams, and
rivers drains the broad savannas, dense rainforests,extensive uplands, and tepuis of the Guiana Shield.
Early European explorers and colonists were impressed
not only by the many unusual fish species dwelling in
these water systems but also by the diversity of the
ichthyofauna. Accounts of fishes from those drainage
systems commenced with descriptions by pre-Linnaean
naturalists (e.g., Gronovius 1754, 1756) based on
specimens returned to Europe, with Linnaeus (1758)formally describing a number of these species.
Much of the early descriptive activity involving
fishes of the Guiana Shield centered on the ichthyo-
fauna of British Guiana (5 Guyana). Commentaries
on fishes from that colony by Bancroft (1769) and
Hilhouse (1825) preceded the formal description of
catfish species from the Demerara by Hancock (1828).
Cuvier and Valenciennes summarized the availableinformation on the ichthyofauna of all of the Guianas
in their series entitled Histoire Naturelle des Poissons
that documented the fish species known worldwide to
science to that time; with the catfishes being the first of
the groups inhabiting the shield discussed by those
authors (Cuvier & Valenciennes 1840a, b). These and
the other treatments of that era were, however, largely
opportunistic accounts based on scattered samplesreturned to Europe rather than derivative of focused
studies on the fish fauna of any region on the shield.
Consequently, the scale of the species-level diversity of
that ichthyofauna remained unknown and underap-
preciated.
Indications of the scale of the richness of the fish
fauna inhabiting the rivers of the Guiana Shield
commenced with the expeditions of the Schomburgkbrothers, Richard and Robert. In a remarkable
endeavor for that era, Robert collected fishes from
1835 to 1839 both in the more accessible shorter
northerly flowing rivers of the Guianas and through
portions of the Rıo Orinoco and Rıo Negro and the
Rio Branco. Drawings of fishes prepared during
Schomburgk’s travels across the shield served as the
basis for 83 species accounts in Jardine’s ‘‘Naturalist’sLibrary’’ (1841, 1843), including the formal descrip-
tions of a series of species. Unfortunately, the speci-
mens that were the basis for the drawings were not
preserved, and some illustrations combined details of
more than one species. Subsequent expeditions by the
Schomburgks traversed portions of what are now
Guyana, Venezuela, Suriname, and Brazil and yielded,
what was for that time, large numbers of fish specimens.In a series of publications Muller & Troschel (1845,
1848, 1849) recognized 141 species in the Schomburgk
collections and provided the first detailed illustrations
of fishes from South American freshwaters.
Diverse factors resulted in a lag in the state of
knowledge of the fishes inhabiting many portions of
the shield, with the comparative difficulty in accessi-
bility to inland regions clearly a paramount issue for
many areas. Supplementing that impediment were a
series of misadventures that bedeviled collectors who
sampled the fish fauna of the western portions of the
shield. Alexandre Rodriques Ferreira headed an
expedition that explored a significant portion of the
Rio Negro basin, commencing with a major collecting
effort through the Rio Branco system in 1786 (Ferreira
et al. 2007:12). Confounding Ferreira’s attempts to
publish his results were a string of unfortunate events
that culminated with the 1807 invasion of Portugal by
Napoleonic forces and the seizure and shipment of
Ferreira’s collections to Paris. Ferreira’s report on
animals from the Rio Branco region remained unpub-
lished until long after his death, and even then, only
parts appeared in print (see references in Ferreira
1983). In two expeditions between 1850 and 1852,
Alfred Russel Wallace (of Natural Selection fame)
collected over 200 species of fishes throughout the Rio
Negro basin including rivers draining the shield.
Wallace’s collections were lost with the sinking of the
ship returning him to England. Nonetheless, his field
sketches (Wallace 2002) document that the lost
collection included a number of species of fishes then
unknown to science (Regan 1905a, Toledo-Piza et al.
1999, Vari & Ferraris 2006).
An accelerating pace of ichthyological collecting
across many portions of the Guiana Shield during the
latter part of the nineteenth century resulted in the
discovery and description of numerous species. These
collections also documented the presence on the Shield
of many species originally described from elsewhere in
cis-Andean South America. Notwithstanding those
advances, the information was dispersed through
revisionary (e.g., Regan 1905b, c) and monographic
studies (e.g., Eigenmann & Eigenmann 1890), general
ichthyofaunal summaries of regions on the shield (e.g.,
Pellegrin 1908), and species descriptions in multiple
languages.
Exceptions to this pattern of scattered publication
were limited to a handful of papers focused on subsets
of the ichthyofauna from comparatively small regions.
Among the more notable of these were the analysis of
the catfishes of Suriname (Bleeker 1862), discussions of
the fishes present in portions of French Guiana
(Vaillant 1899, 1900), and a semi-popular overview of
the fishes of French Guiana (Pellegrin 1908). Compen-
dia of the freshwater fish species known from
individual colonies, countries, or regions were not
developed, let alone summaries of the fishes inhabiting
in the numerous streams, rivers, and lakes across the
shield. The dispersed literature prevented an appreci-
ation of the scale of the diversity of the shield
ichthyofauna.
The first overview of the freshwater fishes of
northeastern South America, including the Guiana
Shield, was Eigenmann’s (1912) treatise on the
freshwater fishes of British Guiana. Although Eigen-
mann sampled the fish fauna of only a comparatively
small section of British Guiana, his collections were
extensive for that era. In a series of papers, he and his
students described 128 new species from those collec-
tions (Eigenmann 1912:133). Eigenmann’s monograph
included data from his own collections, information
from the literature, and records of fishes that
originated on the Shield in various museums. Summary
tables (Eigenmann 1912:64) detailed the fish species
known from ten subunits that fall, at least in part,
within the boundaries of the Guiana Shield (Rıo
Orinoco basin, ‘‘West Coast’’ of British Guiana
[5 Barima River basin], Rio Branco basin, Rupununi
River, Lower Essequibo River, Lower Potaro River,
Demerara River, Dutch Guiana [5 Suriname], and
French Guiana [5 Guyane Francaise]).
Eigenmann (1912) reported 493 species from those
ten geographic units; a total that was in excess of the
species reported to that time from the rivers of the
Shield. These additional species were a function of two
factors. His total included all of the fish species then
known to inhabit the Rıo Orinoco; however, that vast
river system extends far beyond the Shield boundaries
with approximately only 40% of that watershed
overlying the Shield. Many of the fish species known
at the end of the first decade of the twentieth century
from the Rıo Orinoco basin originated in the llanos
(savannas) of the north central and western portions of
the basin. Aquatic habitats and the fish faunas in these
floodplain savanna settings differ dramatically from
the ecosystems and fish communities of the more
rapidly flowing rivers that drain the forested northern
slope of the Guiana Shield. Further inflating Eigen-
mann’s species total was his inclusion of some
primarily marine forms. Such species penetrate the
lower reaches of the rivers draining the Guianas during
periods of low river flow and consequent increased
estuarine salinity. Few, if any, of these species are likely
to range upriver onto the Shield even during the height
of the dry season.
The decades since Eigenmann’s monograph have
seen numerous ichthyological collecting expeditions in
many systems on the Shield. Two wide-ranging and
productive collecting endeavors through that region
during the first half of the twentieth century remain
relatively poorly known. John Haseman, who collected
throughout the Rio Branco basin and the southern-
most portion of the Rupununi River system in 1912
and 1913, made the first of these. Haseman deposited
these extensive collections in the Naturhistorisches
Museum in Vienna where he studied them for a year in
collaboration with Franz Steindachner. Nonetheless,
only one major publication based on those collections
was published (Steindachner 1915), most likely because
of the onset of World War I, disruptions during and
immediately after the conflict, and the death of
Steindachner soon after the cessation of hostilities.
Various revisionary studies in recent decades incorpo-
rated subsets of Haseman’s collection; nonetheless,
much of the material is yet-to-be analyzed critically.
The second collector, Carl Ternetz, sampled fishes
through the Rio Negro, Rıo Casiquiare, and Rıo
Orinoco basins during 1924 and 1925. Myers
(1927:107) remarked that the collection was ‘‘a
magnificent series of fishes, most of them hitherto
unexplored systematically by an ichthyologist.’’ Not-
withstanding the description of some new species
collected by Ternetz in rivers of the shield by Myers
(1927) and other authors and the use of portions of
that collection in some studies (e.g., Myers & Weitz-
man 1960), most of the material remains unstudied,
even after its transfer from Indiana University to the
California Academy of Sciences.
The 1960s brought a resurgence of major ichthyo-
logical collecting efforts in many of the river systems
on the shield (e.g., the Brokopondo Project; Boeseman
1968:4), with the pace of these endeavors accelerating
during recent decades. A compendium of these
collecting efforts lies beyond the purpose and scope
of this paper; however, as summarized in the next
section many of these expeditions were integral to
checklists, regional revisionary studies, and summaries
of the ichthyofauna in river basins or regions of the
Shield.
State of Knowledge of the Shield Fish Fauna
The nearly ten decades since the preparation of
Eigenmann’s 1912 magnum opus saw numerous
publications on fishes of the Guiana Shield. Many
were revisionary studies of genera or families whose
ranges extend far beyond the limits of the Shield, often
across major portions of cis-Andean South America
and in some instances into trans-Andean regions or
occasionally Central America. Other publications were
restricted to the members of a genus, subfamily, or
family from a country within the Shield region (e.g.,
Suriname: Boeseman 1968, Nijssen 1970, Kullander &
Nijssen 1989) or across a major portion of that area
(e.g., Boeseman 1982). Relatively few of these papers
involved broad surveys of an entire ichthyofauna in a
river system or country on the Shield with those that
10 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
did so summarized below arranged by the geographic
subdivisions in the checklist. Many include associated
ecological and life history information for fish com-
munities and individual species.
Brazil, Para (PA). The single publication of note
from this region is Ferreira (1993) that summarized the
results of intensive collecting efforts at sites within the
Rio Trombetas, one of the major northern tributaries
of the Amazon River east of the Rio Negro.
Brazil, Roraima (RO). Ferreira et al. (1988) summa-
rized the ichthyofauna at several closely situated
localities in the Rio Mucajai, a tributary of the Rio
Branco. More recently, Ferreira et al. (2007) provided
detailed information on the ichthyofauna across the
expanse of the Rio Branco basin, supplemented by
numerous color photographs, discussions of habitats,
and comments on the anthropogenic impact on the
aquatic systems within the basin.
French Guiana (FG). French Guiana has the most
intensely studied ichthyofauna of any portion of the
Guyana Shield. The first attempt to summarize
information on the fishes of the entire department
was that of Puyo (1949). Gery (1972) followed up with
studies of the characiforms (his characoids) from the
Guianas with a particular focus on French Guiana.
Planquette et al. (1996), Keith et al. (2000), and Le Bail
et al. (2000), in a groundbreaking series of publica-
tions, brought together information on the spectrum of
the freshwater fish fauna in that department. Each
species account includes a description, illustration, and
comments on its biology. Distributions within and
beyond French Guiana are discussed, and the sites of
occurrences of the species in the department are
plotted.
Guyana (GU). Notwithstanding the title of the
publication, Eigenmann’s (1912) monographic study
was based primarily on collections from the northern
portions of Guyana, in particular the Potaro River and
lower courses of the Essequibo and Demerara rivers,
albeit with that data supplemented with information
from the literature. Hardman et al. (2002) reported on
the fish fauna captured at Eigenmann’s collecting
localities nine decades after his expedition and provid-
ed a checklist of the 272 species collected in that survey.
Lowe (McConnell) (1964) included lists of fishes from
the southern most reaches of the Essequibo River
system along with observations on their ecology and on
the movements of various species during the yearly
flood and drought cycles. Watkins et al. (2004)
provided a summary of the fishes of the Iwokrama
Forest Reserve.
Suriname (SU). The ichthyofauna of Suriname
remains relatively poorly documented, with the listing
of the freshwater fishes in the country by Eigenmann
(1912:64–73) largely derived from literature informa-
tion. Boeseman (1952, 1953, 1954) supplemented
Eigenmann’s listing. Ouboter & Mol (1993) presented
the most comprehensive published list of the freshwater
fishes of Suriname. Kullander & Nijssen (1989)
published a detailed analysis of the cichlids of
Suriname.
Venezuela, Amazonas (VA). The state of Amazonas,
Venezuela, includes portions of the south-flowing Rıo
Negro of the Amazon basin, the north-draining Rıo
Orinoco and the entirety of the intervening Rıo
Casiquiare. Mago-Leccia (1971) produced the first
summary of the fishes of the Rıo Casiquiare. Lasso
(1992), Royero et al. (1992), and Lasso et al. (2004a,
2004b) provided information on the fish faunas of
various river systems within the state.
Venezuela, Bolivar (BO). A series of studies treated
the fish fauna of several right bank tributaries of the
Rıo Orinoco that drain the northern slopes of the
Guyana Shield. These included summaries of species in
various basins, with those listings supplemented in
some instances by information on fish biology and
distribution. Significant publications on the ichthyo-
fauna of the Rıo Caroni were published by Lasso
(1991) and Lasso et al. (1991a, b) and for the Rıo
Caura by Lasso et al. (2003a, b), Rodrıquez-Olarte et
al. (2003), and Vispo et al. (2003). The Rıo Cuyuni, a
western tributary of the Essequibo River, drains the
eastern portions of the Shield in the state of Bolivar.
Machado-Allison et al. (2000) summarized the fish
fauna of the Venezuelan portions of that river system.
Lasso et al. (2004a) and Girardo et al. (2007) provide
supplemental information on the ichthyofauna of that
drainage basin.
Ichthyofaunal Richness
This checklist of fishes known from the water
systems of the Guiana Shield includes 1168 species.
Included in that total are representatives of 376 genera,
49 families, and 15 orders. Five orders are dominant in
terms of number of species living on the shield and
account for 96.7% of the species (Characiformes, 478
species and 41.0%; Siluriformes, 425 species and 36.4%;
Perciformes, 126 species and 10.8%; Gymnotiformes,
52 species and 4.5%; Cyprinodontiformes, 47 species
and 4.0%). This sum of 1168 species attests to the
dramatic improvement of our knowledge of the
freshwater fish fauna on the Shield in slightly less than
a century since Eigenmann (1912) documented fewer
than 500 species from that region. The 1168 species are
approximately 4.1% of the 28,400 fish species recently
estimated to be present in all marine and freshwater
systems worldwide (Nelson 2006), a percentage that
amply testifies to the striking diversity of the ichthyo-
fauna within that region. All the more noteworthy is
the species-level richness of the ichthyofauna within the
context of the overall Neotropical freshwater fish
fauna. According to a recent summary, approximately
5000 species of freshwater fishes occur across the
NUMBER 17 11
entirety of Central and South America (Reis et al.
2003). Thus, the drainage systems of the Guiana Shield
are home to approximately 23% of the freshwater fish
species that occur across the vast expanse between
southern South America and the southern border of
Mexico. Many factors contributed to the Shield region
being a repository of freshwater ichthyological diver-
sity, with a few particularly worthy of comment.
Physiography
The Guiana Shield is the ancient Precambrian
Guianan formation resulting from the uplift of the
underlying craton (Gibbs & Barron 1993) and demon-
strates attributes that generally lead to high levels of
biodiversity: geological diversity, a topographically
variable landscape, and transitions between ecosystems
(Killeen et al. 2002). Overall the region has a primarily
low to somewhat hilly physiography, albeit with some
abrupt changes in topography in the regions proximate
to the tepui formations that extend across much of the
region in an approximately east to west alignment.
Some river valleys have marked shifts in topography
with resultant waterfalls, rapids, and riffles that
increase the complexity of drainage system structure.
These topographic factors result in multiple aquatic
habitats with differing levels of physical complexity. At
one extreme are the lentic waters of swamps, wetlands,
channels, and lowland rivers. With increasing gradient,
the drainage systems progress through variably flowing
waters interrupted by higher energy settings such as
riffles and isolated lower scale rapids. Finally, there are
regions of greater gradients with rapidly flowing waters
and major repetitive rapids and waterfalls. To the
degree that differences in stream structure directly
correlate with elevational gradients, there also occur
differences in water temperatures.
Water Chemistry
Physical river system attributes are the most obvious
manifestation of variation in aquatic systems across the
Shield but represent a distinct subset of the spectrum of
factors that contribute to shifts in the composition and
relative biomass of fish communities across that region.
Complementing diversity in river structure are varia-
tions in water chemistry that occur not only between
but also within drainage basins across the Shield. Three
major water types occur in the tropics. The first of
these are white waters carrying nutrient-rich sediment
loads for at least part of the year and which, despite
their name, are actually brown (e.g., Rio Branco;
Goulding et al. 2003:42). A second major group are
clear water streams and rivers, including those draining
many regions of the eastern portion of the Amazonian
portion of the Shield (Para and Amapa; Goulding et al.
2003:43). Finally there are acidic black water rivers
that drain heavily leached soils where decomposing
plant matter produces high levels of fluvic and humic
acids but with the water poor in dissolved solids and
nutrients (e.g., Rio Negro; Goulding et al. 2003:44;
Savanna Belt rivers in Suriname, Ouboter & Mol
1993:134). Such water type differences occur both at
the level of major river systems and at a much smaller
scale within some river basins (Arbelaez et al. 2008).
Admixtures of water types resulting from within basin
water type differences yield conjoined drainages with
variably intermediate chemistries (e.g., downriver of
where the black water Rio Negro empties into the
white water Rio Solimoes at Manaus, Goulding et al.
2003:44; or where the white water Rupununi River
empties into the black water upper Essequibo River,
Watkins et al. 2005:40).
Species of freshwater fishes demonstrate physiolog-
ical, morphological, and behavioral adaptations that
often allow them to specialize for life in particular
water types but often simultaneously exclude them
from other water types. Some species or genera are,
therefore, more common in, or effectively limited to
waters with particular chemical characteristics (Lowe-
McConnell 1995). Exemplifying this situation are
acidic black waters such as those in the Rio Negro
that appear to be the primary, if not exclusive, habitat
for some fish species (Goulding et al. 1988, table 2).
These acidic waters are at the same time apparently
inimical to other species and some genera notwith-
standing the presence of these taxa in adjoining rivers
of different water types (Goulding et al. 1988:98). The
different water types also differ in degrees of primary
productivity and dependence on detritus-based energy
systems (De Jesus & Kohler 2004), factors that further
impact fish diversity and community composition.
Allochthonous Influences
Terrestrial habitats further influence freshwater
systems via the shift of nutrients and organic matter,
with the often-substantial input into water bodies
mediated by both water and wind. Terrestrial to
freshwater inputs and their impacts on aquatic systems
span a broad spectrum of scale. At one extreme, both
continuing upland erosion and periodic floods trans-
port dissolved and particulate matter into water bodies
(Sioli 1975, Polis et al. 1997). Alternative forms of
riparian vegetation also affect the amount and types of
allochthonous materials, including detritus, seeds and
fruits, and animals (primarily terrestrial insects) input
into the aquatic food web via runoff and wind. The
input of allochthonous detritus into Neotropical water
systems supports particularly large populations of
detritivorous fishes (Flecker 1996). Most notably,
many species and groups of fishes specialize on
exploiting allochthonous seeds, fruits, and insects, with
variation in the input of these items impacting the
12 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
composition of ichthyofaunal communities (see Gould-
ing et al. 1988, Boujard et al. 1990).
At the other end of the scale spectrum, alternate soil
types in conjunction with factors such as rainfall
regimes and temperature also support dramatically
different plant communities ranging from dense rain-
forests through open savannas. Physical attributes of
differing marginal plant communities directly influence
fish community composition and structure in the water
bodies that they border. Most obvious of these effects
is differential shading by riparian forests, an influence
that is particularly significant in terms of the species
dwelling in streams and smaller rivers. Marginal
vegetation and submerged macrophytes affect the
physical complexity of water bodies and thus the
composition of the resident ichthyofaunas in various
fashions. Most noteworthy among these are differential
inputs to the aquatic systems in the amount and type of
woody and leafy debris, variation in the submerged
portions of overhanging terrestrial plants along water
margins, and differing amounts and types of sub-
merged emergent vegetation. Synergy of drainage
structure and energy, differences in water types,
variation in associated riparian animals and plants,
and differences in input of nutrients yield dramatically
different fish communities. That variation at the local
level is a major contributor to the overall species-level
richness of the ichthyofauna across the totality of the
Shield.
Drainage System Interconnections
Physiological and physical factors closely tie fresh-
water fishes to drainage patterns. Historical separa-
tions of, and associations between, drainage systems
thereby contributed to the present day distributions of
many species on the Shield and the richness of that
fauna. Notwithstanding the tectonic quiescence of the
Guiana Shield for over 550 million years, the highlands
resulting from the uplift of the craton underwent
progressive erosion of the sedimentary layers overlying
that base (Gibbs & Barron 1993). The pronounced
degree of endemicity in many of the basins across the
Shield is indicative of their long isolation, with factors
including differences in water types being influential in
this regard. Nonetheless, there have been changes in
the water flow patterns on and along the margins of the
Shield that influenced the present composition of the
ichthyofauna in those systems.
At the large scale, tectonic events resulted in the
broader details of the present Orinoco and Amazon
basins, both of which contribute to the Shield
ichthyofauna. A large paleo-drainage encompassing
much of the present Orinoco and Amazon basin
drained north into the Caribbean Sea approximately
at the present location of Lago Maracaibo. Tectonic
events at the end of the Miocene resulted in separation
of the Amazon from Orinoco basin (Hoorn 1994).
Another consequence of these changes was the shift
eastward of the mainstream Orinoco along the
northern boundary of the Shield to its present mouth
slightly north of the northeastern margin of the Shield.
This dramatic realignment led to its capture of
drainages flowing from the northern slopes of the
Shield. The shift of the mainstream Amazon to it
present mouth similarly resulted in the capture of the
southern draining rivers of the Shield. Disrupting the
continuity between many of those freshwater systems
for varying periods were subsequent marine transgres-
sions into the eastern portions of the Amazon valley.
Superimposed on these large-scale drainage pattern
changes were long-term, often pronounced, climate
changes through the region (Baker et al. 2001) with
resultant sequential contraction and expansion of
suitable aquatic habitats. This combination of hydro-
graphic and climatic changes resulted in disruptions, in
some instances repeated disruptions, of previously
continuous species ranges, thereby setting the stage
for subsequent speciation.
The complex hydrological history of the drainages
on the Shield involved not only division of previously
continuous drainages but also in new connections
between various river systems. Connectivity resulted
from landscape tilting during uplift events and via
headwater stream capture. Those events permitted, and
continue to permit, movements of fishes between what
have been isolated basins on the Shield. Subsequent
disruptions of connectivity provided an opportunity
for the evolution of species. Such past connections may
account for unusual present day distribution patterns
previously highlighted by some authors (e.g., Armbrus-
ter 2005). Prominent among these possible connections
was the river hypothesized to have drained directly
from the south slope of the Guiana Shield into the
Atlantic Ocean through what is now northeastern
Guyana (Hammond 2005:137). Faunal similarities and
close phylogenetic relationships among included fish
species also point to possible past associations between
the upper Rıo Caroni of the Rıo Orinoco basin and the
Rıo Cuyuni, a western tributary of the Essequibo River
(Lasso et al. 1991a, Sabaj Perez & Birindelli 2008). An
interconnection between the Amazon and the upper
portions of the Maroni River was proposed by
Cardoso & Montoya-Burgos (2009) who also proposed
that temporary connections between adjoining river
systems during periods of lower sea levels permitted
dispersal of freshwater fishes along the coasts of the
Guianas.
The paramount example of an extant interconnec-
tion between major river systems on the Shield, or
indeed across the continent, is the Rio Casiquiare. This
over 300 km long natural canal connects the Rıo Negro
of the Amazon River basin with the upper portions of
the Rıo Orinoco. This unusual drainage begins as a
NUMBER 17 13
bifurcation of the upper portion of the Rıo Orinoco
and represents an ongoing capture of the upper portion
of the latter river system by the Rio Negro (Sternberg
1975, Winemiller at al. 2008). Despite its substantial
size at the divergence, where it is approximately 100 m
wide, and the fact that it carries a significant portion of
the total flow of that portion of the Rıo Orinoco, the
Rıo Casiquiare does not provide unimpeded transit for
all species of fishes between those basins. Lack of
interbasin species panmixis is, in part, a function of the
fact that the Rıo Casiquiare conjoins headwaters
habitats inhabited by a subset of the species resident
across the entirety of each basin. Equally, or perhaps
more significantly, the gradient in water types along
the course of the Rıo Casiquiare acts as a partial filter
that impedes movement of many fish species (Wine-
miller et al. 2008), thereby maintaining differences in
ichthyofaunal composition between the upper Rıo
Negro and upper Rıo Orinoco.
Although the Rıo Casiquiare is the most notable
connection between major rivers on the Guiana Shield
and the only year-round continuity, some degree of
seasonal connectivity occurs between the upper reaches
of the Rio Branco and the southern most portions of
the Essequibo River in the Rupununi Savannas of
southwestern Guyana. That region is an expansive
floodplain where the headwaters of the Rupununi
River (Essequibo basin) and the Takutu and Ireng
rivers (both components of the Rio Branco of the
Amazon River basin) come into close proximity. This
proximity and varying degree of continuity of the
headwaters of these rivers across a vast flooded plan
during high water periods [Lowe (McConnell) 1964,
Watkins et al. 2005] facilitate movement of at least
some fish species between the headwaters of the Rio
Branco and Essequibo River basins. Although such
movements potentially enrich the ichthyofaunas of
each of those river systems, they do not add to the
overall richness of the fish fauna of the shield.
Future Directions
Notwithstanding the series of papers listed under
‘‘State of Knowledge of the Shield Fish Fauna’’ and
many other publications, the continuing discovery and
description of freshwater species from water systems on
the Shield testifies to the incomplete state of our
knowledge of that fish fauna. The primary impediment
is the lack of exhaustive ichthyological collecting across
that vast region, with many river systems still
effectively unsampled (e.g., the upper Mazaruni River
in Guyana; Taphorn et al. 2008). Headwater tributary
streams, deep mainstream channels, and difficult-to-
sample habitats such as swamps and rapids remain
unsampled or poorly sampled even in those drainage
systems that have been the subject of ichthyological
collecting efforts. Those habitats and some largely
ichthyologically unexplored drainage systems hold the
greatest promise as sources of undescribed species;
however, we must not lose sight of the fact that areas
that have been long the foci of ichthyological sampling
continue to yield new species and are deserving of
continued attention. Recent monographic studies of
speciose genera and families of Neotropical freshwater
fishes demonstrate that collections in museums, uni-
versities, and research institutions of North and South
America and Europe house numerous species as of yet
unknown to science. Indeed in many groups, the vast
majority, if not all, of recently described species were
already represented in collections and awaited discov-
ery, often for many decades.
Thorough sampling of the freshwater fish fauna is
vital as is the thorough examination of materials in
collections, but equally or perhaps more important for
furthering our knowledge of the fishes on the Shield are
comprehensive revisionary studies of all groups of
fishes represented in that ichthyofauna. Such in-depth
studies are critical given our inadequate understanding
of the species-level diversity of many Neotropical
freshwater groups (Vari & Malabarba 1998). Inclusive
revisions of complex groups of Neotropical freshwater
fishes have repeatedly demonstrated that the sum of
long recognized species within a genus typically
underestimates the actual number of species in a taxon,
sometimes to a pronounced degree. An example is the
67 species now recognized in Creagrutus; a total three
and one-half times the number of species (19)
recognized in the genus prior to 1994 (Harold & Vari
1994, Vari & Harold 2001, Vari & Lima 2003, Ribeiro
et al. 2004, Torres-Mejia & Vari 2005). Comprehensive
revisions similarly further the subsequent identification
of additional previously unrecognized species by other
researchers (Reis 2004).
It is impossible to estimate the degree to which the
total number of freshwater fish species summarized in
this checklist falls short of the actual count of species
dwelling on the Shield. Nonetheless, it is clear that the
rate of continued additions to this speciose fish fauna,
the many regions and habitats that have not yet been
thoroughly explored ichthyologically, and the large
numbers of groups of fishes in the region that have not
yet been exhaustively studied portend a significant
increase in the species total.
Conservation Challenges
Deleterious anthropogenic activities impact freshwa-
ter ichthyofaunas across the Neotropics (Killeen 2007),
with many affecting various portions of the Guiana
Shield and adversely influencing fish communities in
the region. Adverse impacts are pervasive across
freshwater ichthyofaunas worldwide (Millennium Eco-
system Assessment 2005, Revenga et al. 2005), with
freshwater fishes consequently the most threatened
14 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
groups of vertebrates across the world in terms of
affected species (Dudgeon et al. 2006, Chapman et al.
2008). Major impacts on freshwater ichthyofaunas of
the Guiana Shield cover the spectrum of human
activities. These include overfishing for human con-
sumption and the aquarium trade, pollution from
agricultural, domestic, industrial and mining sources,
diversion of water for agricultural, domestic, and
industrial purposes, mining within river channels,
introductions of exotic species, transplanting of native
species between separate drainage systems, deforesta-
tion within drainage basins with consequent changes in
water flow patterns and quality, increased erosion and
siltation as a consequence of development, agriculture,
and mining operations, and impoundments for hydro-
electric and irrigation systems with disruption of
migration routes for fishes.
Major advances are necessary before we approach a
definitive understanding of the species-level diversity
for the fishes inhabiting the rivers, streams, lakes, and
other water bodies on the Guiana Shield. Nonetheless,
the following Checklist can serve as a foundation for
future studies, leading to a better appreciation of the
diversity of that ichthyofauna. Such information is
vital to inform decisions by resource managers,
government agencies, and members of the public
interested in protecting both the fish fauna and the
broader aquatic communities, both of which provide
essential and important ecosystem services across the
Shield.
Species of the Guiana Shield
The Checklist includes species recognized at the time
that contributors completed their accounts (mid-2008)
with these supplemented whenever possible by infor-
mation on new species described from the Shield
through early 2009. Readers interested in further
information on the families and species included in
the listing can refer to CLOFFSCA (Reis et al. 2003).
That listing includes bibliographic information for all
fish species in Central and South America, including
those known to occur on the Guiana Shield, through
the end of 2002. References to the original descriptions
of species published post that date are listed under the
following Guide to the Checklist. The regions utilized
in the checklist correspond to those used for terrestrial
vertebrates in Hollowell & Reynolds (2005). Abell et al.
(2008) recently proposed a hydrographically delimited
series of zones for South America. The more fine-
scaled resolution of that system is potentially more
informative in terms of areas of regional endemism for
aquatic organisms. We defer, however, from applying
it to the freshwater fishes of the Guiana Shield given
the large degree of uncertainty as to distributional
limits for most species in that fish fauna.
Acknowledgments
Completion of this Bulletin was facilitated by S.
Raredon who prepared the various images that face
each section. Numerous individuals assisted our fieldefforts and studies of the fishes on the Shield over the
years, with particular thanks to L. Aguana, A.
Machado-Allison, O. Castillo, J. Fernandez, S. Jewett,
C. Lasso, H. Madarie, F. Mago-Leccia, L. Parenti, F.
Provenzano, and D. Taphorn.
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18 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
GUIDE TO THE CHECKLIST
ALEKSANDAR RADOSAVLJEVIC
The cornerstone of this project was the exhaustive
Checklist of the freshwater fishes of South and Central
America (abbreviated hereafter as CLOFFSCA; Reis et
al. 2003). Only those species with confirmed collections
within the geographic bounds of the Guiana Shield
(after Gibbs & Barron 1993; excluding outlier forma-
tions in Colombia) were extracted from the
CLOFFSCA accounts. Records from the main channel
of the Amazon River that lies at a distance from Shield
boundaries were excluded, but those from the main
channel of the Rıo Orinoco that runs closer to the
northern limit of the Shield were included. This
preliminary assemblage was updated from the litera-
ture published since CLOFFSCA’s release and the
resultant summaries sent to relevant authorities for
review. Whenever possible, the families and/or sub-
families were reviewed by the original CLOFFSCA
authors. References include papers describing new
species from the area of interest that appeared after
the completion of CLOFFSCA.
Classification in the checklist follows CLOFFSCA.
Orders and families are listed in systematic order with
genera and species listed alphabetically within families/
subfamilies and genera, respectively. Subfamilies are
limited to the Characidae (Characiformes) and Lori-
cariidae (Siluriformes). Synonyms and common names
were not included, but that information is available in
CLOFFSCA.
The Guiana Shield was divided into 11 regions (see
Table 3 for abbreviations and Fig. 1 for map) to
illustrate distributions. The four regions in Brazil
include only those parts of each state (Amapa,
Amazonas, Para, Roraima) falling within the Shield’s
boundaries. Species from rivers forming borders
between two regions were listed as occurring in both
regions. Records from river systems not totally
contained within the Guiana Shield were only included
if it could be confirmed that a particular species was
collected in the portion of the drainage basin overlying
Shield. Regional abbreviations in the checklist are
followed by a ‘‘?’’ in instances when distributions were
uncertain or questionable.
Literature Cited
(Including papers describing new species published after completion
of CLOFFSCA.)
Armbruster, J. W. 2003a. The species of the Hypostomus cochliodon
group (Siluriformes: Loricariidae).—Zootaxa 249:1–60.
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Shield (Siluriformes: Loricariidae).—Zootaxa 344:1–12.
———. 2004. Pseudancistrus sidereus, a new species from southern
Venezuela (Siluriformes: Loricariidae) with a redescription of
Pseudancistrus.—Zootaxa 628:1–15.
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with descriptions of two new species.—Neotropical Ichthyol-
ogy 3(4):549–569.
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species and a reanalysis of the phylogeny of the genera of the
Hypostominae (Siluriformes: Loricariidae).—Zootaxa 1822:
1–76.
———, & L. S. de Souza. 2005. Hypostomus macushi, a new species
of the Hypostomus cochliodon group (Siluriformes: Loricar-
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from the Rıo Caronı, Venezuela (Siluriformes: Loricarii-
dae).—Zootaxa 1731:33–41.
———, & D. C. Werneke. 2005. Peckoltia cavatica, a new loricariid
catfish from Guyana and a redescription of P. braueri
(Eigenmann 1912) (Siluriformes).—Zootaxa 882:1–14.
———, N. K. Lujan, & D. C. Taphorn. 2007. Four new Hypancistrus
(Siluriformes: Loricariidae) from Amazonas, Venezuela.—
Copeia 2007(1):62–79.
———, L. A. Tansey, & N. K. Lujan. 2007. Hypostomus rhantos
(Siluriformes: Loricariidae), a new species from southern
Venezuela.—Zootaxa 1553:59–68.
Benine, R. C., & G. A. M. Lopes. 2007. A new species of
Hemigrammus Gill, 1858 (Characiformes: Characidae) from
Rıo Caura, Venezuela.—Zootaxa 1610:53–59.
———, G. Z. Pelicao, & R. P. Vari. 2004. Tetragonopterus
lemniscatus (Characiformes: Characidae), a new species from
the Corantijn River basin in Suriname.—Proceedings of the
Biological Society of Washington 117:339–345.
Birindelli, J. L. O., M. H. Sabaj, & D. C. Taphorn. 2007. New species
of Rhynchodoras from the Rıo Orinoco, Venezuela, with
comments on the genus (Siluriformes: Doradidae).—Copeia
2007(3):672–684.
Buhrnheim, C. M., & L. R. Malabarba. 2007. Redescription of
Odontostilbe pulchra (Gill, 1858) (Teleostei: Characidae:
Cheirodontinae), and description of two new species from
the rıo Orinoco basin.—Neotropical Ichthyology 5(1):1–20.
Buitrago-Suarez, U. A., & B. M. Burr. 2007. Taxonomy of the catfish
genus Pseudoplatystoma Bleeker (Siluriformes: Pimelodidae)
with recognition of eight species.—Zootaxa 1512:1–38.
Chernoff, B., & A. Machado-Allison. 2005. Bryconops magoi and
Bryconops collettei (Characiformes: Characidae), two new
freshwater fish species from Venezuela, with comments on B.
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de Chambrier, S., & J. I. Montoya-Burgos. 2008. Pseudancistrus
corantijniensis, a new species from the Guyana Shield
Table 3.—Regions of the Guiana Shield used in checklist in
approximately west to east order.
CG Colombian Guayana
VA Venezuela—Amazonas
BO Venezuela—Bolıvar
DA Venezuela—Delta Amacuro
BA Brazil—Amazonas
RO Brazil—Roraima
PA Brazil—Para
AP Brazil—Amapa
GU Guyana
SU Suriname
FG French Guiana
(Siluriformes: Loricariidae) with a molecular and morpholog-
ical description of the Pseudancistrus barbatus group.—
Zootaxa 1918:45–58.
Deynat, P. 2006. Potamotrygon marinae n. sp., a new species of
freshwater stingrays from French Guiana (Myliobatiformes:
Potamotrygonidae).—Comptes Rendus Biologies 329(7):483–
493.
DoNascimiento, C., & J. G. Lundberg. 2005. Myoglanis aspredi-
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Rıo Ventuari, Venezuela.—Zootaxa 1009:37–49.
Friel, J. P. 2008. Pseudobunocephalus, a new genus of banjo catfish
with the description of a new species from the Orinoco River
system of Colombia and Venezuela (Siluriformes: Aspredini-
dae).—Neotropical Ichthyology 6(3):293–300.
Garutti, V. 2003. Revalidacao de Astyanax rupununi Fowler, 1914
(Teleostei, Characidae) e descricao de duas especies novas para
o genero.—Papeis Avulsos de Zoologia (Sao Paulo) 43(1):1–9.
Gibbs, A. K., & C. N. Barron. 1993. The geology of the Guiana
Shield. Oxford University Press, New York, 246 pp.
Hrbek, T., D. C. Taphorn, & 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.
Jegu, M., P. Keith, & P.-Y. Le Bail. 2003. Myloplus planquettei sp. n.
(Teleostei: Characidae) une nouvelle espece de grand Serra-
salminae phytophage du bouclier guyanais.—Revue Suisse de
Zoologie 1024:833–853.
Keith, P., L. Nandrin, & P.-Y. Le Bail. 2006. Rivulus gaucheri, a new
species of rivuline (Cyprinodontiformes: Rivulidae) from
French Guiana.—Cybium 30(2):133–137.
Kullander, S. O., & E. J. G. Ferreira. 2005. Two new species of
Apistogramma Regan (Teleostei: Cichlidae) from the rio
Trombetas, Para State, Brazil.—Neotropical Ichthyology
3(3):361–371.
———, & ———. 2006. A review of the South American cichlid
genus Cichla, with descriptions of nine new species (Teleostei:
Cichlidae).—Ichthyological Exploration of Freshwaters 17(4):
289–398.
Lasso, C. A., & F. Provenzano. 2002. Dos nuevas especies de bagres
del genero Trichomycterus (Siluriformes: Trichomycteridae)
de la Gran Sabana, Escudo de las Guayanas, Venezuela.—
Revista de Biologıa Tropical 50(3/4):1139–1149.
———, J. I. Mojica, J. S. Usma, J. A. Maldonaldo O., C.
DoNascimiento, D. C. Taphorn, F. Provenzano, O. M. Lasso
Alcala, G. Galvis, L. Vasquez, M. Lugo, A. Machado Allison,
R. Royero, C. Suarez, & A. Ortega Lara. 2004. Peces de las
cuenca del rıo Orinoco. Parte I: lista de especies y distribucion
por subcuencas.—Biota Colombiana 5:95–157.
Lasso-Alcala, O. M., D. C. Taphorn B., C. A. Lasso, & O. Leon-
Mata. 2006. Rivulus sape, a new species of killifish (Cyprino-
dontiformes: Rivulidae) from the Paragua River system,
Caronı River drainage, Guyana Shield, Venezuela.—Zootaxa
1275:21–29.
Lehman, A. P., & R. E. Reis. 2004. Callichthys serralabium: a new
species of neotropical catfish from the upper Orinoco and
Negro Rivers (Siluriformes: Callichthyidae).—Copeia 2004(2):
336–343.
Lopez-Fernandez, H., & D. C. Taphorn. 2004. Geophagus abalios, G.
dicrozoster and G. winemilleri (Perciformes: Cichlidae), three
new species from Venezuela.—Zootaxa 439:1–27.
———, D. C. Taphorn Baechle, & S. O. Kullander. 2006. Two new
species of Guianacara from the Guiana Shield of eastern
Venezuela (Perciformes: Cichlidae).—Copeia 2006(3):384–
395.
Lujan, N. K. 2008. Description of a new Lithoxus (Siluriformes:
Loricariidae) from the Guayana Highlands with a discussion
of Guiana Shield biogeography.—Neotropical Ichthyology
6(3):413–418.
———, M. Arce, & J. W. Armbruster. 2009. A new black
Baryancistrus with blue sheen from the upper Orinoco
(Siluriformes: Loricariidae).—Copeia 2009(1):50–56.
———, J. W. Armbruster, & M. H. Sabaj. 2007. Two new species of
Pseudancistrus from southern Venezuela (Siluriformes: Lor-
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163–174.
Lundberg, J. G., & W. M. Dahdul. 2008. Two new cis-Andean
species of the South American catfish genus Megalonema
allied to trans-Andean Megalonema xanthum, with description
of a new subgenus (Siluriformes: Pimelodidae).—Neotropical
Ichthyology 6(3):439–454.
Malabarba, M. C. S. L. 2004. Revision of the Neotropical genus
Triportheus Cope, 1872 (Characiformes: Characidae).—Neo-
tropical Ichthyology 2(4):167–204.
Montana, C. G., H. Lopez-Fernandez, & D. C. Taphorn. 2008. A
new species of Crenicichla (Perciformes: Cichlidae) from the
Ventuari River, Upper Orinoco River Basin, Amazonas State,
Venezuela.—Zootaxa 1856:33–40.
de Pinna, M., & P. Keith. 2003. A new species of the catfish genus
Ituglanis from French Guyana (Osteichthyes: Siluriformes:
Trichomycteridae).—Proceedings of the Biological Society of
Washington 116:873–882.
Provenzano, R. F., A. Machado-Allison, B. Chernoff, P. Willink, &
P. Petry. 2005. Harttia merevari, a new species of catfish
(Siluriformes: Loricariidae) from Venezuela.—Neotropical
Ichthyology 3(4):519–524.
Radda, A. C. 2004. Description of a new species of the rivuline genus
Rivulus Poey, 1869 (Rivulidae, Osteichthyes) from Rio Caura,
Bolivar State, Venezuela.—Annalen des Naturhistorischen
Museums in Wien, Serie B, 105B:21–25.
Ramos, R. T. C. 2003. Systematic review of Apionichthys (Pleuronecti-
formes: Achiridae), with description of four new species.—
Ichthyological Exploration of Freshwaters 14(2):97–126.
Reis, R. E., S. O. Kullander, & C. J. Ferraris, Jr. (eds.). 2003. Check
list of the freshwater fishes of South and Central America.
Edipucrs, Porto Alegre, Brazil, 729 pp.
Roman-Valencia, C., D. C. Taphorn B., & R. I. Ruiz-C. 2008. Two
new Bryconamericus: B. cinarucoense n. sp. and B. singularis n.
sp. (Characiformes, Characidae) from the Cinaruco River,
Orinoco Basin, with keys to all Venezuelan species.—Animal
Biodiversity and Conservation 31(1):15–27.
Romer, U., I. Hahn, & A. Conrad. 2006. Apistogramma wapisana
sp. n. Description of a dwarf cichlid from northern Brazil.
Cichlid Atlas 2. Mergus Verlag, Melle, Germany, pp. 748–
763.
Rosa, R. S., M. R. de Carvalho, & C. de Almeida Wanderley. 2008.
Potamotrygon boesemani (Chondrichthyes: Myliobatiformes:
Potamotrygonidae), a new species of Neotropical freshwater
stingray from Surinam.—Neotropical Ichthyology 6(1):1–8.
Sabaj, M. H. 2005. Taxonomic assessment of Leptodoras (Siluri-
formes: Doradidae) with descriptions of three new species.—
Neotropical Ichthyology 3(4):637–678.
———, D. C. Taphorn, & O. E. Castillo G. 2008. Two new species of
thicklip thornycats, genus Rhinodoras (Teleostei: Siluriformes:
Doradidae).—Copeia 2008(1):209–226.
Sabaj Perez, M. H., & J. L. O. Birindelli. 2008. Taxonomic revision of
extant Doras Lacepede, 1803 (Siluriformes: Doradidae) with
descriptions of three new species.—Proceedings of the
Academy of Natural Sciences of Philadelphia 157:189–233.
de Santana, C. D., & D. C. Taphorn. 2006. Sternarchorhynchus
gnomus, a new species of electric knifefish from the Lower Rıo
Caroni, Venezuela (Gymnotiformes: Apteronotidae).—Ich-
thyological Exploration of Freshwaters 17(1):1–8.
———, & R. P. Vari. 2009. The South American electric fish genus
Platyurosternarchus (Gymnotiformes: Apteronotidae).—Co-
peia 2009(2):233–244.
22 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Sarmento-Soares, L. M., & R. F. Martins-Pinheiro. 2008. A
systematic revision of Tatia (Siluriformes: Auchenipteridae:
Centromochlinae).—Neotropical Ichthyology 6(3):495–542.
Schaefer, S. A., & F. Provenzano, 2008. The Lithogeninae
(Siluriformes, Loricariidae): anatomy, interrelationships, and
description of a new species.—American Museum Novitates
3637:1–49.
———, ———, M. de Pinna, & J. N. Baskin. 2005. New and
noteworthy Venezuelan glanapterygine catfishes (Siluriformes,
Trichomycteridae), with discussion of their biogeography and
psammophily.—American Museum Novitates 3496:1–27.
Schindler, I., & W. Staeck. 2006. Geophagus gottwaldi sp. n. – a new
species of cichlid fish (Teleostei: Perciformes: Cichlidae) from
the drainage of the upper rıo Orinoco in Venezuela.—
Zoologische Abhandlungen (Dresden) 56:91–97.
———, & ———. 2008. Dicrossus gladicauda sp. n. – a new species of
crenicarine dwarf cichlids (Teleostei: Perciformes: Cichlidae)
from Colombia, South-America.—Vertebrate Zoology 58(1):
67–73.
Sidlauskas, B. L., & G. M. dos Santos. 2005. Pseudanos winterbot-
tomi: a new anostomine species (Teleostei: Characiformes:
Anostomidae) from Venezuela and Brazil, and comments on
its phylogenetic relationships.—Copeia 2005(1):109–123.
———, J. C. Garavello, & J. Jellen. 2007. A new Schizodon
(Characiformes: Anostomidae) from the Rıo Orinoco system,
with a redescription of S. Isognathus from the Rıo Paraguay
system.—Copeia 2007(3):711–725.
Staeck, W. 2003. Cichliden-Lexikon, Teil 3: Sudamerikanische
Zwergbuntbarsche. Dahne Verlag, Ettlingen, Germany, 219 pp.
———, & I. Schindler. 2007. Description of Laetacara fulvipinnis sp.
n. (Teleostei: Perciformes: Cichlidae) from the upper drain-
ages of the rio Orinoco and rio Negro in Venezuela.—
Vertebrate Zoology 57(1):63–71.
Suijker, W. H., & G. E. Collier. 2006. Rivulus mahdiaensis, a new
killifish from central Guyana (Cyprinodontiformes: Rivuli-
dae).—Zootaxa 1246:1–13.
Taphorn, B. D. C., H. Lopez-Fernandez, & C. R. Bernard. 2008.
Apareiodon agmatos, a new species from the upper Mazaruni
river, Guyana (Teleostei: Characiformes: Parodontidae).—
Zootaxa 1925:31–38.
Taphorn, B., C. G. Montana, & P. Buckup. 2006. Characidium
longum (Characiformes: Crenuchidae), a new fish from
Venezuela.—Zootaxa 1247:1–12.
Thomas, M. R., & L. H. Rapp Py-Daniel. 2008. Three new species of
the armored catfish genus Loricaria (Siluriformes: Loricar-
iidae) from river channels of the Amazon basin.—Neotropical
Ichthyology 6(3):379–394.
Vari, R. P., & C. J. Ferraris, Jr. 2006. The catfish genus Tetra-
nematichthys (auchenipteridae).—Copeia 2006(2):168–180.
———, ———, & M. C. C. de Pinna. 2005. The Neotropical whale
catfishes (Siluriformes: Cetopsidae: Cetopsinae), a revisionary
study.—Neotropical Ichthyology 3(2):127–238.
Vermeulen, F. B. M., & T. Hrbek. 2005 Kryptolebias sepia n. sp.
(Actinopterygii: Cyprinodontiformes: Rivulidae), a new killi-
fish from the Tapanahony River drainage in southeast
Surinam.—Zootaxa 928:1–20.
Werneke, D. C., J. W. Armbruster, N. K. Lujan, & D. C. Taphorn.
2005. Hemiancistrus guahiborum, a new suckermouth armored
catfish from Southern Venezuela (Siluriformes: Loricarii-
dae).—Neotropical Ichthyology 3(4):543–548.
———, M. H. Sabaj, N. K. Lujan, & J. W. Armbruster. 2005.
Baryancistrus demantoides and Hemiancistrus subviridis, two
new uniquely colored species of catfishes from Venezuela
(Siluriformes: Loricariidae).—Neotropical Ichthyology 3(4):
533–542.
Willink, P. W., B. Chernoff, A. Machado-Allison, F. Provenzano, &
P. Petry. 2003. Aphyocharax yekwanae, a new species of
bloodfin tetra (Teleostei: Characiformes: Characidae) from
the Guyana Shield of Venezuela.—Ichthyological Exploration
of Freshwaters 14(1):1–8.
Zanata, A. M., & M. Toledo-Piza. 2004. Taxonomic revision of the
South American fish genus Chalceus Cuvier (Teleostei:
Ostariophysi: Characiformes) with the description of three
new species.—Zoological Journal of the Linnean Society
140(1):103–135.
Zarske, A., P.-Y. Le Bail, & J. Gery. 2006. New and poorly known
Characiform fishes from French Guiana. 1. Two new Tetras
of the genera Hemigrammus and Hyphessobrycon (Teleostei:
Characiformes: Characidae).—Zoologische Abhandlungen
(Dresden) 55:17–30.
———, J. Gery, & I. Isbrucker. 2004. Moenkhausia rara sp. n. – eine
neue, bereits bestandsgefahrdete Salmler-Art (Teleostei: Char-
aciformes: Characidae) aus Surinam und Franzosisch Guayana
mit einer erganzenden Beschreibung von M. simulata (Eigen-
mann in Pearson, 1924).—Zoologische Abhandlungen (Dres-
den) 54:19–30.
NUMBER 17 23
CHECKLIST OF THE FISHES OF THE GUIANA SHIELD
Order: Pristiformes
Family: Pristidae—John D. McEachran & M. R. de Carvalho
Pristis pectinata Latham, 1794 DA GU?
Pristis pristis (Linnaeus, 1758) DA
Order: Myliobatiformes
Family: Potamotrygonidae—Marcelo R. de Carvalho & Ricardo
S. Rosa
Paratrygon aiereba (Muller & Henle, 1841) BO DA RO
Potamotrygon boesemani Rosa, Carvalho & Almeida, 2008 GU SU
Potamotrygon marinae Deynat, 2006 FG
Potamotrygon motoro (Muller & Henle, 1841) CG VA BO RO
Potamotrygon orbignyi (Castelnau, 1855) VA BO DA GU SU FG
Potamotrygon schroederi Fernandez-Yepez, 1957 VA BO DA
Potamotrygon scobina Garman, 1913 RO PA
Order: Osteoglossiformes
Family: Osteoglossidae—Carl J. Ferraris, Jr.
Osteoglossum bicirrhosum (Cuvier, 1829) CG RO AP GU FG
Family: Arapaimidae—Carl J. Ferraris, Jr.
Arapaima gigas (Schinz, 1822) GU
Order: Anguilliformes
Family: Ophichthidae—Sven O. Kullander
Stictorhinus potamius (Bohlke & McCosker, 1975) BO DA
Order: Clupeiformes
Family: Clupeidae—Carl J. Ferraris, Jr.
Rhinosardinia bahiensis (Steindachner, 1879) BO DA AP GU SU FG
Family: Engraulidae—Carl J. Ferraris, Jr. & Sven O. Kullander
Anchoa spinifer (Valenciennes, 1848) DA GU SU FG
Anchovia surinamensis (Bleeker, 1866) BO DA GU SU FG
Anchoviella brevirostris (Gunther, 1868) BO DA GU SU FG
Anchoviella cayennensis (Puyo, 1946) SU FG
Anchoviella guianensis (Eigenmann, 1912) BO DA GU SU FG
Anchoviella jamesi (Jordan & Seale, 1926) VA BO BA
Anchoviella lepidentostole (Fowler 1911) GU SU FG
Lycengraulis batesii (Gunther, 1868) BO DA GU SU FG
Lycengraulis grossidens (Spix & Agassiz, 1829) DA SU FG
Pterengraulis atherinoides (Linnaeus, 1766) BO DA SU FG
Family: Pristigasteridae—Fabio di Dario
Pellona castelnaeana (Valenciennes, 1847) CG VA BO DA
Pellona flavipinnis (Valenciennes, 1836) CG VA BO DA GU SU FG
Order: Characiformes
Family: Parodontidae—Carla S. Pavanelli
Apareiodon agmatos Taphorn, Lopez-Fernandez & Bernard,
2008
GU
Apareiodon gransabana Starnes & Schindler, 1993 BO GU SU FG
Apareiodon orinocensis Bonilla et al., 1999 VA BO
Parodon apolinari Myers, 1930 CG VA BO
Parodon bifasciatus Eigenmann, 1912 RO GU
Parodon guyanensis Gery, 1959 VA BO GU SU FG
Parodon suborbitalis Valenciennes, 1850 BO
Family: Curimatidae—Richard P. Vari
Curimata cerasina Vari, 1984 VA BO DA
Curimata cisandina (Allen, 1942) BA
Curimata cyprinoides (Linnaeus, 1766) DA GU SU FG
Curimata incompta Vari, 1984 VA BO DA
Curimata ocellata (Eigenmann & Eigenmann, 1889) CG VA BO BA
Curimata roseni Vari, 1989 VA BO DA GU
Curimata vittata (Kner, 1858) CG VA BO GU
Curimatella alburna (Muller & Troschel, 1844) RO GU
Curimatella dorsalis (Eigenmann & Eigenmann, 1889) CG VA BO DA
Curimatella immaculata (Fernandez-Yepez, 1948) CG VA BO DA RO GU
Curimatopsis crypticus Vari, 1982 AP GU SU FG
Curimatopsis evelynae Gery, 1964 CG VA BA?
Curimatopsis macrolepis (Steindachner, 1876) CG VA BO DA BA
Cyphocharax abramoides (Kner, 1859) CG VA BO PA
Cyphocharax festivus Vari, 1992 VA BO GU
Cyphocharax helleri (Steindachner, 1910) BO AP GU SU FG
Cyphocharax leucostictus (Eigenmann & Eigenmann, 1889) VA RO
Cyphocharax meniscaprorus Vari, 1992 VA BO
Cyphocharax mestomyllon Vari, 1992 BA
Cyphocharax microcephalus (Eigenmann & Eigenmann, 1889) GU SU FG?
Cyphocharax multilineatus (Myers, 1927) VA BO BA
Cyphocharax notatus (Steindachner, 1908) VA
Cyphocharax oenas Vari, 1992 VA BO DA
Cyphocharax punctatus (Vari & Nijssen, 1986) SU FG
Cyphocharax spilurus (Gunther, 1864) CG VA BO BA RO GU SU FG
Potamorhina altamazonica (Cope, 1878) CG VA BO DA
Psectrogaster ciliata (Muller & Troschel, 1844) CG VA BO DA RO GU
Psectrogaster essequibensis (Gunther, 1864) GU
Steindachnerina argentea (Gill, 1858) CG VA BO DA
Steindachnerina bimaculata (Steindachner, 1876) BO DA
Steindachnerina guentheri (Eigenmann & Eigenmann, 1889) BO DA GU
Steindachnerina planiventris Vari & Vari, 1989 RO
Steindachnerina pupula Vari, 1991 BO
Steindachnerina varii Gery, Planquette & Le Bail, 1991 AP SU FG
Family: Prochilodontidae—Richard P. Vari
Prochilodus mariae Eigenmann, 1922 CG VA BO DA
Prochilodus rubrotaeniatus Jardine, 1841 VA BO RO GU SU FG
Semaprochilodus insignis (Jardine, 1841) CG GU
Semaprochilodus kneri (Pellegrin, 1909) VA BO DA
Semaprochilodus laticeps (Steindachner, 1879) CG VA BO DA
Semaprochilodus varii Castro, 1988 SU FG
Family: Anostomidae—Richard P. Vari
Abramites hypselonotus (Gunther, 1868) VA BO DA GU
26 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Anostomoides atrianalis Pellegrin, 1908 VA BO
Anostomoides laticeps (Eigenmann, 1912) VA BO GU
Anostomus anostomus (Linnaeus, 1758) GU
Anostomus brevior Gery, 1961 FG
Anostomus plicatus Eigenmann, 1912 GU SU
Anostomus spiloclistron Winterbottom, 1974 SU
Anostomus ternetzi Fernandez-Yepez, 1949 VA BO DA GU SU FG
Gnathodolus bidens Myers, 1927 VA BO
Laemolyta fernandezi Myers, 1950 VA BO DA
Laemolyta orinocensis (Steindachner, 1879) VA BO DA
Laemolyta proximate (Garman, 1890) BA GU
Laemolyta taeniata (Kner, 1859) CG VA BO
Leporinus acutidens (Valenciennes, 1836) FG
Leporinus affinis Gunther, 1864 BO
Leporinus agassizi Steindachner, 1876 VA BO
Leporinus alternus Eigenmann, 1912 DA GU
Leporinus arcus Eigenmann, 1912 VA BO GU SU FG
Leporinus badueli Puyo, 1948 FG
Leporinus brunneus Myers, 1950 VA BO BA
Leporinus desmotes Fowler, 1914 GU SU
Leporinus despaxi Puyo, 1943 RO FG
Leporinus fasciatus (Bloch, 1794) SU FG
Leporinus friderici (Bloch, 1794) GU SU FG
Leporinus gossei Gery, Planquette & Le Bail, 1991 SU FG
Leporinus granti Eigenmann, 1912 GU FG
Leporinus latofasciatus Steindachner, 1910 VA BO DA
Leporinus lebaili Gery & Planquette, 1983 SU FG
Leporinus leschenaulti Valenciennes, 1850 FG
Leporinus maculatus Muller & Troschel, 1844 GU SU FG
Leporinus megalepis Gunther, 1863 GU
Leporinus melanostictus Norman, 1926 AP FG
Leporinus nigrotaeniatus (Jardine, 1841) GU
Leporinus nijsseni Garavello, 1990 SU FG
Leporinus ortomaculatus Garavello, 2000 VA BO
Leporinus paralternus Fowler, 1914 GU
Leporinus pellegrinii Steindachner, 1910 GU SU FG
Leporinus pitingai Santos & Jegu, 1996 BA
Leporinus punctatus Garavello, 2000 VA BO
Leporinus spilopleura Norman, 1926 AP FG
Leporinus steyermarki Inger, 1956 VA BO DA
Leporinus uatumaensis Santos & Jegu, 1996 BA
Leporinus yophorus Eigenmann, 1922 VA BO
Pseudanos gracilis (Kner, 1858) VA BO BA
Pseudanos irinae Winterbottom, 1980 VA BO
Pseudanos trimaculatus (Kner, 1858) GU
Pseudanos winterbottomi Sidlauskas & Santos, 2005 VA BO
Sartor elongatus Santos & Jegu, 1987 PA
Schizodon fasciatus Spix & Agassiz, 1829 BO FG
NUMBER 17 27
Schizodon scotorhabdotus Sidlauskas, Garavello & Jellen, 2007 VA BO DA
Synaptolaemus cingulatus Myers & Fernandez-Yepez, 1950 VA BO
Family: Chilodontidae—Richard P. Vari
Caenotropus labyrinthicus (Kner, 1858) CG VA BO DA GU SU
Caenotropus maculosus (Eigenmann, 1912) BO GU SU FG
Caenotropus mestomorgmatos Vari, Castro & Raredon, 1995 VA BO BA
Chilodus punctatus Muller & Troschel, 1844 CG VA BO DA GU SU
Chilodus zunevei Puyo, 1946 FG
Family: Crenuchidae—Paulo A. Buckup
Ammocryptocharax elegans Weitzman & Kanazawa, 1976 CG VA BO
Ammocryptocharax lateralis (Eigenmann, 1909) GU
Ammocryptocharax minutus Buckup, 1993 CG VA BA
Ammocryptocharax vintonae (Eigenmann, 1909) BO GU
Characidium boaevistae Steindachner, 1915 VA BO RO
Characidium chupa Schultz, 1944 BO
Characidium declivirostre Steindachner, 1915 VA BO BA
Characidium hasemani Steindachner, 1915 BO RO
Characidium longum Taphorn, Montana & Buckup, 2006 VA BO BA
Characidium pellucidum Eigenmann, 1909 GU SU FG
Characidium pteroides Eigenmann, 1909 GU
Characidium steindachneri Cope, 1878 CG VA BO BA RO PA GU
Characidium zebra Eigenmann, 1909 CG VA BO DA BA RO PA AP GU SU FG
Crenuchus spilurus Gunther, 1863 CG VA BO GU SU FG
Elachocharax geryi Weitzman & Kanazawa, 1978 CG VA BO BA
Elachocharax mitopterus Weitzman, 1986 VA BA
Elachocharax pulcher Myers, 1927 CG VA BO BA PA
Leptocharacidium omospilus Buckup, 1993 VA BO BA RO
Melanocharacidium blennioides (Eigenmann, 1909) BO GU
Melanocharacidium compressus Buckup, 1993 VA BO
Melanocharacidium depressum Buckup, 1993 VA BA
Melanocharacidium dispilomma Buckup, 1993 CG VA BO BA RO PA FG
Melanocharacidium melanopteron Buckup, 1993 VA BO
Melanocharacidium nigrum Buckup, 1993 BA RO
Melanocharacidium pectorale Buckup, 1993 VA BO BA RO PA
Microcharacidium eleotrioides (Gery, 1960) SU FG
Microcharacidium gnomus Buckup, 1993 CG VA BO BA
Microcharacidium weitzmani Buckup, 1993 CG VA BO BA PA
Odontocharacidium aphanes (Weitzman & Kanazawa, 1977) CG VA BA
Poecilocharax bovalii Eigenmann, 1909 GU
Poecilocharax weitzmani Gery, 1965 CG VA BO
Skiotocharax meizon Presswell, Weitzman & Bergquist, 2000 GU
Family: Hemiodontidae—Francisco Langeani Neto
Anodus elongatus Agassiz, 1829 PA
Anodus orinocensis (Steindachner, 1887) CG VA BO DA
Argonectes longiceps (Kner, 1858) PA SU FG
Bivibranchia bimaculata Vari, 1985 GU SU FG
Bivibranchia fowleri (Steindachner, 1908) CG VA BO DA BA RO GU
Bivibranchia simulata Gery, Planquette & Le Bail, 1991 AP FG
28 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Bivibranchia velox (Eigenmann & Myers, 1927) AP
Hemiodus amazonum (Humboldt, 1821) GU
Hemiodus argenteus Pellegrin, 1908 CG VA BO DA GU SU
Hemiodus atranalis (Fowler, 1940) RO PA GU
Hemiodus goeldii Steindachner, 1908 AP
Hemiodus gracilis Gunther, 1864 VA BO
Hemiodus huraulti (Gery, 1964) SU FG
Hemiodus immaculatus Kner, 1858 CG VA BO DA PA
Hemiodus microlepis Kner, 1858 VA BO
Hemiodus quadrimaculatus Pellegrin, 1908 RO AP GU SU FG
Hemiodus semitaeniatus Kner, 1858 CG VA BO GU
Hemiodus thayeria Bohlke, 1955 CG VA BO BA
Hemiodus unimaculatus (Bloch, 1794) AP GU SU FG
Hemiodus vorderwinkleri (Gery, 1964) BO GU
Family: Gasteropelecidae—Stanley H. Weitzman
Carnegiella marthae Myers, 1927 CG VA BO DA BA
Carnegiella strigata (Gunther, 1864) CG VA BO DA GU SU
Gasteropelecus sternicla (Linnaeus, 1758) BO DA GU FG
Thoracocharax stellatus (Kner, 1858) CG VA BO DA
Family: Characidae
Genera Incerta Sedis—Flavio C. T. Lima
Aphyocharacidium melandetum (Eigenmann, 1912) GU
Aphyodite grammica Eigenmann, 1912 GU
Astyanax bimaculatus (Linnaeus, 1758) CG VA BO DA GU SU FG
Astyanax clavitaeniatus Garutti, 2003 RO
Astyanax fasciatus (Cuvier, 1819) BO
Astyanax guianensis Eigenmann, 1909 VA BO GU
Astyanax leopoldi Gery, Planquette & Le Bail, 1988 AP FG
Astyanax metae Eigenmann, 1914 VA BO
Astyanax mutator Eigenmann, 1909 GU
Astyanax myersi (Fernandez-Yepez, 1950) VA BO
Astyanax rupununi Garutti, 2003 GU
Astyanax scintillans Myers, 1928 VA BO
Astyanax siapae Garutti, 2003 VA
Astyanax validus Gery, Planquette & Le Bail, 1991 FG
Astyanax venezuelae Schultz, 1944 VA BO
Aulixidens eugeniae Bohlke, 1952 VA BO
Bryconamericus alpha Eigenmann, 1914 CG VA BO
Bryconamericus beta Eigenmann, 1914 VA BO
Bryconamericus cinarucoense Roman-Valencia, Taphorn & Ruiz-
C., 2008
GU
Bryconamericus cismontanus Eigenmann, 1914 VA BO
Bryconamericus cristiani Roman-Valencia, 1998 CG
Bryconamericus deuterodonoides Eigenmann, 1914 VA BO
Bryconamericus hyphesson Eigenmann, 1909 GU
Bryconamericus macropthalmus Roman-Valencia, 2003 VA
Bryconamericus orinocoense Roman-Valencia, 2003 VA
Bryconamericus subtilisform Roman-Valencia, 2003 BO
Bryconexodon trombetasi Jegu, Santos & Ferreira, 1991 PA
NUMBER 17 29
Bryconops affinis (Gunther, 1864) GU SU FG
Bryconops alburnoides Kner, 1858 CG VA BO
Bryconops caudomaculatus (Gunther, 1864) VA BO DA GU SU FG
Bryconops colanegra Chernoff & Machado-Allison, 1999 VA BO
Bryconops colaroja Chernoff & Machado-Allison, 1999 BO
Bryconops collettei Chernoff & Machado-Allison, 2005 VA BO
Bryconops cyrtogaster (Norman, 1926) AP FG
Bryconops disruptus Machado-Allison & Chernoff, 1997 VA BA
Bryconops giacopinii (Fernandez-Yepez, 1950) VA BO
Bryconops humeralis Machado-Allison, Chernoff & Buckup,
1996
VA BO
Bryconops imitator Chernoff & Machado-Allison, 2002 VA BO
Bryconops inpai Knoppel, Junk & Gery, 1968 VA BA
Bryconops melanurus (Bloch, 1794) GU SU FG
Bryconops vibex Machado-Allison, Chernoff & Buckup, 1996 VA BO
Ceratobranchia joanae Chernoff & Machado-Allison, 1990 VA BO
Chalceus epakros Zanata & Toleda-Piza, 2004 VA BO RO PA GU
Chalceus macrolepidotus Cuvier, 1816 CG VA BO DA GU SU FG
Creagrutus bolivari Schultz, 1944 CG VA BO DA
Creagrutus ephippiatus Vari & Harold, 2001 VA BO DA
Creagrutus gyrospilus Vari & Harold, 2001 VA BO
Creagrutus machadoi Vari & Harold, 2001 BO
Creagrutus magoi Vari & Harold, 2001 VA BO
Creagrutus maxillaris (Myers, 1927) CG VA BO BA
Creagrutus melanzonus Eigenmann, 1909 BO GU FG
Creagrutus melasma Vari, Harold & Taphorn, 1994 VA BO
Creagrutus menezesi Vari & Harold, 2001 RO
Creagrutus phasma Myers, 1927 CG VA BO BA
Creagrutus planquettei Gery & Renno, 1989 FG
Creagrutus provenzanoi Vari & Harold, 2001 VA BO
Creagrutus runa Vari & Harold, 2001 CG VA BO BA
Creagrutus veruina Vari & Harold, 2001 VA
Creagrutus vexillapinnus Vari & Harold, 2001 CG VA BA?
Creagrutus xiphos Vari & Harold, 2001 BO
Creagrutus zephyrus Vari & Harold, 2001 CG VA BA?
Ctenobrycon spilurus (Valenciennes, 1850) VA BO DA GU SU FG
Deuterodon potaroensis Eigenmann, 1909 GU
Exodon paradoxus Muller & Troschel, 1844 GU
Gymnocorymbus thayeri Eigenmann, 1908 CG VA BO DA GU
Gymnotichthys hildae Fernandez-Yepez, 1950 VA BO
Hemibrycon surinamensis Gery, 1962 SU FG
Hemigrammus aereus Gery, 1959 FG
Hemigrammus analis Durbin, 1909 CG VA BO BA GU
Hemigrammus barrigonae Eigenmann & Henn, 1914 CG VA BO
Hemigrammus belottii (Steindachner, 1882) FG
Hemigrammus boesemani Gery, 1959 SU FG
Hemigrammus cylindricus Durbin, 1909 GU
Hemigrammus erythrozonus Durbin, 1909 GU
Hemigrammus guyanensis Gery, 1959 FG
30 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Hemigrammus iota Durbin, 1909 GU
Hemigrammus lunatus Durbin, 1918 SU
Hemigrammus marginatus Ellis, 1911 VA BO DA
Hemigrammus micropterus Meek, 1907 VA BO DA
Hemigrammus microstomus Durbin, 1918 VA BO
Hemigrammus mimus Bohlke, 1955 VA BO
Hemigrammus newboldi (Fernandez-Yepez, 1949) VA BO DA
Hemigrammus ocellifer (Steindachner, 1882) GU SU FG
Hemigrammus ora Zarske, Le Bail & Gery, 2006 FG
Hemigrammus orthus Durbin, 1909 GU
Hemigrammus rhodostomus Ahl, 1924 CG VA BO
Hemigrammus rodwayi Durbin, 1909 GU SU FG
Hemigrammus schmardae (Steindachner, 1882) CG VA BO BA
Hemigrammus stictus (Durbin, 1909) CG VA BO GU
Hemigrammus taphorni Benine & Lopes, 2007 BO
Hemigrammus unilineatus (Gill, 1858) GU SU FG
Hyphessobrycon albolineatum Fernandez-Yepez, 1950 VA BO
Hyphessobrycon borealis Zarske, Le Bail & Gery, 2006 AP SU FG
Hyphessobrycon catableptus (Durbin, 1909) GU
Hyphessobrycon copelandi Durbin, 1908 FG
Hyphessobrycon diancistrus Weitzman, 1977 CG VA BO
Hyphessobrycon eos Durbin, 1909 GU
Hyphessobrycon epicharis Weitzman & Palmer, 1997 CG VA BA
Hyphessobrycon eques (Steindachner, 1882) FG
Hyphessobrycon georgettae Gery, 1961 SU
Hyphessobrycon hildae Fernandez-Yepez, 1950 VA BO
Hyphessobrycon metae Eigenmann & Henn, 1914 CG VA BO
Hyphessobrycon minimus Durbin, 1909 GU
Hyphessobrycon minor Durbin, 1909 GU
Hyphessobrycon rosaceus Durbin, 1909 GU SU
Hyphessobrycon roseus (Gery, 1960) FG
Hyphessobrycon simulatus (Gery, 1960) FG
Hyphessobrycon sweglesi (Gery, 1961) CG VA BO
Hyphessobrycon takasei Gery, 1964 AP FG
Jupiaba abramoides (Eigenmann, 1909) CG VA BO GU SU FG
Jupiaba atypindi Zanata, 1997 RO
Jupiaba essequibensis (Eigenmann, 1909) GU
Jupiaba keithi (Gery, Planquette & Le Bail, 1996) FG
Jupiaba maroniensis (Gery, Planquette & Le Bail, 1996) FG
Jupiaba meunieri (Gery, Planquette & Le Bail, 1996) SU FG
Jupiaba mucronata (Eigenmann, 1909) GU
Jupiaba ocellata (Gery, Planquette & Le Bail, 1996) AP? FG
Jupiaba pinnata (Eigenmann, 1909) GU SU
Jupiaba polylepis (Gunther, 1864) PA GU SU
Jupiaba potaroensis (Eigenmann, 1909) GU
Jupiaba scologaster (Weitzman & Vari, 1986) CG VA BO BA
Knodus heteresthes (Eigenmann, 1908) VA BO
Microschemobrycon callops Bohlke, 1953 VA BO PA
NUMBER 17 31
Microschemobrycon casiquiare Bohlke, 1953 VA BA
Microschemobrycon melanotus (Eigenmann, 1912) GU
Microschemobrycon meyburgi Meinken, 1975 RO
Moenkhausia browni Eigenmann, 1909 GU
Moenkhausia chrysargyrea (Gunther, 1864) VA BO GU FG
Moenkhausia collettii (Steindachner, 1882) CG VA BO DA GU SU FG
Moenkhausia copei (Steindachner, 1882) CG VA BO
Moenkhausia cotinho Eigenmann, 1908 CG VA BO
Moenkhausia dichroura (Kner, 1858) VA BO DA
Moenkhausia georgiae Gery, 1965 SU FG
Moenkhausia grandisquamis (Muller & Troschel, 1845) VA BO DA GU SU FG
Moenkhausia hemigrammoides Gery, 1965 SU FG
Moenkhausia inrai Gery, 1992 FG
Moenkhausia intermedia Eigenmann, 1908 CG VA BO FG
Moenkhausia lata Eigenmann, 1908 AP FG
Moenkhausia lepidura (Kner, 1858) VA BO DA GU SU
Moenkhausia megalops (Eigenmann, 1907) FG
Moenkhausia miangi Steindachner, 1915 BO RO
Moenkhausia moisae Gery, Planquette & Le Bail, 1995 FG
Moenkhausia oligolepis (Gunther, 1864) CG VA BO DA GU SU FG
Moenkhausia rara Zarske, Gery & Isbrucker, 2004 PA SU FG
Moenkhausia shideleri Eigenmann, 1909 GU
Moenkhausia surinamensis Gery, 1965 AP SU FG
Paracheirodon axelrodi (Schultz, 1956) CG VA BO BA
Paracheirodon simulans (Gery, 1963) CG VA BO BA
Paragoniates alburnus Steindachner, 1876 VA BO
Parapristella aubynei (Eigenmann, 1909) GU
Pristella maxillaris (Ulrey, 1894) VA BO GU FG
Salminus hilarii Valenciennes, 1850 CG VA BO RO
Scissor macrocephalus Gunther, 1864 SU
Serrabrycon magoi Vari, 1986 CG VA BO BA
Thayeria ifati Gery, 1959 FG
Triportheus angulatus (Spix & Agassiz, 1829) GU?
Triportheus auritus Malabarba, 2004 VA BO DA GU?
Triportheus brachipomus Malabarba, 2004 VA AP? GU SU FG
Triportheus elongatus (Gunther, 1864) VA BO
Triportheus venezuelensis Malabarba, 2004 BO
Xenagoniates bondi Myers, 1942 CG VA BO DA
Subfamily: Agoniatinae—Angela M. Zanata
Agoniates anchovia Eigenmann, 1914 PA
Agoniates halecinus Muller & Troschel, 1845 VA BO RO PA AP? GU
Subfamily: Iguanodectinae—Cristiano R. Moreira
Iguanodectes adujai Gery, 1970 VA BA
Iguanodectes geisleri Gery, 1970 VA BA
Iguanodectes spilurus (Gunther, 1864) CG VA BO DA BA RO AP GU
Piabucus dentatus (Koelreuter, 1763) VA DA GU SU FG
Subfamily: Bryconinae—Flavio C. T. Lima
Brycon amazonicus (Spix & Agassiz, 1829) CG VA BO GU
32 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Brycon bicolor Pellegrin, 1909 VA BO DA
Brycon coquenani Steindachner, 1915 VA BO
Brycon falcatus Muller & Troschel, 1844 CG VA BO RO PA GU SU FG
Brycon pesu Muller & Troschel, 1845 CG VA BO DA RO PA AP GU SU FG
Subfamily: Serrasalminae—Michel Jegu
Acnodon oligacanthus (Muller & Trochel, 1844) GU SU FG
Acnodon senai Jegu & Santos, 1990 PA AP
Catoprion mento (Cuvier, 1819) CG VA BO DA BA RO PA AP GU
Colossoma macropomum (Cuvier, 1818) CG VA BO DA
Metynnis altidorsalis Ahl, 1923 GU SU
Metynnis argenteus Ahl, 1924 VA BO DA GU
Metynnis hypsauchen (Muller & Troschel, 1844) VA BO RO GU
Metynnis lippincottianus (Cope, 1870) CG VA AP? FG
Metynnis luna (Cope, 1878) CG VA BO GU
Metynnis orinocensis (Steindachner, 1908) DA
Mylesinus paraschomburgkii Jegu, Santos & Ferreira, 1989 BA PA AP
Mylesinus schomburgkii Valenciennes, 1850 GU
Myleus knerii (Steindachner, 1881) FG
Myleus setiger Muller & Troschel, 1844 CG VA BO BA RO PA GU SU
Myloplus asterias (Muller & Troschel, 1844) VA BO PA AP GU
Myloplus lobatus (Valenciennes, 1850) PA
Myloplus planquettei Jegu, Keith & Le Bail, 2003 GU SU FG
Myloplus rhomboidalis (Cuvier, 1818) VA RO PA AP GU SU FG
Myloplus rubripinnis (Muller & Troschel, 1844) CG VA BO RO AP GU SU FG
Myloplus schomburgkii (Jardine & Schomburgk, 1841) CG VA BO BA PA
Myloplus ternetzi (Norman, 1929) AP SU FG
Myloplus torquatus (Kner, 1858) VA BO BA RO GU
Mylossoma aureum (Agassiz, 1829) BO DA
Mylossoma duriventre (Cuvier, 1818) VA BO DA AP
Piaractus brachypomus (Cuvier, 1818) VA BO DA GU
Pristobrycon aureus (Spix & Agassiz, 1829) GU
Pristobrycon calmoni (Steindachner, 1908) BO DA BA AP GU
Pristobrycon careospinus Fink & Machado-Allison, 1992 CG VA
Pristobrycon eigenmanni (Norman, 1929) VA BO DA BA RO PA AP GU SU FG
Pristobrycon maculipinnis Fink & Machado-Allison, 1992 VA
Pristobrycon striolatus (Steindachner, 1908) VA BO DA BA PA AP GU SU FG
Pygocentrus cariba (Humboldt & Valenciennes, 1821) CG VA BO DA
Pygocentrus nattereri Kner, 1858 AP GU
Pygopristis denticulata (Cuvier, 1819) VA BO BA PA AP GU SU FG
Serrasalmus altispinis Merckx, Jegu & Santos, 2000 BA PA
Serrasalmus altuvei Ramırez, 1965 VA BO DA
Serrasalmus elongatus Kner, 1858 BO DA BA PA
Serrasalmus gouldingi Fink & Machado-Allison, 1992 VA BO BA PA
Serrasalmus hastatus Fink & Machado-Allison, 2001 BA RO
Serrasalmus irritans Peters, 1877 VA BO DA
Serrasalmus maculatus Kner, 1858 AP
Serrasalmus manueli (Fernandez-Yepez & Ramırez, 1967) CG VA BA AP
Serrasalmus medinai Ramırez, 1965 BO DA
NUMBER 17 33
Serrasalmus nalseni Fernandez-Yepez, 1969 VA BO
Serrasalmus rhombeus (Linnaeus, 1766) CG VA BO DA BA RO PA AP GU SU FG
Serrasalmus serrulatus (Valenciennes, 1850) GU?
Tometes lebaili Jegu, Keith & Belmont-Jegu, 2002 SU FG
Tometes makue Jegu, Santos & Belmont-Jegu, 2002 CG VA BO BA
Tometes trilobatus Valenciennes, 1850 AP FG
Subfamily: Aphyocharacinae—Rosana S. Lima
Aphyocharax avary Souza-Lima, 2003 VA BO GU
Aphyocharax colifax Taphorn & Thomerson, 1991 VA BO
Aphyocharax erythrurus Eigenmann, 1912 GU
Aphyocharax yekwanae Willink, Chernoff, Machado-Allison,
Provenzano & Petry, 2003
BO
Subfamily: Characinae—Naercio A. Menezes & Carlos A.S.
Lucena
Acanthocharax microlepis Eigenmann, 1912 GU
Acestrocephalus sardina (Fowler, 1913) VA BO BA GU
Charax apurensis Lucena, 1987 VA BO DA
Charax gibbosus (Linnaeus, 1758) GU SU
Charax hemigrammus (Eigenmann, 1912) GU
Charax metae Eigenmann, 1922 BO
Charax michaeli Lucena, 1989 RO
Charax notulatus Lucena, 1987 VA BO DA
Charax rupununi Eigenmann, 1912 PA GU
Cynopotamus bipunctatus Pellegrin, 1909 VA BO
Cynopotamus essequibensis Eigenmann, 1912 GU SU FG
Gnathocharax steindachneri Fowler, 1913 CG VA BO GU
Heterocharax leptogrammus Toledo-Piza, 2000 VA BA
Heterocharax macrolepis Eigenmann, 1912 CG VA BO GU
Heterocharax virgulatus Toledo-Piza, 2000 VA BA
Lonchogenys ilisha Myers, 1927 CG VA BO BA
Phenacogaster apletostigma Lucena & Gama, 2007 AP
Phenacogaster carteri (Norman, 1934) GU
Phenacogaster megalostictus Eigenmann, 1909 GU
Phenacogaster microstictus Eigenmann, 1909 GU SU
Priocharax ariel Weitzman & Vari, 1987 VA BO
Roeboides affinis (Gunther, 1868) VA BO
Roeboides araguaito Lucena, 2003 CG
Roeboides dientonito Schultz, 1944 VA BO DA GU
Roeboides myersii Gill, 1870 VA BO DA
Roeboides numerosus Lucena, 2000 VA BO
Roeboides oligistos Lucena, 2000 RO PA
Roeboides thurni Eigenmann, 1912 GU SU FG
Subfamily: Stethaprioninae—Roberto E. Reis
Brachychalcinus orbicularis (Valenciennes, 1850) GU SU
Poptella brevispina Reis, 1989 RO PA GU SU
Poptella compressa (Gunther, 1864) CG VA BO DA GU
Poptella longipinnis (Popta, 1901) CG VA BO DA SU
Subfamily: Tetragonopterinae—Roberto E. Reis
Tetragonopterus chalceus Spix & Agassiz, 1829 CG VA BO DA GU SU FG
34 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Tetragonopterus lemniscatus Benine, Pelicao & Vari, 2004 SU
Subfamily: Cheirodontinae—Luiz R. Malabarba
Cheirodontops geayi Schultz, 1944 VA BO DA
Odontostilbe gracilis (Gery, 1960) FG
Odontostilbe littoris (Gery, 1960) FG
Odontostilbe pulchra (Gill, 1858) VA BO DA RO
Odontostilbe splendida Buhrnheim & Malabarba, 2007 BO
Subfamily: Glandulocaudinae—Stanley H. Weitzman
Ptychocharax rhyacophila Weitzman, Fink, Machado-Allison &
Royero, 1994
VA
Family: Acestrorhynchidae—Naercio A. Menezes
Acestrorhynchus falcatus (Bloch, 1794) VA GU SU FG
Acestrorhynchus falcirostris (Cuvier, 1819) CG VA BA GU
Acestrorhynchus grandoculis Menezes & Gery, 1983 VA BO BA
Acestrorhynchus heterolepis (Cope, 1878) VA
Acestrorhynchus microlepis (Schomburgk, 1841) CG VA BO DA BA GU SU FG
Acestrorhynchus minimus Menezes, 1969 VA BO PA
Acestrorhynchus nasutus Eigenmann, 1912 VA GU
Family: Cynodontidae—Monica Toledo-Piza
Cynodon gibbus Spix & Agassiz, 1829 CG BO DA GU
Cynodon meionactis Gery, Le Bail & Keith, 1999 SU FG
Cynodon septenarius Toledo-Piza, 2000 VA BO GU
Hydrolycus armatus (Jardine, 1841) CG VA BO DA GU
Hydrolycus tatauaia Toledo-Piza, Menezes & Santos, 1999 VA BO GU
Hydrolycus wallacei Toledo-Piza, Menezes & Santos, 1999 VA BO
Rhaphiodon vulpinus Spix & Agassiz, 1829 VA BO DA GU
Roestes ogilviei (Fowler, 1914) RO GU
Family: Erythrinidae—Based on Cloffsca account (Osvaldo O.
Oyakawa)
Erythrinus erythrinus (Bloch & Schneider, 1801) CG VA BO DA GU SU FG
Hoplerythrinus gronovii (Valenciennes, 1847) FG
Hoplerythrinus unitaeniatus (Agassiz, 1829) CG VA BO DA AP GU SU FG
Hoplias aimara (Valenciennes, 1847) VA BO FG
Hoplias macrophthalmus (Pellegrin, 1907) CG VA BO DA GU SU FG
Hoplias malabaricus (Bloch, 1794) CG VA BO DA GU SU FG
Hoplias patana (Valenciennes, 1847) FG
Family: Lebiasinidae—Marilyn Weitzman & Stanley H.
Weitzman
Copella arnoldi (Regan, 1912) PA GU SU FG
Copella carsevennensis (Regan, 1912) AP SU FG
Copella compta (Myers, 1927) CG VA BA
Copella eigenmanni (Regan, 1912) BO DA PA AP GU SU FG
Copella metae (Eigenmann, 1914) CG VA BO DA? BA
Copella nattereri (Steindachner, 1876) CG VA BO BA RO GU
Derhamia hoffmannorum Gery & Zarske, 2002 GU
Lebiasina provenzanoi Ardila Rodrıguez, 1999 BO
Lebiasina taphorni Ardila Rodrıguez, 2004 BO
Lebiasina uruyensis Fernandez-Yepez, 1967 BO
Lebiasina yuruaniensis Ardila Rodrıguez, 2000 BO
NUMBER 17 35
Nannostomus anduzei Fernandez & Weitzman, 1987 VA BO
Nannostomus beckfordi Gunther, 1872 AP GU SU FG
Nannostomus bifasciatus Hoedeman, 1954 SU FG
Nannostomus digrammus (Fowler, 1913) BA RO GU
Nannostomus espei (Meinken, 1956) GU
Nannostomus harrisoni (Eigenmann, 1909) GU
Nannostomus marginatus Eigenmann, 1909 CG VA BO DA GU SU
Nannostomus marilynae Weitzman & Cobb, 1975 CG VA BA
Nannostomus minimus Eigenmann, 1909 GU
Nannostomus trifasciatus Steindachner, 1876 RO? GU
Nannostomus unifasciatus Steindachner, 1876 CG VA BO DA GU GU
Piabucina unitaeniata Gunther, 1864 BO GU
Pyrrhulina filamentosa Valenciennes, 1847 VA BO DA GU SU FG
Pyrrhulina lugubris Eigenmann, 1922 CG VA BO DA
Pyrrhulina semifasciata Steindachner, 1876 RO GU
Pyrrhulina stoli Boeseman, 1953 GU SU
Family: Ctenoluciidae—Richard P. Vari
Boulengerella cuvieri (Agassiz, 1829) CG VA BO DA BA AP FG
Boulengerella lateristriga (Boulenger, 1895) VA BA
Boulengerella lucius (Cuvier, 1816) CG VA BO BA PA AP
Boulengerella maculata (Valenciennes, 1850) CG VA BO DA
Boulengerella xyrekes Vari, 1995 VA BO
Order: Siluriformes
Family: Cetopsidae—Richard P. Vari
Cetopsidium ferreirai Vari, Ferraris & de Pinna, 2005 PA
Cetopsidium minutum Vari, Ferraris & de Pinna, 2005 GU SU FG
Cetopsidium morenoi Vari, Ferraris & de Pinna, 2005 VA BO
Cetopsidium orientale Vari, Ferraris & de Pinna, 2005 AP GU SU FG
Cetopsidium pemon Vari, Ferraris & de Pinna, 2005 BO RO GU
Cetopsidium roae Vari, Ferraris & de Pinna, 2005 GU
Cetopsis coecutiens (Lichtenstein, 1819) CG VA BO
Cetopsis orinoco Vari, Ferraris & de Pinna, 2005 BO
Denticetopsis iwokrama Vari, Ferraris & de Pinna, 2005 GU
Denticetopsis macilenta Vari, Ferraris & de Pinna, 2005 GU
Denticetopsis praecox Vari, Ferraris & de Pinna, 2005 VA
Denticetopsis royeroi Ferraris, 1996 VA
Denticetopsis sauli Ferraris, 1996 VA
Helogenes castaneus (Dahl, 1960) CG
Helogenes marmoratus Gunther, 1863 VA BA GU SU FG
Helogenes uruyensis Fernandez-Yepez, 1967 BO
Family: Aspredinidae—John P. Friel
Acanthobunocephalus nicoi Friel, 1995 VA BO
Amaralia hypsiura (Kner, 1855) GU
Aspredinichthys filamentosus (Valenciennes, 1840) DA GU SU FG
Aspredinichthys tibicen (Valenciennes, 1840) DA GU SU FG
Aspredo aspredo (Linnaeus, 1758) DA GU SU FG
Bunocephalus aleuropsis Cope, 1870 VA BO
Bunocephalus amaurus Eigenmann, 1912 VA BO GU SU FG
36 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Bunocephalus chamaizelus Eigenmann, 1912 GU
Bunocephalus verrucosus (Walbaum, 1792) GU SU
Ernstichthys anduzei Fernandez-Yepez, 1953 VA BO
Hoplomyzon sexpapilostoma Taphorn & Marrero, 1990 VA BO
Platystacus cotylephorus Bloch, 1794 DA GU SU FG
Pseudobunocephalus lundbergi Friel, 2008 BO
Pterobunocephalus depressus (Haseman, 1911) VA BO
Family: Trichomycteridae—Wolmar Wosiacki
Ammoglanis pulex de Pinna & Winemiller, 2000 VA BO
Glanapteryx anguilla Myers, 1927 VA BO BA
Glanapteryx niobium de Pinna, 1998 BA
Haemomaster venezuelae Myers, 1927 VA BO
Henonemus taxistigmus (Fowler, 1914) GU
Henonemus triacanthopomus DoNascimiento & Provenzano, 2006 DA
Ituglanis amazonicus (Steindachner, 1882) FG
Ituglanis gracilior (Eigenmann, 1912) GU
Ituglanis metae (Eigenmann, 1917) CG VA? BO?
Ituglanis nebulosus de Pinna & Keith, 2003 FG
Megalocentor echthrus de Pinna & Britski, 1991 VA BO
Ochmacanthus alternus Myers, 1927 CG VA BO DA
Ochmacanthus flabelliferus Eigenmann, 1912 BO? GU
Ochmacanthus orinoco Myers, 1927 VA BO
Ochmacanthus reinhardtii (Steindachner, 1882) FG
Paracanthopoma parva Giltay, 1935 RO
Pseudostegophilus haemomyzon (Myers, 1942) VA BO
Pygidianops cuao Schaefer, Provenzano, de Pinna & Baskin,
2005
VA
Pygidianops magoi Schaefer, Provenzano, de Pinna & Baskin,
2005
BO DA
Schultzichthys bondi (Myers, 1942) VA BO
Stauroglanis gouldingi de Pinna, 1989 BA
Stegophilus septentrionalis Myers, 1927 VA BO
Trichomycterus celsae Lasso & Provenzano, 2002 BO
Trichomycterus conradi (Eigenmann, 1912) BO? GU
Trichomycterus guianensis (Eigenmann, 1909) VA BO GU FG
Trichomycterus lewi Lasso & Provenzano, 2002 BO
Trichomycterus santanderensis Castellanos-Morales, 2007 CG
Typhlobelus guacamaya Schaefer, Provenzano, de Pinna &
Baskin, 2005
VA
Typhlobelus lundbergi Schaefer, Provenzano, de Pinna & Baskin,
2005
BO DA
Vandellia beccarii Di Caporiacco, 1935 CG VA BO DA GU
Vandellia sanguinea Eigenmann, 1917 VA BO GU
Family: Callichthyidae—Roberto E. Reis
Callichthys callichthys (Linnaeus, 1758) CG VA BO DA GU SU FG
Callichthys serralabium Lehmann & Reis, 2004 VA BA
Corydoras aeneus (Gill, 1858) CG VA BO GU SU FG
Corydoras amapaensis Nijssen, 1972 AP FG
Corydoras approuaguensis Nijssen & Isbrucker, 1983 FG
Corydoras axelrodi Rossel, 1962 CG VA BO
NUMBER 17 37
Corydoras baderi Geisler, 1969 PA SU
Corydoras bicolor Nijssen & Isbrucker, 1967 SU
Corydoras blochi Nijssen, 1971 VA BO RO GU
Corydoras boehlkei Nijssen & Isbrucker, 1982 VA BO
Corydoras boesemani Nijssen & Isbrucker, 1967 SU
Corydoras bondi Gosline, 1940 VA BO GU SU
Corydoras breei Isbrucker & Nijssen, 1992 SU
Corydoras brevirostris Fraser-Brunner, 1947 VA BO SU
Corydoras concolor Weitzman, 1961 VA BO
Corydoras condiscipulus Nijssen & Isbrucker, 1980 AP FG
Corydoras coppenamensis Nijssen, 1970 SU
Corydoras delphax Nijssen & Isbrucker, 1983 CG VA
Corydoras ephippifer Nijssen, 1972 AP
Corydoras filamentosus Nijssen & Isbrucker, 1983 SU
Corydoras geoffroy La Cepede, 1803 SU FG
Corydoras griseus Holly, 1940 GU
Corydoras guianensis Nijssen, 1970 SU FG
Corydoras habrosus Weitzman, 1960 CG? VA BO
Corydoras heteromorphus Nijssen, 1970 SU
Corydoras melanistius Regan, 1912 GU SU FG
Corydoras melini Lonnberg & Rendahl, 1930 CG VA? BO?
Corydoras metae Eigenmann, 1914 CG VA BO
Corydoras nanus Nijssen & Isbrucker, 1967 SU FG
Corydoras oiapoquensis Nijssen, 1972 AP FG
Corydoras osteocarus Bohlke, 1951 VA BO SU
Corydoras oxyrhynchus Nijssen & Isbrucker, 1967 SU
Corydoras potaroensis Myers, 1927 GU
Corydoras punctatus (Bloch, 1794) SU FG
Corydoras sanchesi Nijssen & Isbrucker, 1967 SU
Corydoras saramaccensis Nijssen, 1970 SU
Corydoras septentrionalis Gosline, 1940 CG? VA BO
Corydoras sipaliwini Hoedeman, 1965 GU SU
Corydoras solox Nijssen & Isbrucker, 1983 AP FG
Corydoras spilurus Norman, 1926 SU FG
Corydoras surinamensis Nijssen, 1970 SU
Corydoras trilineatus Cope, 1872 SU
Hoplosternum littorale (Hancock, 1828) CG VA BO DA GU SU FG
Megalechis picta (Muller & Troschel, 1848) VA BO DA BA GU
Megalechis thoracata (Valenciennes, 1840) VA BO GU SU FG
Family: Loricariidae
Subfamily: Lithogeninae—Carl J. Ferraris, Jr.
Lithogenes villosus Eigenmann, 1909 GU
Lithogenes wahari Schaefer & Provenzano, 2008 VA
Subfamily: Hypoptopomatinae—Scott A. Schaefer
Acestridium dichromum Retzer, Nico & Provenzano, 1999 VA
Acestridium martini Retzer, Nico & Provenzano, 1999 VA BO
Hypoptopoma guianense Boeseman, 1974 GU SU
Nannoptopoma spectabile (Eigenmann, 1914) CG? VA? BO
38 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Niobichthys ferrarisi Schaefer & Provenzano, 1998 VA
Otocinclus huaorani Schaefer, 1997 CG
Otocinclus mariae Fowler, 1940 BA
Otocinclus vittatus Regan, 1904 CG? VA BO
Oxyropsis acutirostra Miranda Ribeiro, 1951 CG VA BO
Parotocinclus britskii Boeseman, 1974 VA BO GU SU
Parotocinclus collinsae Schmidt & Ferraris, 1985 GU
Parotocinclus eppleyi Schaefer & Provenzano, 1993 VA BO
Parotocinclus polyochrus Schaefer, 1988 VA
Subfamily: Loricariinae—Carl J. Ferraris, Jr.
Cteniloricaria platystoma (Gunther, 1868) SU
Farlowella nattereri Steindachner, 1910 GU
Farlowella oxyrryncha (Kner, 1853) VA BO
Farlowella reticulata Boeseman, 1971 GU SU FG
Farlowella rugosa Boeseman, 1971 GU SU FG
Farlowella vittata Myers, 1942 CG VA BO DA
Harttia fowleri (Pellegrin, 1908) AP FG
Harttia guianensis Rapp Py-Daniel & Oliveira, 2001 SU FG
Harttia maculata (Boeseman, 1971) SU FG
Harttia merevari Provenzano, Machado-Allison, Chernoff,
Willink & Petry, 2005
BO
Harttia surinamensis Boeseman, 1971 SU
Harttia trombetensis Rapp Py-Daniel & Oliveira, 2001 PA
Harttiella crassicauda (Boeseman, 1953) SU
Hemiloricaria castroi Isbrucker & Nijssen, 1984 PA
Hemiloricaria eigenmanni (Pellegrin, 1908) VA BO
Hemiloricaria fallax (Steindachner, 1915) RO GU
Hemiloricaria formosa Isbrucker & Njissen, 1979 CG VA BO DA
Hemiloricaria platyura (Muller & Troschel, 1848) GU FG
Hemiloricaria stewarti (Eigenmann, 1909) GU SU FG
Hemiodontichthys acipenserinus (Kner, 1853) AP GU FG
Limatulichthys griseus (Eigenmann, 1909) CG VA BO DA GU
Loricaria cataphracta Linnaeus, 1758 GU SU FG
Loricaria lundbergi Thomas & Rapp Py-Daniel, 2008 VA
Loricaria nickeriensis Isbrucker, 1979 SU FG
Loricaria parnahybae Steindachner, 1907 AP? FG
Loricaria simillima Regan, 1904 VA BO
Loricaria spinulifera Thomas & Rapp Py-Daniel, 2008 RO
Loricariichthys maculatus (Bloch, 1794) SU
Loricariichthys microdon (Eigenmann, 1909) GU
Metaloricaria nijsseni (Boeseman, 1976) SU
Metaloricaria paucidens Isbrucker, 1975 SU FG
Pseudoloricaria laeviuscula (Valenciennes, 1840) RO
Reganella depressa (Kner, 1853) RO
Sturisoma monopelte Fowler, 1914 GU
Subfamily: Hypostominae—Jonathan W. Armbruster & Claude
Weber
Acanthicus hystrix Spix & Agassiz, 1829 VA BO DA
Ancistrus hoplogenys (Gunther, 1864) GU
NUMBER 17 39
Ancistrus leucostictus (Gunther, 1864) GU SU? FG?
Ancistrus lithurgicus Eigenmann, 1912 GU
Ancistrus macrophthalmus (Pellegrin, 1912) VA BO
Ancistrus nudiceps (Muller & Troschel, 1848) GU
Ancistrus temminckii (Valenciennes, 1840) SU
Ancistrus triradiatus Eigenmann, 1918 CG VA BO DA
Baryancistrus beggini Lujan, Arce & Armbruster, 2009 CG VA
Baryancistrus demantoides Werneke, Sabaj, Lujan &
Armbruster, 2005
VA
Baryancistrus niveatus (Castelnau, 1855) PA
Chaetostoma jegui Rapp Py-Daniel, 1991 RO
Chaetostoma vasquezi Lasso & Provenzano, 1998 VA BO
Corymbophanes andersoni Eigenmann, 1909 GU
Corymbophanes kaiei Armbruster & Sabaj, 2000 GU
Dekeyseria niveata (La Monte, 1929) VA
Dekeyseria pulcher (Steindachner, 1915) CG VA BA?
Dekeyseria scaphirhyncha (Kner, 1854) BA
Exastilithoxus fimbriatus (Steindachner, 1915) BO
Exastilithoxus hoedemani Isbrucker & Nijssen, 1985 BA
Hemiancistrus guahiborum Werneke, Armbruster, Lujan &
Taphorn, 2005
VA BO
Hemiancistrus medians (Kner, 1854) SU FG
Hemiancistrus sabaji Armbruster, 2003 VA GU
Hemiancistrus subviridis Werneke, Sabaj, Lujan & Armbruster,
2005
VA
Hypancistrus contradens Armbruster, Lujan & Taphorn, 2007 VA
Hypancistrus debilittera Armbruster, Lujan & Taphorn, 2007 VA
Hypancistrus furunculus Armbruster, Lujan & Taphorn, 2007 VA
Hypancistrus inspector Armbruster, 2002 VA BO
Hypancistrus lunaorum Armbruster, Lujan & Taphorn, 2007 VA
Hypostomus coppenamensis Boeseman, 1969 SU
Hypostomus corantijni Boeseman, 1968 SU
Hypostomus crassicauda Boeseman, 1968 SU
Hypostomus gymnorhynchus (Norman, 1926) SU FG
Hypostomus hemicochliodon Armbruster, 2003 VA BO
Hypostomus hemiurus (Eigenmann, 1912) GU
Hypostomus macrophthalmus Boeseman, 1968 SU
Hypostomus macushi Armbruster & de Souza, 2005 GU
Hypostomus micromaculatus Boeseman, 1968 SU
Hypostomus nematopterus Isbrucker & Nijssen, 1984 FG
Hypostomus nickeriensis Boeseman, 1969 SU
Hypostomus occidentalis Boeseman, 1968 SU
Hypostomus paucimaculatus Boeseman, 1968 SU
Hypostomus plecostomoides (Eigenmann, 1922) BO
Hypostomus plecostomus (Linnaeus, 1758) GU SU
Hypostomus pseudohemiurus Boeseman, 1968 SU
Hypostomus rhantos Armbruster, Tansey & Lujan, 2007 VA
Hypostomus saramaccensis Boeseman, 1968 SU
Hypostomus sculpodon Armbruster, 2003 VA
40 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Hypostomus sipaliwini Boeseman, 1968 SU
Hypostomus squalinus (Jardine, 1841) RO GU
Hypostomus surinamensis Boeseman, 1968 SU
Hypostomus tapanahoniensis Boeseman, 1969 SU
Hypostomus taphorni (Lilyestrom, 1984) BO GU
Hypostomus tenuis (Boeseman, 1968) SU?
Hypostomus ventromaculatus Boeseman, 1968 SU FG
Hypostomus villarsi (Lutken, 1874) VA BO
Hypostomus watwata Hancock, 1828 DA GU SU FG
Lasiancistrus schomburgkii (Gunther, 1864) GU
Lasiancistrus tentaculatus Armbruster, 2005 VA BO
Leporacanthicus galaxias Isbrucker & Nijssen, 1989 VA BO
Leporacanthicus triactis Isbrucker, Nijssen & Nico, 1992 CG VA BO
Lithoxus boujardi Muller & Isbrucker, 1993 FG
Lithoxus bovallii (Regan, 1906) GU
Lithoxus jantjae Lujan, 2008 VA
Lithoxus lithoides Eigenmann, 1910 GU SU
Lithoxus pallidimaculatus Boeseman, 1982 SU
Lithoxus planquettei Boeseman, 1982 FG
Lithoxus stocki Nijssen & Isbrucker, 1990 FG
Lithoxus surinamensis Boeseman, 1982 SU
Neblinichthys pilosus Ferraris, Isbrucker & Nijssen, 1986 VA
Neblinichthys roraima Provenzano, Lasso & Ponte, 1995 BO
Neblinichthys yaravi (Steindachner, 1915) BO
Panaque maccus Schaefer & Stewart, 1993 VA BO
Panaque nigrolineatus (Peters, 1877) CG VA BO
Peckoltia braueri (Eigenmann, 1912) GU
Peckoltia brevis (La Monte, 1935) CG?
Peckoltia cavatica Armbruster & Werneke, 2005 GU
Peckoltia lineola Armbruster, 2008 CG VA
Peckoltia vittata (Steindachner, 1881) CG? VA PA
Pseudacanthicus fordii (Gunther, 1868) SU
Pseudacanthicus leopardus (Fowler, 1914) GU
Pseudacanthicus serratus (Valenciennes, 1840) SU FG
Pseudancistrus barbatus (Valenciennes, 1840) GU SU FG
Pseudancistrus brevispinis (Heitmans, Nijssen & Isbrucker, 1983) SU FG
Pseudancistrus coquenani (Steindachner, 1915) BO
Pseudancistrus corantijniensis De Chambrier & Montoya-
Burgos, 2008
SU
Pseudancistrus depressus (Gunther, 1868) SU
Pseudancistrus guentheri (Regan, 1904) GU
Pseudancistrus longispinis (Heitmans, Nijssen & Isbrucker, 1983) FG
Pseudancistrus macrops (Lutken, 1874) SU
Pseudancistrus megacephalus (Gunther, 1868) GU SU
Pseudancistrus niger (Norman, 1926) FG
Pseudancistrus nigrescens Eigenmann, 1912 GU
Pseudancistrus orinoco Isbrucker, Nijssen & Cala, 1988 CG VA BO
Pseudancistrus pectegenitor Lujan, Armbruster & Sabaj, 2007 VA
Pseudancistrus reus Armbruster & Taphorn, 2008 BO
NUMBER 17 41
Pseudancistrus sidereus Armbruster, 2004 VA
Pseudancistrus yekuana Lujan, Armbruster & Sabaj, 2007 VA
Pseudolithoxus anthrax (Armbruster & Provenzano, 2000) VA BO
Pseudolithoxus dumus (Armbruster & Provenzano, 2000) VA BO
Pseudolithoxus nicoi (Armbruster & Provenzano, 2000) VA
Pseudolithoxus tigris (Armbruster & Provenzano, 2000) VA BO
Pterygoplichthys gibbiceps (Kner, 1854) CG VA BO
Pterygoplichthys multiradiatus (Hancock, 1828) VA BO GU?
Family: Pseudopimelodidae—Oscar A. Shibbata
Batrochoglanis raninus (Valenciennes, 1840) GU SU FG
Batrochoglanis villosus (Eigenmann, 1912) VA BO GU SU
Cephalosilurus albomarginatus (Eigenmann, 1912) GU
Cephalosilurus nigricaudus (Mees, 1974) SU
Microglanis poecilus Eigenmann, 1912 GU FG
Microglanis secundus Mees, 1974 CG? VA BO GU SU
Pseudopimelodus bufonius (Valenciennes, 1840) CG? VA BO GU SU FG
Family: Heptapteridae—Updated from Cloffsca account (Flavio
A. Bockmann & Gizelani M. Guazzeli)
Brachyglanis frenata Eigenmann, 1912 VA BO BA GU
Brachyglanis magoi Fernandez-Yepez, 1967 VA BO
Brachyglanis melas Eigenmann, 1912 GU
Brachyglanis microphthalmus Bizerril, 1991 PA
Brachyglanis phalacra Eigenmann, 1912 GU
Brachyrhamdia heteropleura (Eigenmann, 1912) GU SU
Brachyrhamdia imitator Myers, 1927 VA BO
Cetopsorhamdia insidiosa (Steindachner, 1917) RO
Cetopsorhamdia orinoco Schultz, 1944 BO
Chasmocranus brevior Eigenmann, 1912 GU SU FG
Chasmocranus chimantanus Inger, 1956 BO
Chasmocranus longior Eigenmann, 1912 VA BO BA GU SU
Chasmocranus surinamensis (Bleeker, 1862) SU
Gladioglanis machadoi Ferraris & Mago-Leccia, 1989 VA BO BA?
Goeldiella eques (Muller & Troschel, 1848) CG VA BO BA? GU
Heptapterus bleekeri Boeseman, 1953 AP? SU FG
Heptapterus tapanahoniensis Mees, 1967 SU FG
Heptapterus tenuis Mees, 1986 FG
Imparfinis hasemani Steindachner, 1917 RO
Imparfinis pijpersi (Hoedeman, 1961) SU
Imparfinis pristos Mees & Cala, 1989 CG VA BO
Imparfinis pseudonemacheir Mees & Cala, 1989 CG VA BO
Leptorhamdia essequibensis (Eigenmann, 1912) GU
Leptorhamdia marmorata Myers, 1928 CG? VA BO BA
Mastiglanis asopos Bockmann, 1994 VA BO PA
Medemichthys guayaberensis (Dahl, 1961) CG
Myoglanis aspredinoides DoNascimiento & Lundberg, 2005 VA
Myoglanis potaroensis Eigenmann, 1912 GU
Nemuroglanis pauciradiatus Ferraris, 1988 CG VA BO BA
Phenacorhamdia anisura (Mees, 1987) VA BO
Phenacorhamdia macarenensis Dahl, 1961 CG
42 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Phenacorhamdia provenzanoi DoNascimiento & Milani, 2008 VA BO
Pimelodella altipinnis (Steindachner, 1864) GU
Pimelodella cristata (Muller & Troschel, 1848) GU FG
Pimelodella cruxenti Fernandez-Yepez, 1950 VA BO
Pimelodella figueroai Dahl, 1961 CG
Pimelodella geryi Hoedeman, 1961 SU
Pimelodella linami Schultz, 1944 VA BO
Pimelodella macturki Eigenmann, 1912 GU SU FG
Pimelodella megalops Eigenmann, 1912 GU FG
Pimelodella pallida Dahl, 1961 CG
Pimelodella procera Mees, 1983 FG
Pimelodella wesselii (Steindachner, 1877) GU
Rhamdella leptosoma Fowler, 1914 GU
Rhamdia foina (Muller & Troschel, 1848) RO PA GU
Rhamdia laukidi Bleeker, 1858 CG VA BO GU
Rhamdia muelleri (Gunther, 1864) VA BO DA PA GU
Rhamdia quelen (Quoy & Gaimard, 1824) CG VA BO DA GU SU FG
Family: Pimelodidae—John G. Lundberg
Brachyplatystoma filamentosum (Lichtenstein, 1819) CG VA BO DA GU SU FG
Brachyplatystoma juruense (Boulenger, 1898) CG? VA BO DA
Brachyplatystoma platynemum (Boulenger, 1898) CG VA BO DA
Brachyplatystoma rousseauxii (Castelnau, 1855) BO DA FG
Brachyplatystoma vaillantii (Valenciennes, 1840) CG VA BO DA GU SU FG
Calophysus macropterus (Lichtenstein, 1819) CG VA BO DA PA
Duopalatinus peruanus Eigenmann & Allen, 1942 BO
Exallodontus aguanai Lundberg, Mago-Leccia & Nass, 1991 BO DA
Hemisorubim platyrhynchos (Valenciennes, 1840) CG VA BO DA GU SU FG
Hypophthalmus edentatus Spix & Agassiz, 1829 VA BO DA GU SU
Hypophthalmus fimbriatus Kner, 1858 BO
Hypophthalmus marginatus Valenciennes, 1840 SU FG
Leiarius marmoratus (Gill, 1870) CG VA BO
Leiarius pictus (Muller & Troschel, 1849) CG? VA BO GU
Megalonema amaxanthum Lundberg & Dahdul, 2008 GU
Megalonema platycephalum Eigenmann, 1912 CG VA BO GU
Phractocephalus hemioliopterus (Bloch & Schneider, 1801) CG VA BO DA GU
Pimelodina flavipinnis Steindachner, 1877 BO DA
Pimelodus albofasciatus Mees, 1974 CG VA BO BA SU
Pimelodus blochii Valenciennes, 1840 CG VA BO DA GU SU FG
Pimelodus ornatus Kner, 1858 CG VA BO GU SU FG
Pimelodus pictus Steindachner, 1877 BO
Pinirampus pirinampu (Spix & Agassiz, 1829) CG VA BO DA GU
Platynematichthys notatus (Jardine, 1841) CG VA BO DA
Platysilurus mucosus (Vaillant, 1880) CG VA BO DA
Propimelodus eigenmanni (van der Stigchel, 1946) AP FG
Pseudoplatystoma fasciatum (Linnaeus, 1766) GU SU FG
Pseudoplatystoma metaense Buitrago-Suarez & Burr, 2007 BO DA
Pseudoplatystoma orinocoense Buitrago-Suarez & Burr, 2007 VA BO DA
Sorubim elongatus Littmann, Burr, Schmidt & Isern, 2001 CG VA BO GU
NUMBER 17 43
Sorubim lima (Bloch & Schneider, 1801) CG VA BO DA
Sorubimichthys planiceps (Spix & Agassiz, 1829) CG VA BO DA
Zungaro zungaro (Humboldt, 1821) CG VA BO DA BA GU
Family: Doradidae—Mark H. Sabaj Perez
Acanthodoras cataphractus (Linnaeus, 1758) CG VA BO? BA RO PA AP GU SU FG
Acanthodoras spinosissimus (Eigenmann & Eigenmann, 1888) CG VA BO? DA? BA RO GU
Agamyxis albomaculatus (Peters, 1877) CG VA BO DA BA?
Amblydoras affinis (Kner, 1855) RO GU
Amblydoras bolivarensis (Fernandez-Yepez, 1968) CG VA BO DA? BA?
Amblydoras gonzalezi (Fernandez-Yepez, 1968) CG VA BO DA? BA?
Anadoras regani (Steindachner, 1908) PA AP FG
Anduzedoras oxyrhynchus (Valenciennes, 1821) CG VA BA
Centrodoras hasemani (Steindachner, 1915) CG? VA BA
Doras carinatus (Linnaeus, 1766) BO AP GU SU FG
Doras micropoeus (Eigenmann, 1912) GU SU FG
Doras phlyzakion Sabaj Perez & Birindelli, 2008 RO
Hassar orestis (Steindachner, 1875) CG VA BO DA BA RO? PA? GU
Leptodoras cataniai Sabaj, 2005 VA BA RO?
Leptodoras copei (Fernandez-Yepez, 1968) CG VA BO BA RO? PA?
Leptodoras hasemani (Steindachner, 1915) CG VA BO DA BA? RO GU
Leptodoras linnelli Eigenmann, 1912 VA BO BA? RO GU
Leptodoras praelongus (Myers & Weitzman, 1956) CG VA BA RO PA?
Leptodoras rogersae Sabaj, 2005 CG VA BO DA
Megalodoras guayoensis (Fernandez-Yepez, 1968) CG? VA? BO DA
Megalodoras uranoscopus (Eigenmann & Eigenmann, 1888) CG? VA? BA? RO? PA? GU
Nemadoras leporhinus (Eigenmann, 1912) CG VA BO BA? RO PA GU
Nemadoras trimaculatus (Boulenger, 1898) CG VA BO BA RO GU
Opsodoras morei (Steindachner, 1881) CG VA BO? BA RO? PA? GU
Opsodoras ternetzi Eigenmann, 1925 CG VA BO DA BA RO PA GU
Orinocodoras eigenmanni Myers, 1927 BO DA
Oxydoras niger (Valenciennes, 1821) CG? VA? BA? RO? GU
Oxydoras sifontesi Fernandez-Yepez, 1968 CG? VA? BO DA
Physopyxis ananas Sousa & Rapp Py-Daniel 2005 CG VA BA RO PA GU
Physopyxis cristata Sousa & Rapp Py-Daniel 2005 BA RO?
Platydoras armatulus (Valenciennes 1840) BO DA
Platydoras costatus (Linnaeus, 1758) SU FG
Platydoras hancockii (Valenciennes 1840) CG VA BO? BA RO GU
Pterodoras granulosus (Valenciennes, 1821) CG? VA? BA? RO? PA? GU SU
Pterodoras rivasi (Fernandez-Yepez, 1950) CG? VA? BO DA
Rhinodoras armbrusteri Sabaj, Taphorn & Castillo, 2008 GU
Rhynchodoras woodsi Glodek, 1976 RO GU
Scorpiodoras heckelii (Kner, 1855) CG VA BO BA
Trachydoras brevis (Kner, 1853) BA RO GU
Trachydoras microstomus (Eigenmann, 1912) CG VA BO DA? BA RO PA GU
Family: Auchenipteridae—Carl J. Ferraris, Jr.
Ageneiosus inermis (Linnaeus, 1766) CG VA BO DA GU SU FG
Ageneiosus marmoratus Eigenmann, 1912 GU SU
Ageneiosus piperatus (Eigenmann, 1912) RO GU
44 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Ageneiosus polystictus Steindachner, 1915 RO
Ageneiosus ucayalensis Castelnau, 1855 VA BO DA GU SU
Ageneiosus vittatus Steindachner, 1908 BO DA
Asterophysus batrachus Kner, 1858 VA BO BA?
Auchenipterus ambyiacus Fowler, 1915 CG? VA BO DA GU
Auchenipterus brevior Eigenmann, 1912 GU
Auchenipterus demerarae Eigenmann, 1912 BO GU
Auchenipterus dentatus Valenciennes, 1840 SU FG
Auchenipterus nuchalis (Spix & Agassiz, 1829) RO? FG
Centromochlus concolor (Mees, 1974) SU
Centromochlus existimatus Mees, 1974 VA BO
Centromochlus punctatus (Mees, 1974) SU
Centromochlus reticulatus (Mees, 1974) GU
Entomocorus gameroi Mago-Leccia, 1984 VA BO DA
Gelanoglanis nanonocticolus Soares-Porto, Walsh, Nico & Netto,
1999
VA BO
Glanidium leopardum (Hoedeman, 1961) GU SU FG
Liosomadoras oncinus (Jardine, 1841) RO
Pseudepapterus gracilis Ferraris & Vari, 2000 BO
Tatia brunnea Mees, 1974 SU FG
Tatia creutzbergi (Boeseman, 1953) SU
Tatia galaxias Mees, 1974 VA BO DA
Tatia gyrina (Eigenmann & Allen, 1942) SU
Tatia intermedia (Steindachner, 1877) GU SU FG
Tatia meesi Sarmento-Soares & Martins-Pinheiro, 2008 GU
Tatia musaica Royero, 1992 VA BO
Tatia nigra Sarmento-Soares & Martins-Pinheiro, 2008 PA
Tatia strigata Soares-Porto, 1995 VA BO BA
Tetranematichthys wallacei Vari & Ferraris, 2006 VA BO BA
Trachelyichthys decaradiatus Mees, 1974 GU
Trachelyopterichthys anduzei Ferraris & Fernandez, 1987 VA BO
Trachelyopterichthys taeniatus (Kner, 1858) CG VA BO BA?
Trachelyopterus ceratophysus (Kner, 1858) RO
Trachelyopterus coriaceus Valenciennes, 1840 FG
Trachelyopterus galeatus (Linnaeus, 1766) SU FG
Trachycorystes trachycorystes Valenciennes, 1840 GU
Order: Gymnotiformes
Family: Gymnotidae—Ricardo Campos-da-Paz
Electrophorus electricus (Linnaeus, 1766) CG VA BO DA GU SU FG
Gymnotus anguillaris Hoedeman, 1962 SU FG
Gymnotus carapo Linnaeus, 1758 VA BO DA FG
Gymnotus cataniapo Mago-Leccia, 1994 BO SU
Gymnotus coropinae Hoedeman, 1962 GU SU
Gymnotus pedanopterus Mago-Leccia, 1994 VA BO BA
Gymnotus stenoleucus Mago-Leccia, 1994 VA BO DA
Family: Sternopygidae—James S. Albert
Archolaemus blax Korringa, 1970 RO AP? FG
Distocyclus conirostris (Eigenmann & Allen, 1942) BO DA
NUMBER 17 45
Eigenmannia limbata (Schreiner & Miranda Ribeiro, 1903) BO
Eigenmannia macrops (Boulenger, 1897) BA GU
Eigenmannia nigra Mago-Leccia, 1994 CG VA BO BA GU
Eigenmannia virescens (Valenciennes, 1842) CG VA BO DA BA AP GU SU FG
Rhabdolichops caviceps (Fernandez-Yepez, 1968) BO
Rhabdolichops eastwardi Lundberg & Mago-Leccia, 1986 BO DA
Rhabdolichops electrogrammus Lundberg & Mago-Leccia, 1986 BO DA BA RO
Rhabdolichops jegui Keith & Meunier, 2000 FG
Rhabdolichops stewarti Lundberg & Mago-Leccia, 1986 VA BO
Rhabdolichops troscheli (Kaup, 1856) BO DA
Rhabdolichops zareti Lundberg & Mago-Leccia, 1986 VA BO DA
Sternopygus astrabes Mago-Leccia, 1994 CG? VA BO BA
Sternopygus macrurus (Bloch & Schneider, 1801) CG VA BO DA BA? RO? PA? AP? GU SU FG
Family: Rhamphichthyidae—Carl J. Ferraris, Jr.
Gymnorhamphichthys hypostomus Ellis, 1912 CG VA DA
Gymnorhamphichthys rondoni (Miranda Ribeiro, 1920) VA BO GU SU
Iracema caiana Triques, 1996 BA RO
Rhamphichthys apurensis (Fernandez-Yepez, 1968) VA BO DA
Rhamphichthys rostratus (Linnaeus, 1766) GU SU
Family: Hypopomidae—James S. Albert
Brachyhypopomus beebei (Schultz, 1944) CG VA BO DA GU SU
Brachyhypopomus brevirostris (Steindachner, 1868) CG VA BO DA GU SU
Brachyhypopomus pinnicaudatus (Hopkins, 1991) BO GU SU
Hypopomus artedi (Kaup, 1856) PA? AP? GU SU FG
Hypopygus lepturus Hoedeman, 1962 CG VA BO DA BA? SU FG
Hypopygus neblinae Mago-Leccia, 1994 CG VA BO DA
Microsternarchus bilineatus Fernandez-Yepez, 1968 CG VA BO BA
Racenisia fimbriipinna Mago-Leccia, 1994 VA BO
Steatogenys duidae (La Monte, 1929) VA BO
Steatogenys elegans (Steindachner, 1880) CG VA BO DA
Family: Apteronotidae—James S. Albert
Adontosternarchus clarkae Mago-Leccia, Lundberg & Baskin,
1985
CG VA BA RO
Adontosternarchus devenanzii Mago-Leccia, Lundberg & Baskin,
1985
BO DA
Adontosternarchus sachsi (Peters, 1877) BO DA
Apteronotus albifrons (Linnaeus, 1766) VA BO DA GU SU FG
Apteronotus leptorhynchus (Ellis, 1912) VA BO DA BA GU SU FG
Compsaraia compsus (Mago-Leccia, 1994) BO DA
Megadontognathus cuyuniense Mago-Leccia, 1994 BO
Platyurosternarchus crypticus de Santana &Vari, 2009 RO GU
Platyurosternarchus macrostomus (Gunther, 1870) VA BO DA GU
Porotergus gymnotus Ellis, 1912 GU FG
Sternarchella orthos Mago-Leccia, 1994 VA BO DA
Sternarchella sima Starks, 1913 BO DA
Sternarchorhamphus muelleri (Steindachner, 1881) BO DA
Sternarchorhynchus gnomus de Santana & Taphorn, 2006 BO
Sternarchorhynchus oxyrhynchus (Muller & Troschel, 1849) BO GU FG
Sternarchorhynchus roseni Mago-Leccia, 1994 BO DA
46 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Order: Cyprinodontiformes
Family: Rivulidae—Wilson J. E. M. Costa
Austrofundulus rupununi Hrbek, Taphorn &Thomerson, 2005 GU
Austrofundulus transilis Myers, 1932 VA BO
Gnatholebias hoignei (Thomerson, 1974) VA BO
Gnatholebias zonatus (Myers, 1935) VA BO
Kryptolebias sepia Vermeulen & Hrbek, 2005 SU
Micromoema xiphophora (Thomerson & Taphorn, 1992) VA BO
Rachovia maculipinnis (Radda, 1964) BO DA
Rachovia stellifer (Thomerson & Turner, 1973) VA BO
Renova oscari Thomerson & Taphorn, 1995 VA BO
Rivulus agilae Hoedeman, 1954 GU SU FG
Rivulus altivelis Huber, 1992 CG
Rivulus amphoreus Huber, 1979 SU
Rivulus breviceps Eigenmann, 1909 GU
Rivulus caurae Radda, 2004 BO
Rivulus cladophorus Huber, 1991 FG
Rivulus corpulentus Thomerson & Taphorn, 1993 CG
Rivulus deltaphilus Seegers, 1983 VA BO DA
Rivulus frenatus Eigenmann, 1912 GU SU
Rivulus gaucheri Keith, Nandrin & Le-Bail 2006 FG
Rivulus geayi Vaillant, 1899 AP FG
Rivulus gransabanae Lasso, Taphorn & Thomerson, 1992 BO
Rivulus holmiae Eigenmann, 1909 GU
Rivulus igneus Huber, 1991 AP FG
Rivulus immaculatus Thomerson, Nico & Taphorn, 1991 BO
Rivulus lanceolatus Eigenmann, 1909 GU
Rivulus lungi Berkenkamp, 1984 FG
Rivulus lyricauda Thomerson, Berkenkamp & Taphorn, 1991 VA BO
Rivulus mahdiaensis Suijker & Collier, 2006 GU
Rivulus manaensis Hoedeman, 1961 FG
Rivulus mazaruni Myers, 1924 GU
Rivulus nicoi Thomerson & Taphorn, 1992 VA BO
Rivulus sape Lasso-Alcala, Taphorn, Lasso & Leon-Mata, 2006 BO
Rivulus stagnatus Eigenmann, 1909 GU SU
Rivulus tecminae Thomerson, Nico & Taphorn, 1992 VA BO
Rivulus torrenticola Vermeulen & Isbrucker, 2000 GU
Rivulus waimacui Eigenmann, 1909 GU
Rivulus xiphidius Huber, 1979 AP FG
Terranatos dolichopterus (Weitzman & Wourms, 1967) VA BO
Family: Poeciliidae—Paulo H. J. Lucinda
Fluviphylax palikur Costa & Le Bail, 1999 AP FG
Fluviphylax pygmaeus (Myers & Carvalho, 1955) CG VA BA PA
Fluviphylax simplex Costa, 1996 PA
Micropoecilia bifurca (Eigenmann, 1909) GU SU FG
Micropoecilia parae (Eigenmann, 1894) AP? GU SU FG
Micropoecilia picta (Regan, 1913) GU SU FG
Poecilia reticulata Peters, 1859 BO DA AP? GU SU FG
NUMBER 17 47
Poecilia vivipara Bloch & Schneider, 1801 DA GU SU FG
Tomeurus gracilis Eigenmann, 1909 DA AP GU SU FG
Order: Beloniformes
Family: Belonidae—Bruce B. Collette
Belonion dibranchodon Collette, 1966 VA BA
Potamorrhaphis guianensis (Jardine, 1843) CG VA BA RO PA GU SU FG
Potamorrhaphis petersi Collette, 1974 CG VA BO BA
Pseudotylosurus microps (Gunther, 1866) VA BO DA GU SU
Family: Hemiramphidae—Bruce B. Collette
Hyporhamphus brederi (Fernandez-Yepez, 1948) VA BO DA
Order: Synbranchiformes
Family: Synbranchidae—Sven O. Kullander
Synbranchus marmoratus Bloch, 1795 CG VA BO DA BA RO AP GU SU FG
Order: Perciformes
Family: Sciaenidae—Ning Labbish Chao & Lilian Casatti
Pachypops fourcroi (Lacpede, 1802) VA BO DA AP GU SU FG
Pachypops pigmaeus Casatti, 2002 RO
Pachypops trifilis (Muller & Troschel, 1849) BA RO GU SU
Pachyurus gabrielensis Casatti, 2001 VA BO BA
Pachyurus schomburgkii Gunther, 1860 VA BO
Petilipinnis grunniens (Jardine, 1843) BO GU
Plagioscion auratus (Castelnau, 1855) BO DA GU SU FG
Plagioscion squamosissimus (Heckel, 1840) CG VA BO DA SU FG
Plagioscion surinamensis (Bleeker, 1973) CG DA SU FG
Family: Polycentridae—Richard P. Vari
Monocirrhus polyacanthus Heckel, 1840 BO RO
Polycentrus schomburgkii Muller & Troschel, 1849 DA AP GU SU FG
Family: Cichlidae—Sven O. Kullander
Acaronia nassa (Heckel, 1840) RO AP GU FG
Acaronia vultuosa Kullander, 1989 CG VA BO BA
Aequidens chimantanus Inger, 1956 BO
Aequidens diadema (Heckel, 1840) CG VA BO BA
Aequidens paloemeuensis Kullander & Nijssen, 1989 SU
Aequidens potaroensis Eigenmann, 1912 GU
Aequidens tetramerus (Heckel, 1840) CG VA BO DA PA AP GU SU FG
Aequidens tubicen Kullander & Ferreira, 1991 PA
Apistogramma angayuara Kullander & Ferreira, 2005 PA
Apistogramma diplotaenia Kullander, 1987 VA BO BA
Apistogramma gibbiceps Meinken, 1969 RO
Apistogramma gossei Kullander, 1982 AP FG
Apistogramma hoignei Meinken, 1965 VA BO
Apistogramma hongsloi Kullander, 1979 VA
Apistogramma iniridae Kullander, 1979 CG
Apistogramma inornata Staeck, 2003 BO
Apistogramma ortmanni (Eigenmann, 1912) BO? GU SU
Apistogramma rupununi Fowler, 1914 RO GU
Apistogramma salpinction Kullander & Ferreira, 2005 PA
Apistogramma steindachneri (Regan, 1908) GU SU
48 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Apistogramma velifera Staeck, 2003 VA
Apistogramma viejita Kullander, 1979 CG
Apistogramma wapisana Romer, Hahn & Conrad, 2006 RO
Astronotus ocellatus (Agassiz, 1831) FG
Biotodoma cupido (Heckel, 1840) GU
Biotodoma wavrini (Gosse, 1963) CG VA BO BA
Biotoecus dicentrarchus Kullander, 1989 CG VA BO DA
Caquetaia spectabilis (Steindachner, 1875) BA RO AP GU
Chaetobranchus flavescens Heckel, 1840 CG VA BO AP GU SU FG
Cichla intermedia Machado-Allison, 1971 VA BO
Cichla jariina Kullander & Ferreira, 2006 PA
Cichla monoculus Spix & Agassiz, 1831 PA AP FG
Cichla nigromaculata Jardine, 1843 VA
Cichla ocellaris Schneider, 1801 RO GU SU FG
Cichla orinocensis Humboldt, 1821 CG VA BO DA
Cichla temensis Humboldt, 1821 CG VA BO DA BA
Cichla thyrorus Kullander & Ferreira, 2006 PA
Cichlasoma amazonarum Kullander, 1983 AP FG
Cichlasoma bimaculatum (Linnaeus, 1758) BO RO GU SU FG
Cichlasoma orinocense Kullander, 1983 CG VA BO
Cleithracara maronii (Steindachner, 1881) DA GU SU FG
Crenicara punctulatum (Gunther, 1863) GU
Crenicichla albopunctata Pellegrin, 1904 GU SU FG
Crenicichla alta Eigenmann, 1912 RO GU
Crenicichla coppenamensis Ploeg, 1987 SU
Crenicichla heckeli Ploeg, 1989 PA
Crenicichla hummelincki Ploeg, 1991 PA
Crenicichla johanna Heckel, 1840 CG VA BO AP GU FG
Crenicichla lugubris Heckel, 1840 BA RO GU SU
Crenicichla multispinosa Pellegrin, 1903 SU FG
Crenicichla nickeriensis Ploeg, 1987 SU
Crenicichla pydanielae Ploeg, 1991 PA
Crenicichla reticulata (Heckel, 1840) GU
Crenicichla saxatilis (Linnaeus, 1758) DA GU SU FG
Crenicichla sipaliwini Ploeg, 1987 SU
Crenicichla ternetzi Norman, 1926 FG
Crenicichla tigrina Ploeg, Jegu & Ferreira, 1991 PA
Crenicichla vaillanti Pellegrin, 1903 GU FG
Crenicichla virgatula Ploeg, 1991 RO
Crenicichla wallacii Regan, 1905 GU
Crenicichla zebrina Montana, Lopez-Fernandez & Taphorn,
2008
VA
Dicrossus filamentosus (Ladiges, 1958) CG VA BO
Dicrossus gladicauda Schindler & Staeck, 2008 CG
Geophagus abalios Lopez-Fernandez & Taphorn, 2004 VA BO
Geophagus brachybranchus Kullander & Nijssen, 1989 GU SU
Geophagus brokopondo Kullander & Nijssen, 1989 SU
Geophagus camopiensis Pellegrin, 1903 AP FG
Geophagus dicrozoster Lopez-Fernandez & Taphorn, 2004 VA BO
NUMBER 17 49
Geophagus gottwaldi Schindler & Staeck, 2006 VA
Geophagus grammepareius Kullander & Taphorn, 1992 VA BO
Geophagus harreri Gosse, 1976 SU FG
Geophagus surinamensis (Bloch, 1791) SU FG
Geophagus taeniopareius Kullander & Royero, 1992 VA BO
Geophagus winemilleri Lopez-Fernandez & Taphorn, 2004 VA
Guianacara (Guianacara) cuyunii Lopez-Fernandez, Taphorn, &
Kullander, 2006
BO
Guianacara (Guianacara) geayi (Pellegrin, 1902) AP FG
Guianacara (Guianacara) owroewefi Kullander & Nijssen, 1989 SU FG
Guianacara (Guianacara) sphenozona Kullander & Nijssen, 1989 GU SU
Guianacara (Guianacara) stergiosi Lopez-Fernandez, Taphorn,
& Kullander, 2006
BO
Guianacara (Oelemaria) oelemariensis Kullander & Nijssen,
1989
SU
Heros severus Heckel, 1840 CG VA BO BA
Hoplarchus psittacus (Heckel, 1840) CG VA BO BA
Hypselecara coryphaenoides (Heckel, 1840) CG VA BO BA
Hypselecara temporalis (Gunther, 1862) AP
Ivanacara adoketa (Kullander & Prada-Pedreros, 1993) BA
Ivanacara bimaculata Eigenmann, 1912 GU
Krobia guianensis (Regan, 1905) GU SU
Krobia itanyi (Puyo, 1943) SU FG
Laetacara fulvipinnis Staeck & Schindler, 2007 CG VA BA
Mazarunia mazarunii Kullander, 1990 GU
Mesonauta egregius Kullander & Silfvergrip, 1991 CG
Mesonauta guyanae Schindler, 1998 RO GU
Mesonauta insignis (Heckel, 1840) CG VA BO BA
Nannacara anomala Regan, 1905 GU SU
Nannacara aureocephalus Allgayer, 1983 FG
Pterophyllum altum Pellegrin, 1903 CG VA
Pterophyllum leopoldi (Gosse, 1963) GU
Pterophyllum scalare (Schultze, 1823) BA AP GU SU FG
Retroculus septentrionalis Gosse, 1971 AP FG
Satanoperca acuticeps (Heckel, 1840) RO
Satanoperca daemon (Heckel, 1840) CG VA BO BA
Satanoperca jurupari (Heckel, 1840) BA PA AP
Satanoperca leucosticta (Muller & Troschel, 1849) GU SU
Satanoperca lilith Kullander & Ferreira, 1988 BA RO PA
Satanoperca mapiritensis (Fernandez-Yepez, 1950) VA BO DA
Uaru fernandezyepezi Stawikowski, 1989 VA
Family: Gobiidae—Sven O. Kullander
Awaous flavus Valenciennes, 1837 DA AP GU SU FG
Ctenogobius claytonii (Meek, 1902) DA
Dormitator maculatus (Bloch, 1792) GU SU FG
Eleotris amplyopsis (Cope, 1871) DA GU SU FG
Eleotris pisonis (Gmelin, 1789) DA GU SU FG
Gobioides broussonnetii La Cepede, 1800 DA FG
Gobioides grahamae (Palmer & Wheeler, 1995) GU SU FG
50 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
Gobiomorus dormitor La Cepede, 1800 GU SU
Microphilypnus ternetzi Myers, 1927 VA BO DA
Order: Pleuronectiformes
Family: Achiridae—Robson T. C. Ramos
Achirus achirus (Linnaeus, 1758) GU FG
Achirus novoae Cervigon, 1982 BO DA
Apionichthys dumerili Kaup, 1858 DA PA AP GU SU FG
Apionichthys finis Eigenmann, 1912 RO GU
Apionichthys menezesi Ramos, 2003 VA BO
Hypoclinemus mentalis (Gunther, 1862) VA BO BA RO AP GU
Order: Tetraodontiformes
Family: Tetraodontidae—Carl J. Ferraris, Jr. & Sven O.
Kullander
Colomesus asellus (Muller & Troschel, 1849) BO DA GU
Order: Lepidosireniformes
Family: Lepidosirenidae—Based on Cloffsca account (Gloria
Arratia)
Lepidosiren paradoxa Fitzinger, 1837 CG? VA? BA? FG
NUMBER 17 51
PHOTOGRAPHIC ATLAS OF FISHES OF THE GUIANA SHIELD
MARK H. SABAJ PEREZ
Introduction
The last decade or so has witnessed a surge in
expeditions to both ichthyologically familiar and virgin
waters in southeastern Venezuela, Guyana, Suriname, and
French Guiana. Included are surveys of the Iwokrama
Forest in west-central Guyana (Watkins et al. 2005),
retracing Carl Eigenmann’s 1908 collecting route up the
Essequibo to the Potaro River above Kaiteur Falls
(Hardman et al. 2002), and rapid assessments targeting
species-rich waters such as the upper Essequibo Basin,
Guyana (Lasso et al. 2008), Coppename Basin, Suriname
(Berrenstein 2005, Alonso & Berrenstein 2006, and
references therein), and Venezuelan states of Amazonas
(Lasso et al. 2006, and references therein), and Bolıvar
(Machado-Allison et al. 2003). Systematic fish inventories
of French Guiana began over 50 years ago (see references
in Vari & Ferraris, this volume), and have been recently
expanded by French and Swiss ichthyologists to include
ecological (e.g., Lord et al. 2007) and molecular data, the
latter to investigate the origins of the Guianas’ highly
diversified fish fauna (Cardoso & Montoya-Burgos 2009).
Explorations of remote Shield regions in search of
undescribed catfishes (Sabaj Perez et al. 2009) have
assembled a parade of new taxa led by the sucker-mouth
armored siluriforms in the family Loricariidae. Fifteen
new loricariid species from Guyana, Suriname, and
Amazonas, Venezuela, have been described in the last
five years (e.g., Werneke et al. 2005, Armbruster et al.
2007, de Chambrier & Montoya-Burgos 2008, Lujan et al.
2009) with many more discoveries awaiting description.
This impressive amount of fieldwork has significantly
advanced our taxonomic understanding of fishes in the
Guianas; nevertheless, much must still be accomplished.
Expeditions to remote, previously unsampled waters,
particularly headwater systems above waterfalls or large
cataracts, routinely yield new and sometimes enigmatic
ichthyofaunas (Taphorn et al. 2008; Lujan, pers. comm.;
pers. obs.). More comprehensive collecting efforts (e.g.,
night sampling) in relatively well-sampled waters have
uncovered new species that escaped prior efforts (e.g.,
Armbruster et al. 2000; pers. obs.). Fieldwork aside,
there exists in museums a wealth of specimens of
Guianas fishes that require critical evaluation. The rich
and complex diversity of fishes in the Guianas, and their
systematic placement in the greater context of the
Neotropical fauna, will remain a lodestone for ichthy-
ological studies in decades to come.
Scope
The plates present 130 individuals representing 127
species of 46 families. Fishes were collected in Guyana
(53 species), Suriname (36) and Amazonas State,
Venezuela (38) from 1985 to 2008. Most of the species
occur on or immediately peripheral to the Guiana
Shield, with a few species restricted to lowland, coastal
habitats in fresh and/or estuarine waters (i.e., Rhino-
sardinia amazonica, Sciades parkeri, Tomeurus gracilis,
Anableps anableps, Polycentrus schomburgkii).
Fishes were imaged live or shortly after death (89
species), or from specimens purchased at market (2),preserved in formalin (2), or stored in alcohol (34).
Each image is identified in the plate description by
taxon, condition of specimen at time of photo, museum
and catalog number, size and sex (if so determined),
current status of voucher if other than preserved whole
in alcohol, and complete locality data. Depositories are
The Academy of Natural Sciences, Philadelphia
(ANSP), Auburn University Natural History Museum(AUM), Field Museum of Natural History (FMNH),
Illinois Natural History Survey (INHS), Museo de
Ciencias Naturales de la UNELLEZ, Guanare
(MCNG), National Zoological Collection of Suriname
(NZCS), and University of Guyana, Center for the
Study of Biological Diversity (UG/CSBD). Photos are
by author unless credited otherwise. Abbreviations in
the text are: LEA – length to end of anal fin; SL –standard length; and TL – total length. Scale bars are
presented only for those species in which that indicator
was included in the original photograph.
Fish Photography
There is a variety of techniques for capturing high-
quality color images of fishes, all of which have been
vastly simplified and in many ways improved by the
advent of digital technology. Most of the images
presented here are of live (or recently so) and alcohol
preserved specimens immersed in water in a glass
phototank. Materials and methods are largely the same
whether taken streamside of live specimens (Figs. 3, 4) orin the lab of preserved specimens (Fig. 5), except for the
light source: ambient sunlight in the field vs. incandescent
light in-doors. Other photographers have used electronic
flashes (e.g., Jenkins & Burkhead 1994:129, Planquette et
al. 1996:17) to produce stunning photos of live fishes in
phototanks. I have not tried such techniques, but consider
a cooperative sun to be equally effective and in some ways
less burdensome. In any event, phototank-immersionremains the gold standard for ex-situ fish photography.
Phototank-immersion Method
This method involves three stages: equipment set up,specimen preparation, and image capture and editing.
The techniques described below follow a minimalist
approach with some advice limited to the specific
cameras and conditions involved. For a more sophis-
ticated system and additional tips on fish photography
see Jenkins & Burkhead (1994:127–130).
Equipment set up.—The phototank is made of
ordinary plate glass bonded together with clear silicone
adhesive. Outside dimensions (in inches) of the tanks
used for the photos in this section are: 13.5 length 3
10.25 height 3 2.75 width (field and lab) and 15.75
length 3 12.25 height 3 3.5 width (lab only). Both are
made from one-quarter inch thick glass, except one-
eighth inch glass is used for the front plate of smaller
tank. These dimensions are well suited for lateral and
often dorsal/ventral views of small to medium-sized
fishes up to about 300 mm total length and 63 mm
width for smaller tank, and 370 mm total length and 75
mm width for larger. Two important factors compro-
mise field utility of larger phototanks: the volume of
water necessary to fill it and size of carrying case (see
below). Each tank requires a separate glass plate to
immobilize the subject. The free plate can be one-
eighth (smaller tank) or one-quarter (larger) inch thick
and is slightly shorter and deeper than the inside
dimensions of the tanks (e.g., 13 3 10.25 and 15 3 12
inches for smaller and larger tank, respectively).
Having smooth edges of all plates is recommended.
The tank should be filled with clear bottled or
filtered/deionized tap water to minimize formation of
air bubbles on specimen and glass. Stream or lake
water is unsuitable because it lacks the desired clarity
and suspended debris is a significant distraction in an
otherwise good photo. Any water will accumulate
debris over an extended photo session, and an ample
supply of clean photo water must accompany long
forays to remote locations.
In the lab the phototank is stationed between two
pairs of incandescent bulbs positioned to the side and
slightly above the top of the tank (Fig. 5). Polarizing
filters are useful for reducing glare or overexposed hot
spots on the specimen, particularly on the snout. When
using sunlight, the tank is oriented to maximize the
even distribution of light and minimize glare and
shadows on the subject.
Selection of a camera is important, but the rapid
pace of digital technology soon outdistances specific
recommendations on make or model. By current
standards a digital camera with a good optical zoom
(6X and higher) that records images at or above
resolutions of 12 Megapixels (MP) is generally a safe
Figure 3. Author using phototank-immersion method to photograph fish streamside in Mongolia. Photo by C. Sabaj Perez.
54 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
choice. Most of the photos herein were taken with a
Nikon Coolpix E8700 (8 MP); others with this model’s
predecessors, the E4500 (4 MP) and older E995 (3.1
MP). The most recent photos, all of alcohol preserved
specimens, were taken in the lab with a Nikon D90 D-
SLR (12.3 MP) fitted with a 60 mm f/2.8G micro lens.
Images taken with the E8700 contain a high level of
sharp detail that is slightly exceeded by the D90 (or
other cameras offering greater MP), particularly for
small specimens. The differences, however, are only
visible at high magnification or extremely large print
sizes. The greatest advantage of the D-SLR design and
micro lens is the enhanced ability to reliably focus on
very small specimens. Any camera and lens should be
thoroughly vetted by comparing published reviews
(many available on-line), and then personally tested
with the phototank-immersion method. A few digital
cameras apparently have difficulties rendering a sharp
specimen image through glass and water.
Additional essentials for basic set up are a tripod
(mini-tripods are handy in the field; Figs. 3–5), 4-ply
mat board in several background colors (e.g., flat
black, dull light blue) and 3/16th inch foam board with
flat black surface for camera blind (sizes of all boards
ideally fitted to carrying case), glass cleaner, and paper
towels or lint-free cloth, both long and small forceps,
large metal binder clips, 12-inch plastic metal rulers,
stiff wire, an assortment of needles and insect pins,
calipers, a system for tagging individual specimens
(e.g., dymo-tags in pre-punched number series tied to
strong twine), extra camera batteries and charger,
memory cards and reader, and laptop computer for
image storage. These essentials are best stored with the
phototank in a crushproof and watertight carrying
case. The smaller tank is ideal for field use as it requires
less water and allows for co-storage of accessories and
laptop in a small case suitable for carry-on luggage (see
Fig. 3). Cameras are better stored separately to
facilitate other uses and avoid residual moisture in
the phototank.
Specimen preparation.—The overarching strategy
when photographing fishes for identification purposes
is to maximize the content and accuracy of information
in the image. This aim determines which among
multiple specimens is photographed, how it is illumi-
nated and arranged for display, and which color
background is used. Most striking are photographs of
the most impressive specimens (i.e., in peak coloration
and with fins and scales intact), but even the image of
an impressive fish may be rendered less informative if
the photograph is poorly composed.
Once a live or alcohol specimen is selected it is
carefully inspected and cleaned of foreign debris.
Mucous-laden skin and fins often attract distracting grit
or other suspended particles, and cheesecloth fibers may
adhere to preserved specimens. An anesthetized fish (e.g.,
with a few drops of clove oil) is quickly euthanized in a
container of strong (30–50%) formalin. This often causes
the body to straighten and fins to become completely
erect. Otherwise the anesthetized specimen may be
Figure 4. Author photographing fish streamside in Guyana. Photo by J.W. Armbruster.
NUMBER 17 55
removed to a tray of shallow formalin wherein small
forceps are carefully used to hold the fins erect without
damaging them. The most important consideration
when photographing live specimens is time; bright colors
and iridescences are soon lost in formalin. Fatty skin, as
in pseudopimelodid catfishes, also becomes opaque in
formalin, obscuring any underlying color.
Once the specimen is flat with fins erect, it is care-
fully wedged between the front plate of the phototank
and free plate of glass, the latter set at an angle and
braced against metal binder clips either attached to the
sides of the tank or loosely set between the free plate
and back of tank (Fig. 3). Positioning laterally com-
pressed fishes in this manner is easy. Dorsoventrally
depressed specimens, particularly those with pectoral
spines, require more attention to achieve a vertical
lateral view. Maintaining pectoral spines folded against
the body as one wedges the specimen between the two
glass plates requires practice and patience. Long
forceps, a metal ruler and stiff wire are useful tools
for fine-tuning a specimen’s posture, arranging long
delicate features such as barbels, and dislodging air
bubbles that form on the fish. Information content of a
fish photo is diminished when the specimen is tilted or
otherwise poorly positioned.
Preserved specimens offer fewer options for achiev-
ing an ideal photo-friendly posture. Laterally contorted
specimens often can be made to appear more linear
when tightly wedged between the two plates of glass.
Issues that are more difficult arise with partial or
complete folding of fins. In some cases insect pins
(carefully inserted in the body opposite the side to be
imaged) may be used to prop up the anterior most
portions of fins. This technique, however, may cause
small tears in the fin membranes.
Next is selection of an appropriate background.
Many specimens, particularly dark ones with opaque
fins, often render best with more dramatic effect
against flat black backgrounds. This may pose a
serious drawback for specimens with relatively trans-
parent fins. Black pigment in fin membranes or along
distal fin margins disappears against dark back-
grounds. In such cases, a light blue background
provides better contrast and will highlight dark
pigmentation in fins. Conversely, transparent fins
lacking pigmentation and with clear margins, particu-
larly in live specimens, are often lost against light
backgrounds. This can be alleviated to a certain degree
by adjusting the tank relative to light source to achieve
a small measure of direct side or back lighting. While it
is true that graphics editing software (e.g., Vertus Fluid
Mask) can virtually affect any color background,
specimens may not appear natural if the new back-
ground deviates sharply from the original (i.e., black to
Figure 5. Kyle Luckenbill photographing small alcohol-preserved specimen (above ruler) while holding polarizing filter in lab. Photo
by author.
56 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
white and vice versa). Choice of background color
often involves trade-offs, and is ultimately a reflection
of personal taste determined via trial and error.
The final step is placement of a scale bar. This is
accomplished by cutting out a 10+ mm portion of a
plastic ruler, dipping it in water and adhering it to the
outside front of the phototank beneath the specimen
and within the photographic field.
Image capture and editing.—The camera is mounted
on a tripod, as most exposures are too long to permit
hand-held use, and positioned behind a black foam
board with central circular aperture fitted to lens. The
blind prevents the phototank glass from reflecting the
images of camera and photographer. Whether hori-
zontal or angled the specimen should occupy about
90% of the length of the digital image recorded. To
preserve detail in extremely long and slender fishes
(e.g., belonids; Pl. 14, Fig. E) the specimen is imaged in
two aligned and overlapping parts (anterior and
posterior halves) that are digitally combined. The
shutter is placed on a timer delay and white balance
set appropriately (e.g., sunlight vs. incandescent).
Digital photography frees one from limits imposed
by the amount of available film and developing costs.
In the field, particularly while the sun is dodging
clouds, it is advisable to take multiple photos for
each of several combinations of exposures and aper-
tures (f-stops). Full sunlight often highlights fine
structures (e.g., odontodes in loricariids), but at the
same time may wash out bright colors or result in
overexposed hot spots on the snout or dorsum. The
phototank should be carefully oriented with respect
to the light source, and extra mat boards used to
shadow harsh sunlight and maintain vibrant colors
(Fig. 3).
For the Coolpix E8700 in manual mode, the shutter
speed is set such that the target aperture (i.e., lower
f-stops) lies between f-stops 5 and 7; larger apertures
reduce depth of field, and smaller apertures tend to
reduce resolution. The Nikon D90 D-SLR better
accommodates smaller apertures (f-stop fixed at 16
with ISO set to 200), and the shutter speed is manually
adjusted for the best exposure. Autofocus generally
works fine as long as the active area of focus includes
important features on the fish, not the scale bar or
background. Digital cameras typically have a setting
whereby the user determines the active area of
autofocus. Depending on specimen size, the camera
may need to be manually set to macro mode, and some
cameras (e.g., Nikon CoolPix) also require one to
slightly zoom in on subject for sharp autofocus. Nikon
images presented here are of Fine quality (recorded as
JPEGs with compression ratio of roughly 1:4) and
maximum size (3264 3 2448 and 4288 3 2848 pixels for
E8700 and D90, respectively). Higher quality settings
record either uncompressed TIFF or RAW (NEF)
images, the latter requiring extra software and com-
puting time for conversion to TIFF files (Nikon D90
allows one to record NEF and Fine JPEG images
concurrently). TIFF and RAW files retain the full
quality of the image and the latter maximizes allowable
post exposure processing, whereas JPEGs are com-
pressed often with some visual quality permanently lost
in the process (the loss, however, is barely perceptible).
Larger image files (NEF, RAW, TIFF) do offer
slightly higher resolution, but the improvement is often
negligible, except at high magnification. For any
camera, there is no substitute for testing a variety of
settings and image qualities to optimize the desired
effect and protocol.
While photographing a specimen it is difficult to
know which image will optimize the desired effect; so,
it is best to have ample images from which to choose.
The number of images I generally take is directly
proportional to the impressive and unique nature of
the specimen added to the amount of time expended to
pose it properly in the phototank. It is easy to
accumulate many photos of numerous species, thus it
is critical to have a system for later identification and
management of images. Failure to do so guarantees
extra time and often frustration when attempting to
match images to specimens long after capture. The best
field solution is to take a final photo of the specimen
together with a uniquely numbered tag that is then
secured to the fish. In the case of museum specimens,
the jar label is photographed immediately after imaging
the fish. A photo-log is useful for recording the
standard length of the specimen. Such practices greatly
facilitate subsequent annotation of images with catalog
and measurement data. A new and much welcomed
trend in digital cameras is a built in or accessory global
positioning system (GPS) receiver that records and
embeds latitude, longitude, altitude and universal time
as image metadata.
The final step is image editing, all of which was
performed on the photos in this section using Adobe
Photoshop. This program offers a seemingly endless
myriad of simple to advanced tools for graphics
manipulation. Only a few of the more basic tools and
techniques are mentioned briefly here.
Once an image is selected the background (original)
layer is immediately duplicated and subsequent edits
are made to the duplicate layer. A third blank layer is
added to mask the specimen with a uniform back-
ground. Masking color (e.g., solid black or white, or a
color shade taken from the original background using
the eyedropper tool) is first added as a rough outline
using a large diameter pencil tool, and then completed
with a fine-tipped brush (1–10 pixels) under magnifi-
cation (e.g., $300%) to carefully trace the specimen’s
precise contours. The magic wand and/or magnetic
lasso are more expedient, yet less precise, tools for
masking the specimen with a uniform background.
Next, the duplicate layer is automatically and manually
NUMBER 17 57
adjusted for levels (tonal range and color balance),
brightness/contrast, and hue/saturation. The auto
options often render extreme values that are manually
faded to desired opacity before additional manual fine-
tuning. The cloning stamp tool is useful for removing
small bubbles or debris on the specimen, while the
dodge and burn tools help lighten or darken localized
regions (brush size/shape and exposure/opacity of such
tools are manually adjusted). Under the Filter menu
‘Sharpen.Unsharp Mask’ can sharpen an image with
soft focus, and ‘Noise.Despeckle’ removes graininess,
particularly for images scanned from 35 mm slides.
One final trick is to render a solid scale bar by: 1)
rotating the entire image so the ruler piece in the
photograph is horizontal, 2) using the rectangular
marquee tool to select and copy a 5 or 10 mm long
portion, 3) rotating the entire image back to its final
intended position, 4) pasting the copied selection,
thereby creating a new layer, and 5) adjusting the
brightness/contrast of this layer to extreme values to
render a black or white bar that is then labeled
accordingly with the text tool.
The final edited version of the specimen image (i.e.,
duplicate layer) can be quickly compared to the
original by using the layers window and clicking the
‘eye’ icon to hide or display the corresponding layer.
Creation of additional layers requires the new image to
be saved as an uncompressed TIFF file that is suitable
for archiving and any additional post processing. A
copy of the final edited image is flattened to a single
layer and resaved as a TIFF for print publication (with
Image.Mode set to Grayscale, RGB, or CMYK color
based upon printer specifications) and separately as a
JPEG (with Mode set to 8 bits/channel) for use in
presentations or easy transmission and accession via
the Internet.
In sum, a high-quality fish photo is the product
of preparation, practice, and patience all committed
with keen attention to detail. Additional factors in
the field are perseverance under suboptimal conditions
and a bit of luck with respect to weather and finding
the ideal specimen. The amount and quality of the
images presented herein had more to do with will than
skill.
Acknowledgments
Numerous colleagues contributed valuable assis-
tance both before and after the shutter click. For
assistance in the field I wish to thank in particular:
Mariangeles Arce, Jonathan Armbruster, Michael
Hardman, Oscar J. Leon-Mata, Nathan Lujan, John
Lundberg, Jan Mol, Matthew Thomas, David Wer-
neke, and Philip Willink. For loans of specimens I
thank Michael Retzer, and for help tracking down
specimens and taking measurements I thank Osvaldo
Oyakawa and especially Nathan Lujan. For help
cleaning backgrounds I thank Kyle Luckenbill and
Rebecca Meyer. Special thanks to Pierre-Yves le Bail,
Raphael Covain and Juan Montoya Burgos for
comments on the introduction, and to Jonathan
Armbruster, Hernan Lopez-Fernandez, Nathan Lujan,
Donald Stewart, and Donald Taphorn for graciously
contributing their images to this atlas. Finally I thank
my ichthyological friends for years of help identifying
many of the fishes presented here. Support for this
project was received in part from the All Catfish
Species Inventory (NSF DEB-0315963).
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assessment of the aquatic ecosystems of the Coppename River
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———, M. H. Sabaj, M. Hardman, L. M. Page, & J. H. Knouft.
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Baryancistrus with blue sheen from the upper Orinoco
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Marcano, P. Petry, B. Sidlauskas, & T. Jones. 2003.
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G. Lundberg, & L. M. Page (eds.). 2009. All Catfish Species
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NUMBER 17 59
APPENDIX: PLATES
Plate 1
Potamotrygonidae
A. Paratrygon aiereba (live). AUM 43646 (154 mm maximum disk width). Venezuela,
Amazonas, Rıo Negro (Amazonas drainage), left bank sandy beach and small adjacent
backwater 7.2 km NW of San Carlos de Rio Negro, 01u589110N, 067u069100W, 19 Mar
2005, M. Sabaj, D. Werneke et al.
B. Potamotrygon orbignyi (live). AUM 43201 (171 mm maximum disk width). Venezuela,Amazonas, Rıo Orinoco ca. 60 km E of San Fernando de Atabapo, 03u589260N,
067u099460W, 3 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.
C. Potamotrygon marinae (live). ANSP 187098 (400 mm maximum disk width). Suriname,
Sipaliwini, Lawa River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan
(airstrip), 03u199310N, 054u039480W, 18 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P.
Willink, & K. Wan.
D. Potamotrygon schroederi (live). AUM 44507 (423 mm maximum disk width). Venezuela,
Amazonas, Rıo Orinoco, island W of Puerto Venado, 4.5 km S of Samariapo, 56.5 km
SW of Puerto Ayacucho, 05u129250N, 067u489320W, 28 Feb 2005, M. Sabaj, N. Lujan, D.
Werneke et al.
62 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 63
Plate 2
Arapaimidae
A. Arapaima sp. (live). UG/CSBD uncataloged (174 cm SL, male, skeleton). Guyana, Grass
Pond, Rewa River basin (Rupununi drainage), near Rewa village, Aug 2006, D. Stewart et
al. Photo by D. J. Stewart.
Osteoglossidae
B. Osteoglossum bicirrhosum (live). No voucher (ca. 500 mm TL). Guyana, Crane Pond,
southwestern part of Rupununi basin near Karanambu Ranch, 2007, D. Stewart et al.
Photo by D. J. Stewart.
64 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 65
Plate 3
Clupeidae
A. Rhinosardinia amazonica (alcohol). INHS 49009 (33.7 mm SL). Guyana, East Demerara,
Demerara River (Atlantic drainage), Land of Canaan, 11.8 mi S-SW of Georgetown at
bearing 213u, 06u389390N, 058u119460W, 14 Oct 1998, M. Sabaj, J. Armbruster, M.
Hardman et al.
Pristigasteridae
B. Pellona castelnaeana (live). MCNG 51957. Venezuela, Amazonas, Rıo Casiquiare at
mouth of Cano Caripo, 37 km W-SW of La Esmeralda, 03u069500N, 065u529380W, 5 Mar
2005, M. Sabaj, N. Lujan, D. Werneke et al.
Engraulidae
C. Amazonsprattus scintilla (live). ANSP 181134. Guyana, Pirara River (Ireng-Takutu-
Branco drainage), 3.5 km N-NW of Pirara, 03u389550N, 059u419200W, 2 Nov 2002, M.
Sabaj, J. Armbruster, M. Thomas et al.
D. Anchovia surinamensis (alcohol). ANSP 189252 (69.7 mm SL). Guyana, Rupununi River
(Essequibo drainage), at Massara’s Landing, 1.1 km NE village of Massara, 03u539410N,
059u179370W, 26 Oct 2002. M. Sabaj, J. Armbruster, M. Thomas et al.
E. Lycengraulis batesii (alcohol). ANSP 189251 (52.4 mm SL). Guyana, Cuyuni-Mazaruni,
Mazaruni River (Essequibo drainage) long partially exposed sandy shoal between two
islands in main channel, 6.9 km SW of Bartica, 06u229470N, 058u409320W, 12 Nov 2002,
M. Sabaj, J. Armbruster, M. Thomas et al.
F. Anchoviella sp. (alcohol). ANSP 189234 (29.5 mm SL). Suriname, Sipalawini, Lawa River
(Marowini drainage), small sand beach below cataract upstream from base camp, ca. 9 kmS-SW of Anapaike. 03u199120N, 054u039410W, 19–23 Apr 2007, J. Lundberg, J. Mol, M.
Sabaj, P. Willink, & K. Wan.
Parodontidae
G. Apareiodon agmatos (alcohol). ROM 83755 (52.6 mm SL). Guyana, Mazaruni River
(Essequibo drainage), sandy beach and embayment on right bank, upstream from villageof Jawalla, 05u41935.40N, 060u28911.80 W, 18 Apr 2008, H. Lopez-Fernandez, D.
Taphorn, E. Liverpool, K. Kramer, & C. Thierens. Photo by D. C. Taphorn Baechle & H.
Lopez-Fernandez.
H. Apareiodon orinocensis (live). ANSP 185045. Venezuela, Amazonas, Rıo Orinoco at
Puerto Venado, 4.3 km S of Samariapo, 56.4 km SW of Puerto Ayacucho, 05u129380N,
067u489180W, 26 Feb 2005, M. Sabaj, N. Lujan, D. Werneke et al.
I. Parodon guyanensis (live). ANSP 189204 (ca. 70 mm SL). Suriname, Sipaliwini, Lawa River
(Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip), 03u199310N,
054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.
J. Parodon guyanensis (alcohol). ANSP 189204 (73 mm SL). Same locality as I.
66 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 67
Plate 4
Curimatidae
A. Curimatopsis crypticus (live). ANSP 189091 (32.4 mm SL, female). Suriname, Para,
Coropinae Creek (Suriname drainage), vicinity of Republiek, 05u299570N, 055u129520W,
28 Apr 2007, M. Sabaj & P. Willink.
B. Curimatopsis crypticus (live). ANSP 189091 (28.8 mm SL, male). Same locality as A.
Prochilodontidae
C. Prochilodus rubrotaeniatus (alcohol). ANSP 175495 (160 mm SL). Guyana, Essequibo
River, sandbar ca. 800 m downstream of Essequibo (Maipuri) campsite, 04u459430N,
058u459520W, 29 Jan 1997, W. Saul et al.
D. Semaprochilodus varii (live). ANSP 187435 (210 mm SL). Suriname, Sipaliwini, LawaRiver (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
Crenuchidae
E. Leptocharacidium omospilus (live). ANSP 189272 (60.5 mm SL). Venezuela, Amazonas, 1km upstream from mouth of left bank tributary of Rıo Siapa, mouth below Salto Oso and
above Salto Sardinas on Rıo Siapa, 01u269240N, 065u409010W, 14 Mar 2005, M. Sabaj, N.
Lujan, M. Arce, & T. Wesley.
F. Poecilocharax bovalii (alcohol). INHS 49600 (28.6 mm SL). Guyana, first N bank creek
tributary of Potaro River (Essequibo drainage) downstream from Waratuk Cataract, 27
Oct 1998, L. Page, M. Sabaj, J. Armbruster et al.
Hemiodontidae
G. Argonectes longiceps (alcohol). ANSP 189151 (184 mm SL). Same locality data as D.
H. Bivibranchia bimaculata (live). ANSP 189149 (111 mm SL). Same locality data as D.
Gasteropelecidae
I. Carnegiella strigata (alcohol). INHS 49173 (26 mm SL). Guyana, Mazaruni–Potaro,
‘‘Himarakus’’ Creek, tributary of Essequibo River (Atlantic drainage) at Rockstone,
05u599080N, 058u339030W, 19 Oct 1998, M. Sabaj, J. Armbruster, M. Hardman.
J. Gasteropelecus sternicla (alcohol). ANSP 189193 (46.6 mm SL). Same locality as A.
68 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 69
Plate 5
Anostomidae
A. Anostomus brevior (live). FMNH ex ANSP 189141 (51.6 mm SL). Suriname, Sipaliwini,
Lawa River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
B. Anostomus anostomus (live). ANSP 180172 (72.5 mm SL). Guyana, Essequibo River(Atlantic drainage) at Yukanopito Falls, 44.5 km SW of mouth of Kuyuwini River,
01u549530N, 058u319140W, 9 Nov 2003, M. H. Sabaj, J. Armbruster, N. Lujan et al.
C. Synaptolaemus cingulatus (live). AUM 43269 (78.8 mm SL). Venezuela, Amazonas, Rıo
Orinoco, 147 km SE of San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar
2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.
D. Leporellus vittatus (formalin). ANSP 189270 (photo voucher). Guyana, Rupununi River(Essequibo drainage), road crossing 5.9 km W-SW of village of Sand Creek, 02u579000N,
059u349100W, 4 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
E. Leporinus fasciatus (live). ANSP 189158 (81.7 mm SL). Same locality as A.
F. Leporinus maculatus (live). FMNH ex ANSP 189041 (123.4 mm SL). Same locality as A.
G. Hypomasticus megalepis (live). AUM 37999 (68.3 mm SL). Guyana, Essequibo River atKassi-Attae Rapids, 5.5 km SE of mouth of Kuyuwini River, 02u139360N, 058u179380W, 8
Nov 2003, M. Sabaj, J. Armbruster, M. Hardman et al.
H. Leporinus ortomaculatus (live). AUM 43262 (72.9 mm SL). Same locality as C.
I. Leporinus lebaili (live). FMNH ex ANSP 189043 (56.7 mm SL). Same locality as A.
J. Caenotropus maculosus (live). ANSP 189147 (70 mm SL). Suriname, Sipalawini, LitanieRiver, side channel behind Theo’s bakery, just upstream of confluence with Marowini
River and village of Konya Kondre, 03u179240N, 054u049380W, 23 Apr 2007, J. Lundberg,
J. Mol, M. Sabaj, P. Willink, & K. Wan.
70 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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Plate 6
Characidae
A. Brittanichthys myersi (formalin). INHS 49053 (22.4 mm SL). Guyana, East Demerara,
Maduni River and Conservancy Canal (Mahaica-Atlantic drainage), Maduni Stop-off,
22.3 mi S of Georgetown bearing 176u, 06u309010N, 58u029140W, 15 Oct 1998, M. Sabaj, J.
Armbruster, M. Hardman et al.
B. Bryconops melanurus (live). ANSP 189268 (75.5 mm SL). Suriname, Sipalawini, LitanieRiver at mouth and confluence with Marowini River, just upstream from settlement of
Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P.
Willink, J. Mol et al.
C. Chalceus epakros (live). AUM 43073 (75 mm SL). Venezuela, Amazonas, Rıo Orinoco,
inlet between 2 islands in channel, 84.5 km N of San Fernando de Atabapo, 9 km S of
Monduapo, 1 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.
D. Exodon paradoxus (live). AUM 36843 (61 mm SL). Guyana, Rupununi River (Essequibo
drainage), sand beach and inlet at Karanambo Ranch, 03u459000N, 059u189300W, 30 Oct
2002, M. Sabaj, J. Armbruster, M. Thomas et al.
E. Myleus rubripinnis (live). ANSP 189267 (34 mm SL). Guyana, Suriname, Sipaliwini, Lawa
River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
F. Myleus schomburgkii (live). ANSP 180812 (160 mm SL). Venezuela, Amazonas, Rıo
Orinoco, bedrock outcrop, 52.9 km SE of San Antonio, 102 km W of La Esmerelda,
03u069010N, 066u279460W, 4 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.
G. Myloplus cf. planquettei (live). ANSP 179808 (425 mm SL). Guyana, Essequibo River
(Atlantic drainage) at Yukanopito Falls, 44.5 km SW of mouth of Kuyuwini River,
01u549530N, 058u319140W, 9 Nov 2003, M. H. Sabaj, J. Armbruster, N. Lujan et al.
H. Serrasalmus rhombeus (live). ANSP 180287 (ca. 260 mm SL). Venezuela, Amazonas, Rıo
Ventuari (Orinoco drainage), large rock outcrop, 97 km NE of Macuruco, 163 km E-NE
of San Fernando de Atabapo, 7 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.
72 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 73
Plate 7
Acestrorhynchidae
A. Acestrorhynchus falcatus (live). INHS 48983 (153.9 mm SL). Guyana, East Demerara,
Madewini River (Demerara drainage), 21.5 mi S-SW of Georgetown, bearing 207u, at
Linden highway bridge, 06u 309 05.00 N, 58u 129 44.90 W, 14 Oct 1998, M. Sabaj, J.
Armbruster, M. Hardman et al.
B. Acestrorhynchus microlepis (live). AUM 36753 (123 mm SL). Guyana, Circle West Creek,tributary of Pirara River (Ireng-Takutu drainage), 26.6 km SW of Karanambo Ranch,
03u399140N, 059u319430W, 30 Oct 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
Cynodontidae
C. Cynodon meionactis (alcohol). ANSP 189129 (190 mm SL). Suriname, Sipalawini, Litanie
River at mouth and confluence with Marowini River, just upstream from settlement ofKonya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P.
Willink, J. Mol et al.
D. Hydrolycus armatus (live). MCNG 51983 (ca. 400 mm SL). Venezuela, Amazonas, Rio
Orinoco (Atlantic drainage), Pasaganado, 38 km N of San Fernando de Atabapo,
04u239040N, 067u469280W, 1 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.
Lebiasinidae
E. Copella compta (live). AUM 41327 (37 mm SL). Venezuela, Amazonas, Cano Carmen,
tributary of Rıo Orinoco, 1.5 km S-SE of Manaka, 70 km E of San Fernando de Atabapo,
4 Apr 2004, N. Lujan & D. Werneke.
F. Copella cf. arnoldi (live). ANSP 189192 (30.4 mm SL). Suriname, Para, Coropinae Creek
(Suriname drainage), vicinity of Republiek, 05u299570N, 055u129520W, 28 Apr 2007, M.Sabaj & P. Willink.
Erythrinidae
G. Hoplias aimara (alcohol). ANSP 176723 (172 mm SL). Guyana, Tumble Down Creek,
tributary of Siparuni River (Essequibo drainage), 04u489390N, 058u519110W, 8 Dec 1997,
G. Watkins et al.
H. Hoplias lacerdae group (live). AUM 44674 (152 mm SL). Guyana, Pirara River (Ireng-
Takutu drainage), at Pirara Ranch, 03u379310, 059u409360, 26 Nov 2005, N. Lujan, D.
Taphorn et al. Photo by N. K. Lujan.
I. Erythrinus erythrinus (alcohol). ANSP 175537 (121 mm SL). Guyana, Culvert creek
(Essequibo drainage) crossing Kurupukari-Surama River road, 04u199570N, 058u519130W,
5 Feb 1997, W. Saul et al.
Ctenoluciidae
J. Boulengerella cuvieri (live). AUM 40987 (560 mm SL). Venezuela, Amazonas, Rıo Ventuari
(Orinoco drainage), large rock outcrop, 97 km NE of Macuruco, 163 km E-NE of San Fernando
de Atabapo, 04u259100N, 066u179080W, 7 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.
74 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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Plate 8
Cetopsidae
A. Cetopsis cf. montana (live). ANSP 178782 (78.3 mm SL). Guyana, Ireng River (Takutu-
Branco-Negro drainage), 6.9 km W-SW of village of Karasabai, 04u019100N,
059u369060W, 1 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
B. Helogenes marmoratus (live). AUM 27960 (56 mm SL). Guyana, West Demerara,
Catabuly Creek (Demerara-Atlantic drainage), at Wismar, 1.87 mi. N-NW of Lindenbearing 335u, 06u01937.20N, 58u19925.30W, 18 Oct 1998, L. Page, M. Sabaj, J. Armbruster
et al. Photo by J. W. Armbruster.
Aspredinidae
C. Ernstichthys sp. (alcohol). ANSP 180002 (26.4 mm SL). Same locality as A.
D. Bunocephalus verrucosus (alcohol). ANSP 180015 (36.6 mm SL). Guyana, Hassar Pond
and outlet (Rupununi drainage), 5.4 km S-SE of Massara, 03u509400N, 059u179090W, 27
Oct 2002, J. Armbruster, D. Werneke, C. Allison et al.
Ariidae
E. Sciades parkeri (live). ANSP 178741 (445 mm SL). Guyana, purchased at Georgetown fishmarket, 8–15 Nov 2002, M. Sabaj.
Auchenipteridae
F. Ageneiosus inermis (live). FMNH ex ANSP 187115 (ca. 200 mm SL). Suriname,
Sipalawini, Litanie River at mouth and confluence with Marowini River, just upstream
from settlement of Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg,M. Sabaj, P. Willink, J. Mol et al.
G. Auchenipterichthys punctatus (live). AUM 43416 (131 mm SL). Venezuela, Amazonas, Rıo
Casiquiare, bedrock riffle and outcrop, 74.6 km NE of San Carlos de Rio Negro,
02u219460N, 066u339530W, 9 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.
H. Trachycorystes trachycorystes (live). AUM 35933 (111 mm SL). Guyana, unnamed stream
(Rupununi drainage) at crossing on road between Massara and Karanambo, 10.3 km NWof Karanambo Ranch, 03u489270N, 059u239060W, 28 Oct 2002, M. Sabaj, J. Armbruster,
M. Thomas et al.
I. Gelanoglanis sp. (live). AUM 35908 or ANSP 178738 (male). Guyana, Ireng River
(Takutu-Branco drainage), 6.9 km WSW village of Karasabai, 04u019100N, 059u369060W, 1
Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
J. Glanidium leopardum (live). ANSP 189104 (48.5 mm SL). Same locality as F.
76 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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Plate 9
Doradidae
A. Acanthodoras cataphractus (alcohol). NZCS 1618–1619 (84.5 mm SL). Suriname, Para,
tributary of lower Suriname River at Republiek, 14 Nov 1989, P. Outboter et al.
B. Scorpiodoras heckelii (live). ANSP 182790 (82.5 mm SL, specimen with simple secondary
gas bladder). Venezuela, Amazonas, Rıo Orinoco at Puerto Samariapo, 05u159N,
067u489W, 25 Feb 2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.
C. Leptodoras copei (live). ANSP 182225 (85.8 mm SL). Venezuela, Amazonas, Rıo Ventuari
(Orinoco drainage), beach across river from Picua village, 34 km E-NE of Macuruco, 104
km E of San Fernando de Atabapo, 04u069550N, 066u459520W, 5 Apr 2004, M. Sabaj, N.
Lujan, D. Werneke et al.
D. Leptodoras linnelli (live). AUM 41038 (171 mm SL). Venezuela, Amazonas, Rıo Ventuari
(Orinoco drainage), village of Marueta at landing, 91 kmE-NE of Macuruco, 159 km E-NE of San Fernando de Atabapo, 04u189510N,
066u179320W, 6 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.
E. Centrodoras hasemani (live). ANSP 182227 (211 mm SL). Venezuela, Amazonas, Rıo
Casiquiare, bedrock outcrop 59.5 km SW of La Esmerelda, 02u499070N, 065u579190W, 8
Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.
F. Oxydoras niger (live). AUM 35508 (340 mm SL). Pirara River, tributary of Ireng River(Takutu-Branco drainage), beach at Pirara Ranch on road to Lethem, 03u379170N,
059u409290W, 2 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
G. Doras micropoeus (alcohol). ANSP 187110 (225 mm SL). Suriname, Sipaliwini, Lawa
River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
H. Hassar orestis (live). ANSP 180294 (161 mm SL). Venezuela, Amazonas, Rıo Ventuari
(Orinoco drainage), beach at village of Moriche, 116 km NE of Macuruco, 169 km NE of
San Fernando de Atabapo, 04u459N, 066u219130W, 7 Apr 2004, M. Sabaj, N. Lujan, D.
Werneke et al.
I. Opsodoras morei (live). ANSP 182836 (157 mm SL). Venezuela, Amazonas, Rıo Orinoco
near confluence with Rıo Atabapo, long narrow beach between channel and laguna, San
Fernando de Atabapo, 04u029480N, 067u429170W, 2 Apr 2004, M. Sabaj, N. Lujan, D.Werneke et al.
J. Rhinodoras armbrusteri (live). ANSP 179096 (66.1 mm SL). Guyana, Takutu River (Rio
Branco-Negro drainage), ca. 2.75 km W of Saint Ignatius, 03u219180N, 059u499510W, 5-6
Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
78 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 79
Plate 10
Pseudopimelodidae
A. Pseudopimelodus bufonius (live). ANSP 189098 (87.4 mm SL). Suriname, Sipalawini,
Litanie River at mouth and confluence with Marowini River, just upstream from
settlement of Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M.
Sabaj, P. Willink, J. Mol et al.
Heptapteridae
B. Mastiglanis sp. (live). FMNH ex ANSP 189106 (43.2 mm SL). Same locality as A.
C. Leptorhamdia sp. (live). AUM 43261 (51 mm SL). Venezuela, Amazonas, Rıo Orinoco,
147 km SE of San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar 2005, M.
Sabaj, N. Lujan, M. Arce, & T. Wesley.
D. Pimelodella geryi (live). ANSP 189109 (114 mm SL). Suriname, Sipaliwini, Lawa River
(Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip), 03u199310N,
054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.
Pimelodidae
E. Hypophthalmus marginatus (market). ANSP 187103 (330 mm SL). Suriname, Paramaribo,purchased at main fish market in Paramaribo, presumably from vicinity in Surinam River,
17 Apr 2007, J. Lundberg, M. Sabaj, & P. Willink.
F. Megalonema platycephalum (live). AUM 36018 (97.2 mm SL). Guyana, Rupununi River
(Essequibo drainage), sand beach and inlet at Karanambo Ranch, 03u459000N,
059u189300W, 30 Oct 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
G. Brachyplatystoma filamentosum (live). ANSP 187070 (540 mm SL). Venezuela, Amazonas,Rıo Casiquiare, bedrock outcrop 59.5 km SW of La Esmerelda, 02u499070N,
065u579190W, 8 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.
H. Pseudoplatystoma fasciatum (market). ANSP 187106 (ca. 640 mm SL, skeleton). Same locality
as E.
I. Pimelodus ornatus (live). ANSP 187113 (169 mm SL). Same locality as D.
J. Pinirampus pirinampu (live). AUM 37964. Guyana, Essequibo River, along beach 12.9 km
SE (upstream) of mouth of Kuyuwini River, 02u099430, 058u169350, 10 Nov 2003, M.
Sabaj, J. Armbruster, M. Hardman et al.
80 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 81
Plate 11
Callichthyidae
A. Callichthys callichthys (live). ANSP 179110 (68.2 mm SL). Guyana, Orokang River
(Mazaruni drainage), 1.2 km S of Chi Chi Falls airstrip, 21.1 km S-SW of Imbaimadai,
05u319310N, 060u139560W, 13 Nov 2002, J. Armbruster, M. Sabaj, M. Thomas et al.
B. Megalechis thoracata (alcohol). ANSP 179795 (62 mm SL). Guyana, Hassar Pond and
outlet (Rupununi drainage), 5.4 km S-SE of Massara, 03u509400N, 059u179090W, 27 Oct2002, J. Armbruster, D. Werneke, C. Allison et al.
Trichomycteridae
C. Ituglanis cf. metae (live). AUM 43074 (61 mm SL). Venezuela, Amazonas, Rıo Orinoco,
inlet between two islands in channel, 84.5 km N of San Fernando de Atabapo, 9 km S of
Monduapo, 04u479540N, 067u499160W, 1 Mar 2005, M. Sabaj, N. Lujan, D. Werneke etal.
D. Trichomycterus hasemani (alcohol, top 5 lateral view, bottom 5 dorsal view). ANSP
179154 (13.5 mm SL). Guyana, Rupununi River (Essequibo drainage), 3.7 km S-SE of
village of Massara, 03u519440N, 059u179040W, 27 Oct 2002, M. Sabaj, J. Armbruster, M.
Thomas et al.
E. Henonemus taxistigmus (alcohol). ANSP 179953 (90.6 mm SL). Guyana, Rupununi River(Essequibo drainage) at Kwatamang, 4 km SE of Annai, 03u559030N, 059u069010W, 25 Oct
2002, M. Sabaj, J. Armbruster, M. Thomas et al.
F. Trichomycterus guianensis (alcohol). ANSP 179109 (52.5 mm SL). Guyana, Orokang
River (Mazaruni drainage), at Chi Chi Falls airstrip, 20.1 km S-SW of Imbaimadai,
05u329060N, 060u139590W, 13 Nov 2002, J. Armbruster, M. Sabaj, M. Thomas et al.
G. Typhlobelus sp. (alcohol, dorsal view). AUM 35802 (22.3 mm SL). Guyana, Ireng River(Takutu-Branco-Negro drainage), 6.9 km W-SW of village of Karasabai, 04u019100N,
059u369060W, 1 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
H. Typhlobelus sp. (alcohol, left 5 dorsal view, right 5 ventral view of head). Same data as
G.
I. Pygidianops sp. (live). ANSP 179820 (23.1 mm SL). Guyana, Takutu River (Branco-Negro
drainage), 3.77 km S-SW of Lethem, 03u219180N, 059u499510W, 16 Nov 2003, M. Sabaj, J.Armbruster, M. Hardman et al.
J. Sarcoglanis simplex (live). ANSP 179212 (17 mm SL). Same locality as G.
82 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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Plate 12
Loricariidae
A. Leporacanthicus triactis (live). AUM 39243 (116 mm SL). Venezuela, Amazonas, Rıo
Ventuari (Orinoco drainage), 23 km NE of Macaruco, 94 km E of San Fernando de
Atabapo, 04u049500N, 066u519540W, 5 Apr 2004, N. Lujan, D. Werneke, M. Sabaj et al.
B. Leporacanthicus cf. galaxias (live). AUM 39226 (93 mm SL). Venezuela, Amazonas, Rıo
Orinoco at Cucue Amerindian village, opposite Trapichote village, 60 km E of SanFernando de Atabapo, 03u589260N, 067u099300W, 3 Apr 2004, N. Lujan, D. Werneke, M.
Sabaj et al.
C. Hemiancistrus medians (live). ANSP 187122 (156 mm SL). Suriname, Sipaliwini, Lawa
River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
D. Peckoltia sabaji (live). ANSP 185094 (109.7 mm SL). Guyana, Takutu River (Branco-
Negro drainage), 3.77 km S-SW of Lethem, 03u219180N, 059u499510W, 1 Nov 2003, M.
Sabaj, J. Armbruster, M. Hardman et al.
E. Peckoltia braueri (live). AUM 38882 (103 mm SL). Same locality as D.
F. Lithoxus stocki (live). ANSP 189131 (66.5 mm SL). Suriname, Sipalawini, Litanie River at
mouth and confluence with Marowini River, just upstream from settlement of KonyaKondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P. Willink, J.
Mol et al.
G. Pseudolithoxus dumus (live). AUM 43267 (87 mm SL). Venezuela, Amazonas, Rıo
Orinoco, 147 km SE of San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar
2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.
H. Pseudancistrus pectegenitor (alcohol). MCNG 54797 (241.6 mm SL). Venezuela,Amazonas, Rıo Casiquiare, bedrock in stream, 73 km NE of San Carlos de Rıo Negro,
02u219090N, 066u349310W, 9 Mar 2005, N. Lujan, M. Sabaj, D. Werneke et al.
I. Metaloricaria paucidens (live). ANSP 187325 (146.5 mm SL). Suriname, Sipaliwini, Lawa
River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
J. Rhadinoloricaria macromystax (live). ANSP 182349 (100 mm SL). Guyana, Ireng River
(Takutu-Branco-Negro drainage), 6.9 km W-SW of village of Karasabai, 04u019100N,
059u369060W, 1 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
84 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 85
Plate 13
Gymnotidae
A. Electrophorus electricus (live). MCNG 51982. Venezuela, Amazonas, Rıo Orinoco, 147 km
SE of San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar 2005, M. Sabaj, N.
Lujan, M. Arce, & T. Wesley.
Sternopygidae
B. Sternopygus sp. (live). ANSP 189018 (146 mm TL). Suriname, Sipaliwini, Lawa River
(Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip), 03u199310N,
054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.
Rhamphichthyidae
C. Gymnorhamphichthys hypostomus (alcohol). INHS 49454 (161 mm LEA). Guyana, Potaro
River (Essequibo drainage), beach on N bank, downstream of Tumatumari Cataract,
05u21948.40N, 59u00904.40W, 22 Oct 1998, M. Sabaj, J. Armbruster, & M. Hardman.
Hypopomidae
D. Hypopomus artedi (alcohol). ANSP 189266 (152 mm TL). Suriname, Sipaliwini, LawaRiver (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
Apteronotidae
E. Apteronotus albifrons (live). AUM 40678 (109 mm SL). Venezuela, Amazonas, RıoManapiare (Ventuari-Orinoco drainage), at San Juan de Manapiare landing, 15 Apr 2004,
J. Para.
86 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
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Plate 14
Rivulidae
A. Rivulus waimacui (alcohol). INHS 49635 (55.5 mm SL). Guyana, tributary of Potaro River
(Essequibo drainage) near Tukeit Cataract, 05u12916.50N, 059u279090W, 27-28 Oct 1998,
L. Page, J. Armbruster, M. Hardman et al.
Poeciliidae
B. Tomeurus gracilis (alcohol). INHS 49017 (27 mm SL). Guyana, East Demerara, Demerara
River (Atlantic drainage), Land of Canaan, 11.8 mi S-SW of Georgetown bearing 213u,06u389390N, 058u119460W, 14 Oct 1998, M. Sabaj, J. Armbruster, M. Hardman et al.
Anablepidae
C. Anableps anableps (alcohol). INHS 49016 (140 mm SL). Same locality as B.
Belonidae
D. Potamorrhaphis guianensis (live). ANSP 179480 (228 mm SL). Guyana, Yuora River
(Ireng-Takutu-Branco drainage), 6.7 km NE of village of Karasabai on road to Tiger
Creek village, 04u039140N, 059u299070W, 31 Oct 2002, M. Sabaj, J. Armbruster, M.Thomas et al.
E. Potamorrhaphis petersi (alcohol). ANSP 163026 (231 mm SL). Venezuela, Amazonas, Rıo
Sipapo (Orinoco drainage), backwater channel behind sandbar 6-7 km above Pendare,
04u519N, 67u439W, 12 Nov 1985, B. Chernoff et al.
F. Pseudotylosurus microps (alcohol). ANSP 179633 (178 mm SL). Guyana, Rupununi River
(Essequibo drainage) at Kwatamang, 4 km SE of Annai, 03u559030N, 059u069010W, 25 Oct2002, M. Sabaj, J. Armbruster, M. Thomas et al.
88 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 89
Plate 15
Cichlidae
A. Crenicichla multispinosa (live). ANSP 187101 (223 mm SL). Suriname, Sipaliwini, Lawa
River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
B. Geophagus harreri (live). ANSP 187136 (170 mm SL). Suriname, Sipalawini, Litanie Riverat mouth and confluence with Marowini River, just upstream from settlement of Konya
Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P. Willink, J.
Mol et al.
C. Geophagus sp. (live). AUM 38940 (103 mm SL). Guyana, Kuyuwini River, main channel
and backwater 19.5 km W of confluence with Essequibo River (Atlantic drainage).
02u149280N, 058u309030W, 11 Nov 2003, M. Sabaj, J. Armbruster, M. Hardman et al.
D. Hoplarchus psittacus (live). AUM 41425 (147 mm SL). Venezuela, Rıo Ventuari (Orinoco
drainage), bedrock outcrops 83 km E-NE of Macuruco, 153 km E-NE of San Fernando
de Atabapo, 04u159120N, 066u209410W, 6 Apr 2004, M. Sabaj, N. Lujan, D. Werneke
et al.
E. Mesonauta insignis (live). AUM 40590 (82 mm SL). Venezuela, Rıo Orinoco near mouth
of Rıo Ventuari, Macuruco Landing, 75 km E of San Fernando de Atabapo, 03u579290N,
067u019560W, 4 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.
F. Cichla intermedia (live). ANSP 189269 . Venezuela, Amazonas, Rıo Orinoco, bedrock
outcrop, 52.9 km SE of San Antonio, 102 km W of La Esmerelda, 03u069010N,
066u279460W, 4 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.
G. Pterophyllum altum (live). AUM 40584 (59 mm SL). Same locality as E.
90 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 91
Plate 16
Synbranchidae
A. Synbranchus marmoratus (live). ANSP 187334 (492 mm TL). Suriname, Sipaliwini, Lawa
River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &
K. Wan.
B. Synbranchus marmoratus (live, ventral view). Same data as A.
Sciaenidae
C. Pachyurus schomburgkii (alcohol). ANSP 162800 (195 mm SL). Venezuela, Amazonas,
Rıo Iguapo approximately 1 hour above its confluence with Rıo Orinoco, 03u099N,
065u289W, 13 Mar 1987, H. Lopez et al.
D. Plagioscion squamosissimus (alcohol). ANSP 177421 (257 mm SL). Guyana, Siparuni
River (Essequibo drainage), Black Water camp, 04u449N, 058u599W, 6 Dec 1997, G.
Watkins et al.
Eleotridae
E. Microphilypnus sp. (live). ANSP 180643. Venezuela, Amazonas, Rıo Negro (Amazonasdrainage), left bank sandy beach and small adjacent backwater 7.2 km NW of San Carlos
de Rio Negro, 01u589110N, 067u069100W, 19 Mar 2005, M. Sabaj, D. Werneke et al.
Achiridae
F. Hypoclinemus mentalis (alcohol). ANSP 163857 (80.5 mm SL). Venezuela, Amazonas,
cano of Rıo Orinoco separating island and beach just downstream of Quiratare, 02u599N,066u049W, 11 Mar 1987, B. Chernoff et al.
G. Soleonasus finis (alcohol). ANSP 179510 (75.5 mm TL). Guyana, Essequibo River
(Atlantic drainage), E bank at Kurukupari, 04u399410N, 058u409310W, 24 Oct 2002, M.H.
Sabaj et al.
Polycentridae
H. Polycentrus schomburgkii (live). FMNH ex ANSP 189014 (22.3 mm SL). Suriname, Para,
Coropinae Creek (Suriname drainage), vicinity of Republiek, 05u299570N, 055u129520W,
28 Apr 2007, M. Sabaj & P. Willink.
Tetraodontidae
I. Colomesus asellus (live). AUM 37945 (35 mm SL). Guyana, Essequibo River (E bank) at
Kurukupari, 04u399410N, 058u409310W, 17 Nov 2003, M. Sabaj, M. Hardman, N. Lujan
et al.
92 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON
NUMBER 17 93
INDEX TO ORDERS, FAMILIES, AND SUBFAMILIES
Acestrorhynchidae
Achiridae
Agoniatinae
Anguilliformes
Anostomidae
Aphyocharacinae
Apteronotidae
Arapaimidae
Aspredinidae
Auchenipteridae
Belonidae
Beloniformes
Bryconinae
Callichthyidae
Cetopsidae
Characidae
Characiformes
Characinae
Cheirodontinae
Chilodontidae
Cichlidae
Clupeidae
Clupeiformes
Crenuchidae
Ctenoluciidae
Curimatidae
Cynodontidae
Cyprinodontiformes
Doradidae
Engraulidae
Erythrinidae
Gasteropelecidae
Genera Incerta Sedis, family
Characidae
Glandulocaudinae
Gobiidae
Gymnotidae
Gymnotiformes
Hemiodontidae
Hemiramphidae
Heptapteridae
Hypopomidae
Hypoptopomatinae
Hypostominae
Iguanodectinae
Lebiasinidae
Lepidosirenidae
Lepidosireniformes
Lithogeninae
Loricariidae
Loricariinae
Myliobatiformes
Ophichthidae
Osteoglossidae
Osteoglossiformes
Parodontidae
PerciformesPimelodidae
Pleuronectiformes
Poeciliidae
Polycentridae
Potamotrygonidae
Pristidae
Pristiformes
PristigasteridaeProchilodontidae
Pseudopimelodidae
Rhamphichthyidae
Rivulidae
Sciaenidae
SerrasalminaeSiluriformes
Sternopygidae
Stethaprioninae
Synbranchidae
Synbranchiformes
Tetragonopterinae
TetraodontidaeTetraodontiformes
Trichomycteridae
Bulletin Editor: Stephen L. GardinerReview Editor: Bruce B. Collette
Copies available as the supply lasts from:
The Custodian of PublicationsBiological Society of WashingtonMRC 116National Museum of Natural HistoryP.O. Box 37012Washington, D.C. 20013-7012
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© Biological Society of Washington, 2009
PROCEEDINGS BACK ISSUES, BULLETINS and SPECIAL PUBLICATIONS
Back issues of the Proceedings of the Biological Society of Washington, Volume 90 to present, are available for $10.00 per issue or $40.00 per Volume. The following Bulletins are still available:
• NO. 2. The Panama Biota: Some Observations Prior to a Sea-Level Canal. Meredith L. Jones, editor. 1972. Price: $5.50
• NO. 5. Natural History of Plummers Island, Maryland, by W. Wirth, W. Grogan and T. Erwin. 1981. Price $11.00
• NO. 7. Studies on Polychaetes in Honor of Marian H. Pettibone. Kristian Fauchald, editor. 1987. Price $25.00
• NO. 8. Results of Recent Research on Aldabra Atoll, Indian Ocean. Brian Kensley, editor. 1988. Price $15.00
• NO. 9. An Illustrated Marine Flora of the Pelican Cays, Belize, by Diane Littler and Mark Littler. 1997. Price $20.00
• NO. 10. Commemorative Volume for the 80th Birthday of Frederick M. Bayer in 2001. Stephen Cairns and Roger F. Cressey, editors. 2001. Price $30.00
• NO. 11. Study of the Dorsal Gill-Arch Musculature of Teleostome Fishes, with Special Reference to the Actinopterygii, by Victor G. Springer and G. David Johnson. 2004. Price $75.00
• NO. 12. Gamba, Gabon: Biodiversité d’une forêt équatoriale africaine.
o Gamba, Gabon: Biodiversity of an equatorial African rainforest. o Alfonso Alonso, Michelle E. Lee, Patrick Campbell, Olivier S. G. Pauwels, and Francisco
Dallmeier, editors. 2006. Price $27.00
• NO. 13. Checklist of the terrestrial vertebrates of the Guiana Shield. Tom Hollowell and Robert P. Reynolds, editors. 2005. Price $20.00
• NO. 14. Checklist of the vascular plants of Plummers Island, Maryland. Stanwyn G. Shetler, Sylvia S. Orli, Elizabeth F. Wells, and Marcie Beyersdorfer. 2006. Price $15.00
NO. 15. The Invertebrate Fauna of Plummers Island, Maryland. John W. Brown, editor. 2008. Price $35.00
• No. 16. Supraneural and pterygiophore insertion patterns in carangid fishes, with description of a new Eocene carangid tribe, †Paratrachinotini, and a survey of anterior anal-fin pterygiophore insertion patterns in Acanthomorpha. Victor G. Springer and William F. Smith-Vaniz. 2008 Price: $15.00
Special Publications
Symposium on Natural History Collections, Past, Present, and Future. Daniel M. Cohen and Roger F. Cressey, editors. 1969. Price $10.00
Index to Volumes 1–100 of the Proceedings. Includes an alphabetical listing of 1st author of all titles and an index by familial category or higher, of all taxa. Phyllis Spangler and Brian Robbins, editors. 1990. Price $10.00
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checklist of freshwater fishes of the guiana shield
BULLETIN of the
Biological Society
of Washington
ISSN 0097-0298