Biological Society of Washington - Department of Botany ...

BULLETIN of the 

Biological Society 

of Washington 

ISSN 0097-0298 

checklist of freshwater fishes of the guiana shield 

10 September 2009 

NUMBER 17

Bulletin Editor: Stephen L. Gardiner 

Review Editor: Bruce B. Collette 

Copies available as the supply lasts from: 

The Custodian of Publications 

Biological Society of Washington 

MRC 116 

National Museum of Natural History 

P.O. Box 37012 

Washington, D.C. 20013-7012 

(Cost $20.00, including postage and handling) 

© Biological Society of Washington, 2009

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-Naturwissenschaftlichen 

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 of 

the Biological Society of Washington, no. 17. 

ii

CONTENTS 

CONTRIBUTORS. . . ......................................................... iv 

ABSTRACT . . . ............................................................. vii 

INTRODUCTION. . . ......................................................... 

Vicki A. Funk and Carol L. Kelloff 

1 

FISHES OF THE GUIANA SHIELD. ............................................. 

Richard P. Vari and Carl J. Ferraris, Jr. 

9 

GUIDE TO THE CHECKLIST . ................................................. 

Aleksandar Radosavljevic 

21 

CHECKLIST. . . ............................................................. 25 

Pristiformes . ............................................................. 25 

Mylobatiformes . . ......................................................... 25 

Osteoglossiformes. ......................................................... 25 

Anguilliformes . . . ......................................................... 25 

Clupeiformes ............................................................. 25 

Characiformes . . . ......................................................... 25 

Siluriformes . ............................................................. 36 

Gymnotiformes . . ......................................................... 45 

Cyprinodontiformes . . . ..................................................... 47 

Beloniformes ............................................................. 48 

Synbranchiformes. ......................................................... 48 

Perciformes . ............................................................. 48 

Pleuronectiformes. ......................................................... 51 

Tetraodontiformes ......................................................... 51 

Lepidosireniformes ......................................................... 51 

PHOTOGRAPHIC ATLAS OF FISHES OF THE GUIANA SHIELD. ..................... 

Mark Sabaj Pérez 

INTRODUCTION 

53 

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: jxa4003@louisiana.edu 

Jonathan W. Armbruster, Department of Biological Sciences, 331 Funchess, Auburn University, Alabama 36849, 

U.S.A., e-mail: armbrjw@auburn.edu 

Paulo A. Buckup, Departamento de Vertebrados, Museu Nacional, Quinta da Boa Vista, 20940-040 Rio de 

Janeiro, RJ, Brazil, e-mail: buckup@acd.ufrj.br 

Ricardo Campos-da-Paz, Escola de Ciências Biológicas, Universidade Federal do Estado do Rio de Janeiro, Av. 

Pasteur, 458/sala 408, Urca - Rio de Janeiro, RJ, 22290-240, Brazil, e-mail: rcpaz@acd.ufrj.br 

Marcelo R. de Carvalho, Instituto de Biociências, Universidade de Sâo Paulo, Rua do Matão, Trav. 14, no. 101, 

São Paulo, SP, 05508-900, Brazil, e-mail: mrcarvalho@ib.usp.br 

Lilian Cassati, Laboratório de Ictiologia, Departamento de Zoologia e Botânica, IBILCE-UNESP, Rua Cristóvão 

Colombo, 2265, São José do Rio Preto, SP, 15054-000, Brazil, e-mail: licasatti@hotmail.com 

Ning Labbish Chao, Universidade Federal do Amazonas, Caixa Postal 3695, Manaus, AM 69051-970, Brazil, 

e-mail: piabas@gmail.com 

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: collettb@si.edu 

Wilson J. E. M. Costa, Laboratório de Ictiologia Geral e Aplicada, Departamento de Zoologia -- UFRJ, Caixa 

Postal 68049, 21944-970 Rio de Janeiro, RJ, Brazil, e-mail: wcosta@acd.ufrj.br 

Fabio di Dario, Universidade Federal do Rio de Janeiro, Av. São José do Barreto, São José do Barreto, Caixa 

Postal 119331, Macae, RJ, 27921-550, Brazil, e-mail: didario@gmail.com 

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: carlferraris@comcast.net 

John P. Friel, Museum of Vertebrates, Cornell University, E151 Corson Hall, Ithaca, New York 14853-2701, 

U.S.A., e-mail: jpf19@cornell.edu 

Vicki A. Funk, Department of Botany, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560-0166, U.S.A., e-mail: funkv@si.edu 

Michel Jégu, Antenne IRD, UR 131, Laboratoire d’Ichtyologie, M.N.H.N., 43 rue Cuvier, 75231 Paris Cedex 05, 

France, e-mail: jegu@mnhn.fr 

Carol L. Kelloff, Department of Botany, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560-0166, U.S.A., e-mail: kelloffc@si.edu 

Sven O. Kullander, Department of Vertebrate Zoology, Swedish Museum of Natural History, P. O. Box 50007, SE- 

104 05 Stockholm, Sweden, e-mail: sven.kullander@nrm.se 

Francisco Langeani Neto, Laboratório de Ictiologia, Departamento de Zoologia e Botânica, IBILCE-UNESP, Rua 

Cristóvão Colombo, 2265, São José do Rio Preto, SP, 15054-000, Brazil, e-mail: langeani@zoo.ibilce.unesp.br 

Flávio C. T. Lima, Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42594, São Paulo, SP, 04299-970, 

Brazil, e-mail: fctlima@usp.br 

Rosana S. Lima, Universidade de Estado de Rio de Janeiro, Faculdade de Formação de Professores, Rua Dr. 

Francisco Portela, 794, São Gonçalo, RJ, 24435-000, Brazil, e-mail: rosanasl@yahoo.com.br 

Carlos A. S. Lucena, Laboratory of Ichthyology, Museu de Ciências e Tecnologia PUCRS, Caixa Postal 1429, 

Porto Alegre, RS, 90619-900, Brazil, e-mail: lucena@pucrs.br 

iv

Paulo H. F. Lucinda, Laboratório de Ictiologia, Universidade do Tocantins - Campus de Porto Nacional Caixa 

Postal 25, Porto Nacional, TO, 77500-000, Brazil, e-mail: plucinda@unitins.br 

John G. Lundberg, The Academy of Natural Sciences, Department of Ichthyology, 1900 Benjamin Franklin 

Parkway, Philadelphia, Pennsylvania 19103, U.S.A., e-mail: lundberg@acnatsci.org 

Luiz R. Malabarba, Departamento de Zoologia -- IB, Universidade Federal do Rio Grande do Sul, Av. Bento 

Gonçalves, 9500 - bloco IV - Prédio 43435, Porto Alegre, RS, 91501-970, Brazil, e-mail: malabarb@ufrgs.br 

John D. McEachran, Department of Wildlife & Fishery Science, Texas A&M University, 22587 AMU, College 

Station, Texas 77843-2258, U.S.A., e-mail: jmceachran@tamu.edu 

Naércio A. Menezes, Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42594, São Paulo, SP, 04299- 

970, Brazil, e-mail: naercio@usp.br 

Cristiano Moreira, Departamento de Ciências Biológicas, Universidade Federal de São Paulo, Rua Prof. Artur 

Riedel, 275, Diadema, SP, 09972-270, Brazil, e-mail: Moreira.c.r@gmail.com 

Carla S. Pavanelli, Fundação Universidade Estadual de Maringá, Nupelia, Av. Colombo, 3690, Maringá, PR, 

87020-900, Brazil, e-mail: carlasp@nupelia.uem.br 

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: alex.radosavljevic@gmail.com 

Robson T. C. Ramos, Depto de Sistemática e Ecologia, Universidade Federal da Paraíba, PB, João Pessoa, 58059- 

900, Brazil, e-mail: robtamar@dse.ufpb.br 

Roberto E. Reis, Laboratory of Ichthyology, Museu de Ciências e Tecnologia PUCRS, Caixa Postal 1429, Porto 

Alegre, RS, 90619-900, Brazil, e-mail: reis@pucrs.br 

Ricardo S. Rosa, Depto de Sistemática e Ecologia, Universidade Federal da Paraíba, João Pessoa, PB, 58059-900, 

Brazil, e-mail: rsrosa@dse.ufpb.br 

Mark Sabaj Pérez, The Academy of Natural Sciences, Department of Ichthyology, 1900 Benjamin Franklin 

Parkway, Philadelphia, Pennsylvania 19103, U.S.A., e-mail: sabaj@acnatsci.org 

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: schaefer@amnh.org 

Oscar A. Shibatta, Departamento de Biologia Animal e Vegetal, Centro de Ciências Biológicas, Universidade 

Estadual de Londrina, 86051-990 Londrina, PR, Brazil, e-mail: shibatta@uel.br 

Mônica Toledo-Piza, Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa 

Postal 11461, 05422-970 São Paulo, SP, Brazil, e-mail: mtpiza@usp.br 

Richard P. Vari, Division of Fishes, National Museum of Natural History, Smithsonian Institution, Washington, 

D.C. 20560-0159, U.S.A., e-mail: varir@si.edu 

Claude Weber, Department d’Herpetologie et d’Ichtyologie, Museum d’Histoire Naturelle, P. O. Box 434, CH- 

1211 Genève, Switzerland, e-mail: claude.weber@mhn.ville-ge.ch 

Marilyn Weitzman, Division of Fishes, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560-0159, U.S.A., e-mail: weitzmam@si.edu 

Stanley H. Weitzman, Division of Fishes, National Museum of Natural History, Smithsonian Institution, 

Washington, D.C. 20560-0159, U.S.A., e-mail: weitzmas@si.edu 

Wolmar Wosiacki, Museu Paraense Emílio Goeldi, Laboratório de Ictiologia, Av. Magalhães Barata 376 Caixa 

Postal Box 399, Belém, PA, 66040-170, Brazil, e-mail: wolmar@museu-goeldi.br 

Angela M. Zanata, Departamento de Zoologia, Universidad Federal da Bahia, Rua Barão de Geremoabo, Campus 

de Ondina, Salvador, BA, 40170-290, Brazil, e-mail: a_zanata@yahoo.com.br 

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) accounting 

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 South 

America. 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

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 biodiversity 

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 systematic 

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 biodiversity 

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 available 

in the field. But they do more than that, beyond 

these basics they increase our knowledge of this fascinating 

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 information 

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 

INTRODUCTION 

V. A. FUNK and CAROL L. KELLOFF 

extraction from Reis et al. (2003) but quickly grew into 

a larger and more involved project. As with any 

endeavor 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 of 

Bolívar and Amazonas, and a portion of Delta 

Amaçuro; all of Guyana, Suriname, and French 

Guiana; and parts of northern Brazil. Several geological 

outliers of the Guiana Shield occur west of the 

Orinoco River in Colombia. In Spanish and Portuguese 

speaking countries, the region is often referred to 

as the ‘‘Guayana’’; thus the terms Colombian 

Guayana, Brazilian Guayana, and Venezuelan 

Guayana are often used. The total area of the Guiana 

Shield is approximately 1,520,000 km 2 . 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 Río 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 Río Negro are connected by the 

Río 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

2 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON 

Figure 1. The Guiana Shield; adapted from Gibbs & Barron (1993), with the region of western outliers indicated. 

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 sediments 

deposited primarily by the Orinoco River and is really 

not strictly part of the Shield; however, some mountains 

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 

Table 1.—Divisions of the Guiana Shield in approximately west to 

east arrangement, with abbreviations used and estimated areas. 

Division Abbr. Area (km 2 ) 

*Colombian Guayana CG 120,325 

Amazonas, Venezuela VA 175,750 

Bolívar, Venenzuela BO 238,000 

*Delta Amacuro, Venezuela DA 40,200 

*Amazonas, Brazil BA 125,550 

*Roraima, Brazil RO 173,750 

*Pará, Brazil PA 243,280 

*Amapá, Brazil AP 98,750 

Guyana GU 214,970 

Suriname SU 163,270 

French Guiana FG 91,000 

Total (km 2 ) 1,896,845 

*5 not all parts of these politically defined areas are included in 

the Guiana Shield region. 

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 

(Arbeláez & Callejas 1999) and south into northern 

Brazil (Leechman 1913, Gibbs & Barron 1993). Within 

this area, erosion has resulted in numerous verticalwalled, 

frequently flat-topped mountains called ‘‘tepuis,’’ 

among them are Chimantí-tepui (2550 m), Cerro 

Duida (2358 m), and Auyán-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

NUMBER 17 3 

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 Auyán-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, Suriname, 

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 approximately 

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 easternmost 

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 Bolívar, 

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 populationrelated 

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 monuments, 

and two biosphere reserves covering about 

142,280 km 2 , more than 30% of its share of the


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 km 2 comprising about 3% of 

the country’s area (Kelloff 2003, Kelloff & Funk 

2004), with additional reserves under consideration. 

Guyana’s 3710 km 2 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 km 2 , over 12% 

of its total area. This includes the recently created 

16,000 km 2 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 designated 

protected areas, but 18 proposed sites total 6710 

km 2 , 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 crossborder 

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 destroyed. 

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 

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 

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. The 

mammals 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 the 

Shield 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 composition 

on finer landscape scales, using developing 

abilities to produce customized checklists for research 

and conservation with Geographic Information System 

(GIS) technologies drawing upon comprehensive databases 

that include georeferenced museum specimen 

records. 

Biological Diversity of the Guiana Shield (BDG) 

The ‘‘Biological Diversity of the Guiana Shield 

Program’’ (BDG) is a field-oriented program of the

NUMBER 17 5 

Figure 2. A comparison of the species lists of the political areas of the Guiana Shield Region. 

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 opportunities; 

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.


The Centre houses collections and research space, 

provides a library, and houses a Geographic Information 

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. The 

collections 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 and 

the 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. 

References 

Arbeláez, M. V., & R. Callejas. 1999. Flórula de la Meseta de 

Arensica de la comunidad de Monochoa (Región de 

Araracuara, Medio Caquetá). Tropenbos, Bogotá, Colombia. 

Berry, P. E., B. K. Holst, & K. Yatskievych (eds.). 1995. Flora of the 

Venezuelan Guayana. Vol. 1: Introduction. J. A. Steyermark, 

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 

checklist of the plant diversity of the Iwokrama Forest, 

Guyana.—Sida Botanical Miscellany 21:1–86. 

Doyle, A. C. 1912. The Lost World. Puffin Books, London. 

Ferrier, S., G. V. N. Powell, K. S. Richardson, G. Manion, J. M. 

Overton, T. F. Allnutt, S. E. Cameron, K. Mantle, N. D. 

Burgess, D. P. Faith, J. F. Lamoreux, G. Kier, R. J. Hijmans, 

V. A. Funk, G. A. Cassis, B. L. Fisher, P. Flemons, D. Lees, J. 

C. Lovett, & R. S. A. R. Van Rompaey. 2004. Mapping more 

of terrestrial biodiversity for Global Conservation assessment.—BioScience 

54(12):1101–1109. 

Funk, V. A., & P. E. Berry. 2005. The Guiana Shield. Pp. 76–79 in G. 

A. Krupnick & W. J. Kress, eds., Plant conservation: a natural 

history approach. University of Chicago Press, Chicago, 235 

pp. 

———, & K. S. Richardson. 2002. Systematic data in biodiversity 

studies: use it or lose it.—Systematic Biology 51:303–316. 

———, ———, & S. Ferrier. 2005. Survey-gap analysis in 

expeditionary research: where do we go from here?— 

Biological Journal of the Linnean Society 85:549–567. 

———, A. K. Sakai, & K. Richardson. 2002. Biodiversity: the 

interface between systematics and conservation.—Systematic 

Biology 51:235–237. 

———, T. Hollowell, P. Berry, C. Kelloff, & S. N. Alexander. 2007. 

Checklist of the plants of the Guiana Shield (Venezuela: 

Amazonas, Bolívar, Delta Amaçuro; Guyana, Surinam, 

French Guiana).—Contributions from the United States 

National Herbarium 55:1–584. 

Gibbs, A. K., & C. N. Barron. 1993. The geology of the Guiana 

Shield. Oxford University Press, New York, 246 pp. 

Hollowell, T., & R. P. Reynolds (eds.). 2005. Checklist of the 

terrestrial vertebrates of the Guiana Shield.—Bulletin of the 

Biological Society of Washington 13:i–ix + 1–98. 

Huber, O. 1995a. Geography and physical features. Pp. 1–61 in P. E. 

Berry, B. K. Holst, & K. Yatskievych, eds., Flora of the 

Venezuelan Guayana. Vol. 1: Introduction. J. A. Steyermark, 

P. E. Berry, & B. K. Holst, general eds. Missouri Botanical 

Garden, St. Louis. 

——— 1995b. Vegetation. Pp. 97–160 in P. E. Berry, B. K. Holst, & 

K. Yatskievych, eds., Flora of the Venezuelan Guayana. Vol. 

1: Introduction. J. A. Steyermark, P. E. Berry, & B. K. Holst, 

general eds. Missouri Botanical Garden, St. Louis. 

———, G. Gharbarran, & V. A. Funk. 1995. Preliminary vegetation 

map of Guyana. Biological Diversity of the Guianas Program, 

Smithsonian Institution,Washington, D.C. 

Kelloff, C. L. 2003. The use of biodiversity data in developing 

Kaieteur National Park, Guyana, for ecotourism and 

conservation.—Contributions to the Study of Biological 

Diversity, University of Guyana, Georgetown 1:1–44. 

———, & V. A. Funk. 2004. Phytogeography of the Kaieteur Falls, 

Potaro Plateau, Guyana: floral distributions and affinities.— 

Journal of Biogeography 31:501–513. 

Leechman, A. 1913. The British Guiana handbook. ‘‘The Argosy’’ 

Co., Ltd., Georgetown, British Guiana, and Dulau & Co., 

London, 283 pp. 

Lindeman, J. C., & S. A. Mori. 1989. The Guianas. Pp. 375–391 in D. 

G. Campbell & H. D. Hammond, eds., Floristic inventory of 

tropical countries. New York Botanical Garden, New York. 

Mendoza, V. 1977. Evolución tectónica del Escudo de Guayana.— 

Boletín de Geología. Publicación Especial 7(3):2237–2270. 

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. 

Schubert, C., & O. Huber. 1990. The Gran Sabana: panorama of a 

region. Lagoven Booklets, Caracas, 107 pp. 

Snow, J. W. 1976. Climates of northern South America. Pp. 295–403 

in W. Schwerdtfeger, ed., Climates of Central and South 

America. Elsevier Scientific Publishing Company, Amsterdam.

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 ichthyofauna 

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 available 

information 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 samples 

returned 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 underappreciated. 

Indications of the scale of the richness of the fish 

fauna inhabiting the rivers of the Guiana Shield 

commenced with the expeditions of the Schomburgk 

brothers, 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 Río Orinoco and Río 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’s 

Library’’ (1841, 1843), including the formal descriptions 

of a series of species. Unfortunately, the specimens 

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 Müller & Troschel (1845, 

FISHES OF THE GUIANA SHIELD 

RICHARD P. VARI and CARL J. FERRARIS, JR. 

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 accessibility 

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 unpublished 

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). Compendia 

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 appreciation 

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 Eigenmann 

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 collections 

(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 (Río 

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 Française]). 

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 Río 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 Río 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 Eigenmann’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 southernmost 

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 incorporated 

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, Río Casiquiare, and Río 

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.’’ Notwithstanding 

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 & Weitzman 

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 ichthyological 

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

NUMBER 17 11 

did so summarized below arranged by the geographic 

subdivisions in the checklist. Many include associated 

ecological and life history information for fish communities 

and individual species. 

Brazil, Pará (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) summarized 

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). Géry (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 publications, 

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 provided 

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 information. 

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 Río 

Negro of the Amazon basin, the north-draining Río 

Orinoco and the entirety of the intervening Río 

Casiquiare. Mago-Leccia (1971) produced the first 

summary of the fishes of the Río 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 

Río 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 ichthyofauna 

of the Río Caroni were published by Lasso 

(1991) and Lasso et al. (1991a, b) and for the Río 

Caura by Lasso et al. (2003a, b), Rodríquez-Olarte et 

al. (2003), and Vispo et al. (2003). The Río 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 ichthyofauna 

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


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 diversity, 

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 demonstrates 

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 variations 

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 (Pará and Amapá; 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 (Arbeláez 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 Solimões 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 physiological, 

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 notwithstanding 

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 Jesús & 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 transport 

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

NUMBER 17 13 

composition of ichthyofaunal communities (see Goulding 

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 rainforests 

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 submerged 

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 freshwater 

fishes to drainage patterns. Historical separations 

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 transgressions 

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 hydrographic 

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., Armbruster 

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 Río Caroni of the Río Orinoco basin and the 

Río Cuyuni, a western tributary of the Essequibo River 

(Lasso et al. 1991a, Sabaj Pérez & 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 interconnection 

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 Río Negro 

of the Amazon River basin with the upper portions of 

the Río Orinoco. This unusual drainage begins as a


bifurcation of the upper portion of the Río 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 Río Orinoco, the 

Río 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 Río 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 Río Casiquiare acts as a partial filter 

that impedes movement of many fish species (Winemiller 

et al. 2008), thereby maintaining differences in 

ichthyofaunal composition between the upper Río 

Negro and upper Río Orinoco. 

Although the Río 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-tosample 

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, universities, 

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 discovery, 

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 freshwater 

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 Ecosystem 

Assessment 2005, Revenga et al. 2005), with 

freshwater fishes consequently the most threatened

NUMBER 17 15 

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 consumption 

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, deforestation 

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 hydroelectric 

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 information 

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 finescaled 

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. 


Completion of this Bulletin was facilitated by S. 

Raredon who prepared the various images that face 

each section. Numerous individuals assisted our field 

efforts 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. 

Literature Cited 

Abell, R., M. L. Thieme, C. Revenga, M. Bryer, M. Kottelat, N. 

Bogutskaya, B. Coad, N. Mandrak, S. Contreras Balderas, W. 

Bussing, M. L. J. Stiassny, P. Skelton, G. R. Allen, P. 

Unmack, A. Naseka, R. Ng, N. Sindorf, J. Robertson, E. 

Armijo, J. V. Higgins, T. J. Heibel, E. Wikramanayake, D. 

Olson, H. L. López, R. E. Reis, J. G. Lundberg, M. H. Sabaj 

Pérez, & P. Petry. 2008. Freshwater ecoregions of the world: a 

new map of biogeographic units for freshwater biodiversity 

conservation.—BioScience 58(5):403–414. 

Arbeláez, F., J. F. Duivenvoorden, & J. A. Maldonado-Ocampo. 

2008. Geological differentiation explains diversity and composition 

of fish communities in upland streams in the southern 

Amazon of Colombia.—Journal of Tropical Ecology 24:505– 

515. 

Armbruster, J. A. 2005. The loricariid catfish genus Lasiancistrus 

(Siluriformes) with descriptions of two new species.—Neotropical 

Ichthyology 3(4):549–569. 

Baker, P. A., G. O. Seltzer, S. C. Fritz, R. B. Dunbar, M. J. Grove, P. 

M. Tapia, S. L. Cross, H. D. Rowe, & J. P. Broda. 2001. The 

history of South American tropical precipitation for the past 

25,000 years.—Science 291:640–643. 

Bancroft, E. 1769. An essay on the natural history of Guiana, in 

South America. Containing a description of many curious 

productions in the animal and vegetable systems of that 

country. In several letters from a gentleman of the medical 

faculty during his residence in that country. London. 

Bleeker, P. 1862. Descriptions de quelques espèces nouvelles de 

Silures de Suriname.—Verslagen en Mededelingen van de 

Koninklijke Akademie van Wetenschappen, afdeling Natuurkunde, 

Amsterdam 14:371–399. 

Boeseman, M. 1952. A preliminary list of Surinam fishes not included 

in Eigenmann’s enumeration of 1912.—Zoologische Mededelingen 

31:179–200. 

———. 1953. Scientific results of the Surinam expedition 1948–1949. 

Part II. Zoology. No. 2. The fishes (I).—Zoologische 

Mededelingen 32:1–24. 

———. 1954. On a small collection of Surinam fishes.—Zoologische 

Mededelingen 33:17–24. 

———. 1968. The genus Hypostomus Lacépède, 1803, and its 

Surinam representatives Siluriformes, Loricariidae).—Zoologische 

Verhandelingen 99:1–89. 

———. 1982. The South American mailed catfish genus Lithoxus 

Eigenmann, 1910, with the description of three new species 

from Surinam and French Guyana and records of related 

species (Siluriformes, Loricariidae).—Proceedings of the 

Koninklijke Nederlandse Akademie van Wetenschappen, 

Series C 85(1):41–58. 

Boujard, T., D. Sabatier, R. Rojas-Beltran, M.-F. Prévost, & J.-F. 

Renno. 1990. The food habits of three allochthonous feeding 

characoids in French Guiana.—Revue d’Ecologie (Terre et 

Vie) 45:247–258. 

Cardoso, Y. P., & J. I. Montoya-Burgos. 2009. Unexpected diversity 

in the catfish Pseudancistrus brevispinis reveals dispersal routes


in a Neotropical center of endemism: the Guyanas Region.— 

Molecular Ecology 18:947–964. 

Chapman, L. J., C. A. Chapman, L. Kaufman, F. Witte, & J. 

Balirwa. 2008. Biodiversity conservation in African inland 

waters: lessons of the Lake Victoria region.—Verhandlungen 

Internationale Vereinigung Limnologie 30(1):16–34. 

Cuvier, G., & M. A. Valenciennes. 1840a. Histoire naturelle des 

poissons. Tome quatorzième. Suite du livre seizième. Labroïdes. 

Livre dix-septième. Des Malacoptérygiens, Pitois- 

Levrault, Paris, xxii + 2 + 464 + 4 p., pls. 389–420. [Volume 

dated 1839] 

———, & ———. 1840b. Histoire naturelle de poissons. Tome 

quinzième. Suite du livre dix-septième. Siluroïdes. Ch. Pitois & 

V e Levrault, Paris & Strasbourg, xxxi + 540 pp. 

De Jesús, M. J., & C. C. Kohler. 2004. The commercial fishery of the 

Peruvian Amazon.—Fisheries 29(4):10–16. 

Dudgeon, D., A. H. Arthington, M. O. Gessner, Z.-I. Kawabata, D. 

J. Knowler, C. Lévêque, R. J. Naiman, A.-H. Prieur-Richard, 

D. Soto, M. L. J. Stiassny, & C. A. Sullivan. 2006. Freshwater 

biodiversity: importance, threats, status and conservation 

challenges.—Biological Reviews 81:163–182. 

Eigenmann, C. H. 1912. The freshwater fishes of British Guiana, 

including a study of the ecological grouping of species and the 

relation of the fauna of the plateau to that of the lowlands.— 

Memoirs of the Carnegie Museum 5:1–578. 

———, & R. S. Eigenmann. 1890. A revision of the South American 

Nematognathi or cat-fishes.—Occasional Papers of the 

California Academy of Sciences 1:1–508. 

Ferreira, A. R. 1983. Viagem filosófica ao Rio Negro. Museu 

Paraense Emílio Goeldi, Belém, Brazil, 775 pp. 

Ferreira, E. J. G. 1993. Composição, distribuição e aspectos 

ecológicos da ictiofauna de um trecho do Rio Trombetas, 

na área de influência da futura UHE cachoeira Porteira, 

Estado do Pará, Brasil.—Acta Amazonica 23 (suplemento):1– 

89. 

———, G. M. dos Santos, & M. Jégu. 1988. Aspectos ecológicos da 

ictiofauna do rio Mucajaí, naárea da ilha Paredão, Roraima, 

Brasil.—Amazoniana 10(3):339–352. 

———, J. Zuanon, B. R. Fosberg, M. Goulding, & S. R. Briglia- 

Ferreira. 2007. Rio Branco: peixes, ecología e conservação de 

Roraima. Amazon Conservation Association, Instituto Nacional 

de Pesquisas da Amazônia, Sociedade Civil de 

Mamirauá, Manaus, Brazil, 201 pp. 

Flecker, A. S. 1996. Ecosystem engineering by a dominant detritivore 

in a diverse tropical stream.—Ecology 77(6):1845–1854. 

Géry, J. 1972. Poissons characoïdes des Guyanes. I. Généralitées. II. 

Famille des Serrasalmidae.—Zoologische Verhandelingen No. 

122:1–250. 


Shield. Oxford University Press, New York. 246 pp. 

Girardo, A., C. A. Lasso, H. F. Samudio, & J. H. Hernández- 

Acevedo. 2007. Nuevas adiciones a la ictiofauna de la cuenca 

del río Cuyuní en Venezuela.—Memoria de la Fundación La 

Salle de Ciencias Naturales (for 2006) 66(166):151–154. 

Goulding, M., R. Barthem, & E. J. G. Ferreira. 2003. The 

Smithsonian Atlas of the Amazon. Smithsonian Books, 

Washington, D.C., 253 pp. 

———, M. Leal-Carvalho, & E. J. G. Ferreira. 1988. Rio Negro, rich 

life in poor water: Amazonian diversity and foodchain ecology 

as seen through fish communities. SPB Academic Publishing, 

The Hague, 200 pp. 

Gronovius, L. T. 1754. Museum Ichthyologicum sistens Piscium 

indigenorum & quorundam exoticorum, qui in Museo 

Laurentii Theodorii Gronovii, J. U. D. adservantur, descriptiones 

ordine systematico. Lugduni, Batavorum. 

———. 1756. Musei Ichthyologici tomus secundus sistens piscium 

indigenorum & nonnullorum exoticorum, quorum maxima 

pars in Museo Laurentii Theodorii Gronovii, J. U. D. 

adservantur, nec non quorumdam in aliis Museis observatorum 

descriptiones. Lugduni, Batavorum. 

Hammond, D. S. 2005. Biophysical features of the Guiana Shield. 

Pp. 15–194 in D. S. Hammond, ed., Tropical forests of the 

Guiana Shield: Ancient forests of the modern world. CABI 

Publishing, Cambridge, Massachusetts. 

Hancock, J. 1828. Notes on some species of fishes and reptiles, from 

Demerara, presented to the Zoological Society by John 

Hancock, Esq., corr. memb. Zool. Soc. In a letter addressed 

to the secretary of the Society.—Zoological Journal London 

4:240–247. 

Hardman, M., L. M. Page, M. H. Sabaj, J. W. Armbruster, & J. H. 

Knouft. 2002. A comparison of fish surveys made in 1908 and 

1998 of the Potaro, Essequibo, Demerara, and coastal river 

drainages of Guyana.—Ichthyological Exploration of Freshwaters 

13:225–238. 

Harold, A. S., & R. P. Vari. 1994. Systematics of the trans-Andean 

species of Creagrutus (Ostariophysi: Characiformes: Characidae).—Smithsonian 

Contributions to Zoology 551:1–31. 

Hilhouse, W. 1825. Indian notices: or, sketches of the habits, 

characters, languages, superstitions, soil, and climate, of the 

several nations; with remarks on their capacity for colonization, 

present government, and suggestions for future improvement 

and civilization. Also, the ichthyology of the fresh waters 

of the interior. Printed for the author. 133 pp. 

Hollowell, T., & R. P. Reynolds (eds.). 2005. Checklist of the 

terrestrial vertebrates of the Guiana Shield.—Bulletin of the 

Biological Society of Washington 13:i–ix + 1–98. 

Hoorn, C. 1994. An environmental reconstruction of the palaeo- 

Amazon River system (Middle-Late Miocene, NW Amazonia).—Palaeogeography, 

Palaeoclimatology, Palaeoecology 

112:187–238. 

Jardine, W. 1841. The natural history of the fishes of Guiana. Part I. 

In W. Jardine, ed., The Naturalist’s Library, Ichthyology, 

volume 3. H. Lizars, Edinburgh, 263 pp. 

———. 1843. The natural history of the fishes of Guiana. Part II. In 

W. Jardine, ed., The Naturalist’s Library, Ichthyology, 

volume 5. W. H. Lizars, Edinburgh, 212 pp. 

Keith, P., P.-Y. Le Bail, & P. Planquette. 2000. Atlas des poissons 

d’eau douce de Guyane. Volume 2, part 1. Batrachoidiformes, 

Mugiliformes, Beloniformes, Cyprinodontiformes, Synbranchiformes, 

Perciformes, Pleuronectiformes and Tetraodontiformes. 

Muséum National d’Histoire Naturelle, Institut 

d’Ecologie et de Gestion de la Biodiversité, Service du 

Patrimoine Naturel, Paris, 286 pp. 

Killeen, T. J. 2007. A perfect storm in the Amazon wilderness: 

development and conservation in the context of the Initiative 

for the Integration of the Regional Infrastructure of 

South America (IIRSA).—Advances in Applied Biodiversity 

Science, volume 7. Center for Applied Biodiversity 

Science, Conservation International, Washington, D.C., 

102 pp. 

———, T. M. Siles, T. Grimwood, L. L. Tieszen, M. K. Steininger, 

C. J. Tucker, & S. Panfil. 2002. Habitat heterogeneity on a 

forest-savanna ecotone in Noel Kempff Mercado National 

Park (Santa Cruz, Bolivia): implications for the long-term 

conservation of biodiversity in a changing climate. Pp. 285– 

312 in G. A. Bradshaw & P. A. Marquet, eds., How 

landscapes change: human disturbance and ecosystem fragmentation 

in the Americas. Ecological Studies 162, Springer, 

Berlin. 

Kullander, S. O., & H. Nijssen. 1989. The cichlids of Surinam 

(Teleostei: Labroidei). E. J. Brill, Leiden, 256 pp. 

Lasso, C. A. 1991. Los peces de la Gran Sabana, Alto Caroni, 

Venezuela.—Memoria, Sociedad de Ciencias Naturales La 

Salle 49/50 (131/134):209–285. 

———. 1992. Composicion y aspectos ecologicos de la ictiofauna del 

bajo rio Suapure, Serrania de Los Pijiguaos (Escudo de

NUMBER 17 17 

Guayana), Venezuela.—Memoria, Sociedad de Ciencias 

Naturales La Salle 52(138):5–56. 

———, A. Machado-Allison, & R. Pérez Hernández. 1991a. 

Consideraciones zoogeografica de los peces de la Gran Sabana 

(Alto Caroni), Venezuela, y sus relaciones con los cuencas 

vecinas.—Memoria, Sociedad de Ciencias Naturales La Salle 

49/50 (131/134):109–129. 

———, D. Novoa, & F. Ramos. 1991b. La ictiofauna del Lago de 

Guri: composición, abundancia y potencial pesquero. Parte I: 

consideraciones generales e inventario de la ictiofauna del 

Lago de Guri con breve descripción de las especies de interes 

para la pesca deportiva y comercial.—Memoria, Sociedad de 

Ciencias Naturales La Salle 49/50 (131/134):141–158. 

———, C. R. Vispo, & O. Lasso-Alcalá. 2003b. Floodplain fishes of 

the Caura River, southern Venezuela.—Scientia Guaianae 12: 

273–295. 

———, D. Lew, D. C. Taphorn, C. DoNascimiento, O. Lasso- 

Alcalá, F. Provenzano, & A. Machado-Allison. 2004a. 

Biodiversidade ictiologica continental de Venezuela. Parte 1. 

Lista de especies y distribución por cuencas.—Memorias de la 

Fundación La Salle de Ciencias Naturales 159–160 (for 2003): 

105–195. 

———, A. Machado-Allison, D. Taphorn, D. Rodríguez-Olarte, 

C. R. Vispo, B. Chernoff, F. Provenzano, O. Lasso-Alcalá, 

A. Cervó, K. Nakamura, N. González, J. Meri, C. Silvera, 

A. Bonilla, H. López-Rojas, & D. Machado-Aranda. 2003a. 

The fishes of the Caura River basin, Orinoco drainage, 

Venezuela. Annotated Checklist.—Scientia Guianae 12:223– 

245. 

———, J. I. Mojica, J. S. Usma, J. A. Maldonaldo O., C. 

DoNascimiento, D. C. Taphorn, F. Provenzano, O. M. Lasso 

Alcalá, G. Galvis, L. Vásquez, M. Lugo, A. Machado Allison, 

R. Royero, C. Suárez, & A. Ortega Lara. 2004b. Peces de las 

cuenca del río Orinoco. Parte I: lista de especies y distribución 

por subcuencas.—Biota Colombiana 5:95–157. 

Le Bail, P.-Y. A, P. Keith, & P. Planquette. 2000. Atlas des poissons 

d’eau douce de Guyane. Volume 2, part 2. Siluriformes. 

Muséum National d’Histoire Naturelle, Institut d’Ecologie et 

de Gestion de la Biodiversité, Service du Patrimoine Naturel, 

Paris, 307 pp. 

Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum 

classes, ordines, genera, species, cum characteribus, differentiis, 

synonymis, locis. Tomus I. Editio decima, reformata. 

Holmiae. 

Lowe (McConnell), R. H. 1964. The fishes of the Rupununi savanna 

district of British Guiana, South America. Part 1. Ecological 

groupings of fish species and effects of the seasonal cycle on 

the fish.—Journal of the Linnean Society (Zoology) 45(304): 

103–140. 

Lowe-McConnell, R. H. 1995. Ecological studies in tropical fish 

communities. Cambridge University Press, Cambridge, 382 

pp. 

Machado-Allison, A., B. Chernoff, R. Royero-León, F. Mago- 

Leccia, J. Velázquez, C. Lasso, H. López-Rojas, A. Bonilla- 

Rivero, F. Provenzano, & C. Silvera. 2000. Ictiofauna de la 

cuenca del Río Cuyuní en Venezuela.—Interciencia 25(1):13– 

21. 

Mago-Leccia, F. 1971. La ictiofauna del Casiquiare.—Revista 

Defensa de la Naturaleza (Venezuela) 1(4):5–11. 

Millennium Ecosystem Assessment. 2005. Ecosystems and human 

well-being: synthesis. Island Press, Washington, D.C., 160 pp. 

Müller, J., & F. H. Troschel. 1845. Horae Ichthyologicae. Beschreibung 

und Abbildung neuer Fische. Ertes und zweites Heft. 

Die Familie der Characinen. Verlag von Veit & Comp., Berlin, 

pp. 1–40. 

———, & ———. 1848. Fische. Pp. 618–644 in Reisen in Britisch- 

Guiana in den Jahren 1840–44. Im Auftrag Sr. Mäjestat des 

Königs von Preussen ausgeführt von Richard Schomburgk. 

[Versuch einer Fauna und Flora von Britisch-Guiana.] Vol. 3, 

Berlin. [Date sometimes seen as 1849.] 

———, & ———. 1849. Horae Ichthyologicae. Beschreibung und 

Abbildung neuer Fische. Drittes Heft. Verlag von Veit & 

Comp., Berlin, pp. 1–27 + additional p. 24. 

Myers, G. S. 1927. Descriptions of new South American fresh-water 

fishes collected by Dr. Carl Ternetz.—Bulletin of the Museum 

of Comparative Zoology 68(3):107–135. 

———, & S. H. Weitzman. 1960. A Brazilian pike-characid, 

Boulengerella lateristriga, rediscovered in the Rio Negro.— 

Stanford Ichthyological Bulletin 7(4):201–205. 

Nelson, J. S. 2006. Fishes of the World. 4 th Edition. John Wiley & 

Sons, Hoboken, New Jersey, 624 pp. 

Nijssen, H. 1970. Revision of the Surinam catfishes of the genus 

Corydoras Lacépède, 1803 (Pisces, Siluriformes, Callichthyidae).—Beaufortia 

18(230):1–75. 

Ouboter, P. E., & J. H. A. Mol. 1993. The fish fauna of Suriname. 

Pp. 133–154 in P. E. Ouboter, ed., The Freshwater Ecosystems 

of Suriname. Kluwer Academie Publishers, Dordrecht, The 

Netherlands, 313 pp. 

Pellegrin, J. 1908. Les poissons d’eau douce de la Guyane 

Française.—Revue Coloniale. Publication du Ministère des 

Colonies 67:557–587. 

Planquette, P., P. Keith, & P.-Y. Le Bail. 1996. Atlas des poissons 

d’eau douce de Guyane. Volume 1, Collection du Patrimoine 

Naturel, vol. 22. Institut d’Ecologie et de Gestion de la 

Biodiversité du Muséum National d’Histoire Naturelle, 

Institut national de la Recherche Agronomique, Conseil 

Supérior de la Pêche, Paris, 429 pp. 

Polis, G. A., W. B. Anderson, & R. D. Holt. 1997. Toward an 

integration of landscape and food web ecology: the dynamics 

of spatially subsidized food webs.—Annual Review of 

Ecology and Systematics 28:289–316. 

Puyo, J. 1949. Poissons de la Guyane française. Faune de l’Empire 

français, 12. Librairie Larose, Paris, 280 pp. 

Regan, C. T. 1905a. On drawings of fishes of the rio Negro.— 

Proceedings of the Zoological Society of London 1905:189– 

190. 

———. 1905b. A revision of the fishes of the South-American cichlid 

genera Acara, Nannacara, Acaropsis, and Astronotus.—Annals 

and Magazine of Natural History (Series 7) 15:329–347. 

———. 1905c. A revision of the fishes of the American cichlid genus 

Cichlosoma and some allied genera.—Annals and Magazine of 

Natural History (Series 7) 16:60–77. 

Reis, R. E. 2004. Otocinclus cocama, a new uniquely colored 

loricariid catfish from Peru (Teleostei: Siluriformes), with 

comments on the impact of taxonomic revisions to the 

discovery of new taxa.—Neotropical Ichthyology 2(3):109– 

115. 

———, S. O. Kullander, & C. J. Ferraris, Jr. (eds.). 2003. Check List 

of the freshwater fishes of South and Central America. 


Revenga, C., I. Campbell, R. Abell, P. de Villiers, & M. Bryer. 2005. 

Prospects for monitoring freshwater ecosystems towards the 

2010 targets.—Philosophical Transactions of the Royal 

Society of London B 360(1454):397–413. 

Ribeiro, A. C., R. C. Benine, & C. A. Figueiredo. 2004. A new species 

of Creagrutus Günther (Teleostei: Ostariophysi: Characiformes), 

from the upper Río Paraná basin, central Brazil.— 

Journal of Fish Biology 64:597–611. 

Rodríguez-Olarte, D., D. C. Taphorn, C. Lasso, & C. R. Vispo. 2003. 

Fishes of the lower Caura River, Orinoco basin, Venezuela.— 

Scientia Guaianae 12:181–221. 

Royero, R., A. Machado-Allison, B. Chernoff, & D. Machado-Aranda. 

1992. Peces del Río Atabapo, Territorio Federal Amazona,s 

Venezuela.—Acta Biologica Venezuelica 14(1):41–55. 

Sabaj Pérez, M. H., & J. L. O. Birindelli. 2008. Taxonomic revision of 

extant Doras Lacepède, 1803 (Siluriformes: Doradidae) with


descriptions of three new species.—Proceedings of the 

Academy of Natural Sciences of Philadelphia 157:189–233. 

Sioli, H. 1975. Tropical rivers as expressions of their terrestrial 

environments. Pp. 275–288 in F. B. Golley & E. Medina, eds., 

Tropical ecosystems: Trends in terrestrial and aquatic 

research. Springer, New York. 

Steindachner, F. 1915. Beiträge zur Kenntnis der Flussfische 

Südamerika’s.—V. Denkschriften der Mathematische-Naturwissenschaftenlichen 

Classe der Kaiserlichen Akademie der 

Wissenschaften Wien 93:15–105. [Date based on release of 

separates of main work published in 1917.] 

Sternberg, H. O. 1975. The Amazon River of Brazil.—Geographische 

Zeitschrift 10:1–74. 

Taphorn B, D. C., H. López-Fernández, & C. R. Bernard. 2008. 

Apareiodon agmatos, a new species from the upper Mazaruni 

river, Guyana (Teleostei: Characiformes: Parodontidae).— 

Zootaxa 1925:31–38. 

Toledo-Piza, M., N. A. Menezes, & G. M. dos Santos. 1999. Revision 

of the neotropical fish genus Hydrolycus (Ostariophysi: 

Characiformes: Cynodontidae) with the description of two 

new species.—Ichthyological Exploration of Freshwaters 

10(3):255–280. 

Torres-Mejia, M., & R. P. Vari. 2005. New species of Creagrutus 

(Teleostei: Characiformes: Characidae) from the Río Magdalena 

basin, Colombia.—Copeia. 2005(4):812–817. 

Vaillant, L. L. 1899. Note préliminaire sur les collections ichthyologiques 

recueillies par M. Geay en 1897 et 1898 dans la 

Guyane française et le Contesté franco-brésilien.—Bulletin du 

Muséum d’histoire naturelle, Paris. 5(4):154–156. 

———. 1900. Contribution à l’étude de la fauna ichthyologique de la 

Guyane française et du contesté Franco-Brésilien.—Nouvelles 

Archives du Muséum d’histoire naturelle, Paris. Series 4, 4:126–136. 

Vari, R. P., & C. J. Ferraris, Jr. 2006. The catfish genus Tetranematichthys 

(auchenipteridae).—Copeia 2006 (2):168–180. 

———, & A. S. Harold. 2001. Phylogenetic study of the Neotropical 

fish genera Creagrutus Günther and Piabina Reinhardt 

(Teleostei: Ostariophysi: Characiformes), with a revision of 

the Cis-Andean species.—Smithsonian Contributions to 

Zoology 613:v + 1–237. 

———, & F. C. T. Lima. 2003. New species of Creagrutus (Teleostei: 

Characiformes: Characidae) from the Rio Uaupés Basin, 

Brazil.—Copeia 2003(3):583–587. 

———, & L. Malabarba. 1998. Neotropical Ichthyology: An 

overview. Pp. 23–142 in L. R. Malabarba, R. E. Reis, R. P. 

Vari, Z. M. S. Lucena, & C. A. S. Lucena, eds., Phylogeny and 

classification of neotropical fishes. Edipucrs, Porto Alegre, 

Brazil, 603 pp. 

Vispo, C., C. Lasso, O. Lasso-Alcalá, & N. González. 2003. 

Geographical and temporal variation in fish communities of 

the floodplain lakes of the lower Caura River drainage in the 

Venezuelan Guayana.—Scientia Guaianae 12:297–327. 

Wallace, A. R. 2002. Peixes do Rio Negro, Fishes of the Rio Negro. 

Alfred Russel Wallace (1850–1852). Organization, introductory 

text, and translation by Mônica de Toledo-Piza Ragazzo. 

Editera da Universidade de São Paulo, São Paulo, 517 pp. 

Watkins, G., W. Saul, E. Holm, C. Watson, D. Arjoon, & J. Bicknell. 

2004. The fish fauna of the Iwokrama Forest.—Proceedings of 

the Academy of Natural Sciences of Philadelphia 154(1):39– 

53. 

Winemiller, K. O., H. López-Fernández, D. C. Taphorn, L. G. Nico, 

& A. Barbarino Duque. 2008. Fish assemblages of the 

Casiquiare River, a corridor and zoogeographical filter for 

dispersal between the Orinoco and Amazon basins.—Journal 

of Biogeography 35:1551–1563.

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 formations 

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 Río Orinoco that runs closer to the 

northern limit of the Shield were included. This 

preliminary assemblage was updated from the literature 

published since CLOFFSCA’s release and the 

resultant summaries sent to relevant authorities for 

review. Whenever possible, the families and/or subfamilies 

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 Loricariidae 

(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 (Amapá, 

Amazonas, Pará, 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 

Table 3.—Regions of the Guiana Shield used in checklist in 

approximately west to east order. 

CG Colombian Guayana 

VA Venezuela—Amazonas 

BO Venezuela—Bolívar 

DA Venezuela—Delta Amacuro 

BA Brazil—Amazonas 

RO Brazil—Roraima 

PA Brazil—Pará 

AP Brazil—Amapá 

GU Guyana 

SU Suriname 

FG French Guiana 

GUIDE TO THE CHECKLIST 

ALEKSANDAR RADOSAVLJEVIC 

Shield. Regional abbreviations in the checklist are 

followed by a ‘‘?’’ in instances when distributions were 

uncertain or questionable. 


(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. 

———. 2003b. Peckoltia sabaji, a new species from the Guyana 

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. 

———. 2005. The loricariid catfish genus Lasiancistrus (Siluriformes) 

with descriptions of two new species.—Neotropical Ichthyology 

3(4):549–569. 

———. 2008. The genus Peckoltia with the description of two new 

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: Loricariidae) 

from Guyana.—Zootaxa 920:1–12. 

———, & D. C. Taphorn. 2008. A new species of Pseudancistrus 

from the Río Caroní, Venezuela (Siluriformes: Loricariidae).—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 

Río Caura, Venezuela.—Zootaxa 1610:53–59. 

———, G. Z. Pelição, & 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 Río Orinoco, Venezuela, with 

comments on the genus (Siluriformes: Doradidae).—Copeia 

2007(3):672–684. 

Bührnheim, C. M., & L. R. Malabarba. 2007. Redescription of 

Odontostilbe pulchra (Gill, 1858) (Teleostei: Characidae: 

Cheirodontinae), and description of two new species from 

the río Orinoco basin.—Neotropical Ichthyology 5(1):1–20. 

Buitrago-Suárez, 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. 

caudomaculatus (Günther).—Zootaxa 1094:1–23. 

de Chambrier, S., & J. I. Montoya-Burgos. 2008. Pseudancistrus 

corantijniensis, a new species from the Guyana Shield


(Siluriformes: Loricariidae) with a molecular and morphological 

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 aspredinoides 

(Siluriformes: Heptapteridae), a new catfish from the 

Río 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: Aspredinidae).—Neotropical 


Garutti, V. 2003. Revalidação de Astyanax rupununi Fowler, 1914 

(Teleostei, Characidae) e descrição de duas espécies novas para 

ogênero.—Papéis Avulsos de Zoologia (São Paulo) 43(1):1–9. 


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. 

Jégu, M., P. Keith, & P.-Y. Le Bail. 2003. Myloplus planquettei sp. n. 

(Teleostei: Characidae) une nouvelle espèce de grand Serrasalminae 

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, Pará 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 género Trichomycterus (Siluriformes: Trichomycteridae) 

de la Gran Sabana, Escudo de las Guayanas, Venezuela.— 

Revista de Biología 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 

Alcalá, G. Galvis, L. Vásquez, M. Lugo, A. Machado Allison, 

R. Royero, C. Suárez, & A. Ortega Lara. 2004. Peces de las 

cuenca del río Orinoco. Parte I: lista de especies y distribución 

por subcuencas.—Biota Colombiana 5:95–157. 

Lasso-Alcalá, O. M., D. C. Taphorn B., C. A. Lasso, & O. León- 

Mata. 2006. Rivulus sape, a new species of killifish (Cyprinodontiformes: 

Rivulidae) from the Paragua River system, 

Caroní 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. 

López-Fernández, 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: Loricariidae).—Ichthyological 

Exploration of Freshwaters, 18(2): 

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 


Malabarba, M. C. S. L. 2004. Revision of the Neotropical genus 

Triportheus Cope, 1872 (Characiformes: Characidae).—Neotropical 


Montaña, C. G., H. López-Fernández, & D. C. Taphorn. 2008. A 

new species of Crenicichla (Perciformes: Cichlidae) from the 

Ventuari River, Upper Orinoco River Basin, Amazonas State, 


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 


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 (Pleuronectiformes: 

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. 


Román-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. 

Römer, 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 (Siluriformes: 

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 Pérez, M. H., & J. L. O. Birindelli. 2008. Taxonomic revision of 

extant Doras Lacepède, 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 Río 

Caroni, Venezuela (Gymnotiformes: Apteronotidae).—Ichthyological 

Exploration of Freshwaters 17(1):1–8. 

———, & R. P. Vari. 2009. The South American electric fish genus 

Platyurosternarchus (Gymnotiformes: Apteronotidae).—Copeia 

2009(2):233–244.

NUMBER 17 23 

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 río 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 winterbottomi: 

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 Río Orinoco system, 

with a redescription of S. Isognathus from the Río Paraguay 

system.—Copeia 2007(3):711–725. 

Staeck, W. 2003. Cichliden-Lexikon, Teil 3: Südamerikanische 

Zwergbuntbarsche. Dähne Verlag, Ettlingen, Germany, 219 pp. 

———, & I. Schindler. 2007. Description of Laetacara fulvipinnis sp. 

n. (Teleostei: Perciformes: Cichlidae) from the upper drainages 

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: Rivulidae).—Zootaxa 

1246:1–13. 

Taphorn, B. D. C., H. López-Fernández, & C. R. Bernard. 2008. 



Zootaxa 1925:31–38. 

Taphorn, B., C. G. Montaña, & P. Buckup. 2006. Characidium 

longum (Characiformes: Crenuchidae), a new fish from 


Thomas, M. R., & L. H. Rapp Py-Daniel. 2008. Three new species of 

the armored catfish genus Loricaria (Siluriformes: Loricariidae) 

from river channels of the Amazon basin.—Neotropical 


Vari, R. P., & C. J. Ferraris, Jr. 2006. The catfish genus Tetranematichthys 

(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 killifish 

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: Loricariidae).—Neotropical 


———, 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. Géry. 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. Géry, & I. Isbrücker. 2004. Moenkhausia rara sp. n. – eine 

neue, bereits bestandsgefährdete Salmler-Art (Teleostei: Characiformes: 

Characidae) aus Surinam und Französisch Guayana 

mit einer ergänzenden Beschreibung von M. simulata (Eigenmann 

in Pearson, 1924).—Zoologische Abhandlungen (Dresden) 

54:19–30.

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 (Müller & Henle, 1841) BO DA RO 

Potamotrygon boesemani Rosa, Carvalho & Almeida, 2008 GU SU 

Potamotrygon marinae Deynat, 2006 FG 

Potamotrygon motoro (Müller & Henle, 1841) CG VA BO RO 

Potamotrygon orbignyi (Castelnau, 1855) VA BO DA GU SU FG 

Potamotrygon schroederi Fernández-Yépez, 1957 VA BO DA 

Potamotrygon scobina Garman, 1913 RO PA 

Order: Osteoglossiformes 

Family: Osteoglossidae—Carl J. Ferraris, Jr. 

Osteoglossum bicirrhosum (Cuvier, 1829) 

Family: Arapaimidae—Carl J. Ferraris, Jr. 

CG RO AP GU FG 

Arapaima gigas (Schinz, 1822) GU 

Order: Anguilliformes 

Family: Ophichthidae—Sven O. Kullander 

Stictorhinus potamius (Böhlke & McCosker, 1975) BO DA 

Order: Clupeiformes 

Family: Clupeidae—Carl J. Ferraris, Jr. 

Rhinosardinia bahiensis (Steindachner, 1879) 

Family: Engraulidae—Carl J. Ferraris, Jr. & Sven O. Kullander 

BO DA AP GU SU FG 

Anchoa spinifer (Valenciennes, 1848) DA GU SU FG 

Anchovia surinamensis (Bleeker, 1866) BO DA GU SU FG 

Anchoviella brevirostris (Günther, 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 (Günther, 1868) BO DA GU SU FG 

Lycengraulis grossidens (Spix & Agassiz, 1829) DA SU FG 

Pterengraulis atherinoides (Linnaeus, 1766) 

Family: Pristigasteridae—Fabio di Dario 

BO DA SU FG 

Pellona castelnaeana (Valenciennes, 1847) CG VA BO DA 

Pellona flavipinnis (Valenciennes, 1836) CG VA BO DA GU SU FG 

Family: Parodontidae—Carla S. Pavanelli 

Apareiodon agmatos Taphorn, López-Fernández & Bernard, 

2008 

Order: Characiformes 

Apareiodon gransabana Starnes & Schindler, 1993 BO GU SU FG 

Apareiodon orinocensis Bonilla et al., 1999 VA BO 

GU


Parodon apolinari Myers, 1930 CG VA BO 

Parodon bifasciatus Eigenmann, 1912 RO GU 

Parodon guyanensis Géry, 1959 VA BO GU SU FG 

Parodon suborbitalis Valenciennes, 1850 

Family: Curimatidae—Richard P. Vari 

BO 

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 (Müller & Troschel, 1844) RO GU 

Curimatella dorsalis (Eigenmann & Eigenmann, 1889) CG VA BO DA 

Curimatella immaculata (Fernández-Yépez, 1948) CG VA BO DA RO GU 

Curimatopsis crypticus Vari, 1982 AP GU SU FG 

Curimatopsis evelynae Géry, 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 (Günther, 1864) CG VA BO BA RO GU SU FG 

Potamorhina altamazonica (Cope, 1878) CG VA BO DA 

Psectrogaster ciliata (Müller & Troschel, 1844) CG VA BO DA RO GU 

Psectrogaster essequibensis (Günther, 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 Géry, Planquette & Le Bail, 1991 

Family: Prochilodontidae—Richard P. Vari 

AP SU FG 

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 

Family: Anostomidae—Richard P. Vari 

SU FG 

Abramites hypselonotus (Günther, 1868) VA BO DA GU

NUMBER 17 27 

Anostomoides atrianalis Pellegrin, 1908 VA BO 

Anostomoides laticeps (Eigenmann, 1912) VA BO GU 

Anostomus anostomus (Linnaeus, 1758) GU 

Anostomus brevior Géry, 1961 FG 

Anostomus plicatus Eigenmann, 1912 GU SU 

Anostomus spiloclistron Winterbottom, 1974 SU 

Anostomus ternetzi Fernández-Yépez, 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 Günther, 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 Géry, Planquette & Le Bail, 1991 SU FG 

Leporinus granti Eigenmann, 1912 GU FG 

Leporinus latofasciatus Steindachner, 1910 VA BO DA 

Leporinus lebaili Géry & Planquette, 1983 SU FG 

Leporinus leschenaulti Valenciennes, 1850 FG 

Leporinus maculatus Müller & Troschel, 1844 GU SU FG 

Leporinus megalepis Günther, 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 & Jégu, 1987 PA 

Schizodon fasciatus Spix & Agassiz, 1829 BO FG


Schizodon scotorhabdotus Sidlauskas, Garavello & Jellen, 2007 VA BO DA 

Synaptolaemus cingulatus Myers & Fernández-Yépez, 1950 

Family: Chilodontidae—Richard P. Vari 

VA BO 

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 Müller & Troschel, 1844 CG VA BO DA GU SU 

Chilodus zunevei Puyo, 1946 

Family: Crenuchidae—Paulo A. Buckup 

FG 

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, Montaña & 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 Günther, 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 (Géry, 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 Géry, 1965 CG VA BO 

Skiotocharax meizon Presswell, Weitzman & Bergquist, 2000 

Family: Hemiodontidae—Francisco Langeani Neto 

GU 

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 Géry, Planquette & Le Bail, 1991 AP FG

NUMBER 17 29 

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 Günther, 1864 VA BO 

Hemiodus huraulti (Géry, 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 Böhlke, 1955 CG VA BO BA 

Hemiodus unimaculatus (Bloch, 1794) AP GU SU FG 

Hemiodus vorderwinkleri (Géry, 1964) 

Family: Gasteropelecidae—Stanley H. Weitzman 

BO GU 

Carnegiella marthae Myers, 1927 CG VA BO DA BA 

Carnegiella strigata (Günther, 1864) CG VA BO DA GU SU 

Gasteropelecus sternicla (Linnaeus, 1758) BO DA GU FG 

Thoracocharax stellatus (Kner, 1858) 

Family: Characidae 

Genera Incerta Sedis—Flávio C. T. Lima 

CG VA BO DA 

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 Géry, Planquette & Le Bail, 1988 AP FG 

Astyanax metae Eigenmann, 1914 VA BO 

Astyanax mutator Eigenmann, 1909 GU 

Astyanax myersi (Fernández-Yépez, 1950) VA BO 

Astyanax rupununi Garutti, 2003 GU 

Astyanax scintillans Myers, 1928 VA BO 

Astyanax siapae Garutti, 2003 VA 

Astyanax validus Géry, Planquette & Le Bail, 1991 FG 

Astyanax venezuelae Schultz, 1944 VA BO 

Aulixidens eugeniae Böhlke, 1952 VA BO 

Bryconamericus alpha Eigenmann, 1914 CG VA BO 

Bryconamericus beta Eigenmann, 1914 VA BO 

Bryconamericus cinarucoense Román-Valencia, Taphorn & Ruiz- 

C., 2008 

GU 

Bryconamericus cismontanus Eigenmann, 1914 VA BO 

Bryconamericus cristiani Román-Valencia, 1998 CG 

Bryconamericus deuterodonoides Eigenmann, 1914 VA BO 

Bryconamericus hyphesson Eigenmann, 1909 GU 

Bryconamericus macropthalmus Román-Valencia, 2003 VA 

Bryconamericus orinocoense Román-Valencia, 2003 VA 

Bryconamericus subtilisform Román-Valencia, 2003 BO 

Bryconexodon trombetasi Jégu, Santos & Ferreira, 1991 PA


Bryconops affinis (Günther, 1864) GU SU FG 

Bryconops alburnoides Kner, 1858 CG VA BO 

Bryconops caudomaculatus (Günther, 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 (Fernández-Yépez, 1950) VA BO 

Bryconops humeralis Machado-Allison, Chernoff & Buckup, 

1996 

VA BO 

Bryconops imitator Chernoff & Machado-Allison, 2002 VA BO 

Bryconops inpai Knöppel, Junk & Géry, 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 Géry & 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 Müller & Troschel, 1844 GU 

Gymnocorymbus thayeri Eigenmann, 1908 CG VA BO DA GU 

Gymnotichthys hildae Fernández-Yépez, 1950 VA BO 

Hemibrycon surinamensis Géry, 1962 SU FG 

Hemigrammus aereus Géry, 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 Géry, 1959 SU FG 

Hemigrammus cylindricus Durbin, 1909 GU 

Hemigrammus erythrozonus Durbin, 1909 GU 

Hemigrammus guyanensis Géry, 1959 FG

NUMBER 17 31 

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 Böhlke, 1955 VA BO 

Hemigrammus newboldi (Fernández-Yépez, 1949) VA BO DA 

Hemigrammus ocellifer (Steindachner, 1882) GU SU FG 

Hemigrammus ora Zarske, Le Bail & Géry, 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 Fernández-Yépez, 1950 VA BO 

Hyphessobrycon borealis Zarske, Le Bail & Géry, 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 Géry, 1961 SU 

Hyphessobrycon hildae Fernández-Yépez, 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 (Géry, 1960) FG 

Hyphessobrycon simulatus (Géry, 1960) FG 

Hyphessobrycon sweglesi (Géry, 1961) CG VA BO 

Hyphessobrycon takasei Géry, 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 (Géry, Planquette & Le Bail, 1996) FG 

Jupiaba maroniensis (Géry, Planquette & Le Bail, 1996) FG 

Jupiaba meunieri (Géry, Planquette & Le Bail, 1996) SU FG 

Jupiaba mucronata (Eigenmann, 1909) GU 

Jupiaba ocellata (Géry, Planquette & Le Bail, 1996) AP? FG 

Jupiaba pinnata (Eigenmann, 1909) GU SU 

Jupiaba polylepis (Günther, 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 Böhlke, 1953 VA BO PA


Microschemobrycon casiquiare Böhlke, 1953 VA BA 

Microschemobrycon melanotus (Eigenmann, 1912) GU 

Microschemobrycon meyburgi Meinken, 1975 RO 

Moenkhausia browni Eigenmann, 1909 GU 

Moenkhausia chrysargyrea (Günther, 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 Géry, 1965 SU FG 

Moenkhausia grandisquamis (Müller & Troschel, 1845) VA BO DA GU SU FG 

Moenkhausia hemigrammoides Géry, 1965 SU FG 

Moenkhausia inrai Géry, 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 Géry, Planquette & Le Bail, 1995 FG 

Moenkhausia oligolepis (Günther, 1864) CG VA BO DA GU SU FG 

Moenkhausia rara Zarske, Géry & Isbrücker, 2004 PA SU FG 

Moenkhausia shideleri Eigenmann, 1909 GU 

Moenkhausia surinamensis Géry, 1965 AP SU FG 

Paracheirodon axelrodi (Schultz, 1956) CG VA BO BA 

Paracheirodon simulans (Géry, 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 Günther, 1864 SU 

Serrabrycon magoi Vari, 1986 CG VA BO BA 

Thayeria ifati Géry, 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 (Günther, 1864) VA BO 

Triportheus venezuelensis Malabarba, 2004 BO 

Xenagoniates bondi Myers, 1942 

Subfamily: Agoniatinae—Angela M. Zanata 

CG VA BO DA 

Agoniates anchovia Eigenmann, 1914 PA 

Agoniates halecinus Müller & Troschel, 1845 

Subfamily: Iguanodectinae—Cristiano R. Moreira 

VA BO RO PA AP? GU 

Iguanodectes adujai Géry, 1970 VA BA 

Iguanodectes geisleri Géry, 1970 VA BA 

Iguanodectes spilurus (Günther, 1864) CG VA BO DA BA RO AP GU 

Piabucus dentatus (Koelreuter, 1763) 

Subfamily: Bryconinae—Flávio C. T. Lima 

VA DA GU SU FG 

Brycon amazonicus (Spix & Agassiz, 1829) CG VA BO GU

NUMBER 17 33 

Brycon bicolor Pellegrin, 1909 VA BO DA 

Brycon coquenani Steindachner, 1915 VA BO 

Brycon falcatus Müller & Troschel, 1844 CG VA BO RO PA GU SU FG 

Brycon pesu Müller & Troschel, 1845 

Subfamily: Serrasalminae—Michel Jegú 

CG VA BO DA RO PA AP GU SU FG 

Acnodon oligacanthus (Müller & Trochel, 1844) GU SU FG 

Acnodon senai Jégu & 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 (Müller & 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 Jégu, Santos & Ferreira, 1989 BA PA AP 

Mylesinus schomburgkii Valenciennes, 1850 GU 

Myleus knerii (Steindachner, 1881) FG 

Myleus setiger Müller & Troschel, 1844 CG VA BO BA RO PA GU SU 

Myloplus asterias (Müller & Troschel, 1844) VA BO PA AP GU 

Myloplus lobatus (Valenciennes, 1850) PA 

Myloplus planquettei Jégu, Keith & Le Bail, 2003 GU SU FG 

Myloplus rhomboidalis (Cuvier, 1818) VA RO PA AP GU SU FG 

Myloplus rubripinnis (Müller & 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, Jégu & Santos, 2000 BA PA 

Serrasalmus altuvei Ramírez, 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 (Fernández-Yépez & Ramírez, 1967) CG VA BA AP 

Serrasalmus medinai Ramírez, 1965 BO DA


Serrasalmus nalseni Fernández-Yépez, 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 Jégu, Keith & Belmont-Jégu, 2002 SU FG 

Tometes makue Jégu, Santos & Belmont-Jégu, 2002 CG VA BO BA 

Tometes trilobatus Valenciennes, 1850 

Subfamily: Aphyocharacinae—Rosana S. Lima 

AP FG 

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—Naércio 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 (Günther, 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 

Subfamily: Stethaprioninae—Roberto E. Reis 

GU SU FG 

Brachychalcinus orbicularis (Valenciennes, 1850) GU SU 

Poptella brevispina Reis, 1989 RO PA GU SU 

Poptella compressa (Günther, 1864) CG VA BO DA GU 

Poptella longipinnis (Popta, 1901) 

Subfamily: Tetragonopterinae—Roberto E. Reis 

CG VA BO DA SU 

Tetragonopterus chalceus Spix & Agassiz, 1829 CG VA BO DA GU SU FG

NUMBER 17 35 

Tetragonopterus lemniscatus Benine, Pelição & Vari, 2004 

Subfamily: Cheirodontinae—Luiz R. Malabarba 

SU 

Cheirodontops geayi Schultz, 1944 VA BO DA 

Odontostilbe gracilis (Géry, 1960) FG 

Odontostilbe littoris (Géry, 1960) FG 

Odontostilbe pulchra (Gill, 1858) VA BO DA RO 

Odontostilbe splendida Bührnheim & Malabarba, 2007 

Subfamily: Glandulocaudinae—Stanley H. Weitzman 

BO 

Ptychocharax rhyacophila Weitzman, Fink, Machado-Allison & 

Royero, 1994 

VA 

Family: Acestrorhynchidae—Naércio A. Menezes 

Acestrorhynchus falcatus (Bloch, 1794) VA GU SU FG 

Acestrorhynchus falcirostris (Cuvier, 1819) CG VA BA GU 

Acestrorhynchus grandoculis Menezes & Géry, 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 

Family: Cynodontidae—Mônica Toledo-Piza 

VA GU 

Cynodon gibbus Spix & Agassiz, 1829 CG BO DA GU 

Cynodon meionactis Géry, 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) 

Family: Erythrinidae—Based on Cloffsca account (Osvaldo O. 

Oyakawa) 

RO GU 

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) 

Family: Lebiasinidae—Marilyn Weitzman & Stanley H. 

Weitzman 

FG 

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 Géry & Zarske, 2002 GU 

Lebiasina provenzanoi Ardila Rodríguez, 1999 BO 

Lebiasina taphorni Ardila Rodríguez, 2004 BO 

Lebiasina uruyensis Fernández-Yépez, 1967 BO 

Lebiasina yuruaniensis Ardila Rodríguez, 2000 BO


Nannostomus anduzei Fernandez & Weitzman, 1987 VA BO 

Nannostomus beckfordi Günther, 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 Günther, 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 

Family: Ctenoluciidae—Richard P. Vari 

GU SU 

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 Günther, 1863 VA BA GU SU FG 

Helogenes uruyensis Fernández-Yépez, 1967 

Family: Aspredinidae—John P. Friel 

BO 

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

NUMBER 17 37 

Bunocephalus chamaizelus Eigenmann, 1912 GU 

Bunocephalus verrucosus (Walbaum, 1792) GU SU 

Ernstichthys anduzei Fernández-Yépez, 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) 

Family: Trichomycteridae—Wolmar Wosiacki 

VA BO 

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 

Family: Callichthyidae—Roberto E. Reis 

VA BO GU 

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 & Isbrücker, 1983 FG 

Corydoras axelrodi Rössel, 1962 CG VA BO


Corydoras baderi Geisler, 1969 PA SU 

Corydoras bicolor Nijssen & Isbrücker, 1967 SU 

Corydoras blochi Nijssen, 1971 VA BO RO GU 

Corydoras boehlkei Nijssen & Isbrücker, 1982 VA BO 

Corydoras boesemani Nijssen & Isbrücker, 1967 SU 

Corydoras bondi Gosline, 1940 VA BO GU SU 

Corydoras breei Isbrücker & Nijssen, 1992 SU 

Corydoras brevirostris Fraser-Brunner, 1947 VA BO SU 

Corydoras concolor Weitzman, 1961 VA BO 

Corydoras condiscipulus Nijssen & Isbrücker, 1980 AP FG 

Corydoras coppenamensis Nijssen, 1970 SU 

Corydoras delphax Nijssen & Isbrücker, 1983 CG VA 

Corydoras ephippifer Nijssen, 1972 AP 

Corydoras filamentosus Nijssen & Isbrücker, 1983 SU 

Corydoras geoffroy La Cepède, 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 Lönnberg & Rendahl, 1930 CG VA? BO? 

Corydoras metae Eigenmann, 1914 CG VA BO 

Corydoras nanus Nijssen & Isbrücker, 1967 SU FG 

Corydoras oiapoquensis Nijssen, 1972 AP FG 

Corydoras osteocarus Böhlke, 1951 VA BO SU 

Corydoras oxyrhynchus Nijssen & Isbrücker, 1967 SU 

Corydoras potaroensis Myers, 1927 GU 

Corydoras punctatus (Bloch, 1794) SU FG 

Corydoras sanchesi Nijssen & Isbrücker, 1967 SU 

Corydoras saramaccensis Nijssen, 1970 SU 

Corydoras septentrionalis Gosline, 1940 CG? VA BO 

Corydoras sipaliwini Hoedeman, 1965 GU SU 

Corydoras solox Nijssen & Isbrücker, 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 (Müller & Troschel, 1848) VA BO DA BA GU 

Megalechis thoracata (Valenciennes, 1840) 

Family: Loricariidae 

Subfamily: Lithogeninae—Carl J. Ferraris, Jr. 

VA BO GU SU FG 

Lithogenes villosus Eigenmann, 1909 GU 

Lithogenes wahari Schaefer & Provenzano, 2008 

Subfamily: Hypoptopomatinae—Scott A. Schaefer 

VA 

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

NUMBER 17 39 

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 

Subfamily: Loricariinae—Carl J. Ferraris, Jr. 

VA 

Cteniloricaria platystoma (Günther, 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 Isbrücker & Nijssen, 1984 PA 

Hemiloricaria eigenmanni (Pellegrin, 1908) VA BO 

Hemiloricaria fallax (Steindachner, 1915) RO GU 

Hemiloricaria formosa Isbrücker & Njissen, 1979 CG VA BO DA 

Hemiloricaria platyura (Müller & 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 Isbrücker, 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 Isbrücker, 1975 SU FG 

Pseudoloricaria laeviuscula (Valenciennes, 1840) RO 

Reganella depressa (Kner, 1853) RO 

Sturisoma monopelte Fowler, 1914 

Subfamily: Hypostominae—Jonathan W. Armbruster & Claude 

Weber 

GU 

Acanthicus hystrix Spix & Agassiz, 1829 VA BO DA 

Ancistrus hoplogenys (Günther, 1864) GU


Ancistrus leucostictus (Günther, 1864) GU SU? FG? 

Ancistrus lithurgicus Eigenmann, 1912 GU 

Ancistrus macrophthalmus (Pellegrin, 1912) VA BO 

Ancistrus nudiceps (Müller & 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 Isbrücker & 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 Isbrücker & 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

NUMBER 17 41 

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 (Lütken, 1874) VA BO 

Hypostomus watwata Hancock, 1828 DA GU SU FG 

Lasiancistrus schomburgkii (Günther, 1864) GU 

Lasiancistrus tentaculatus Armbruster, 2005 VA BO 

Leporacanthicus galaxias Isbrücker & Nijssen, 1989 VA BO 

Leporacanthicus triactis Isbrücker, Nijssen & Nico, 1992 CG VA BO 

Lithoxus boujardi Muller & Isbrücker, 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 & Isbrücker, 1990 FG 

Lithoxus surinamensis Boeseman, 1982 SU 

Neblinichthys pilosus Ferraris, Isbrücker & 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 (Günther, 1868) SU 

Pseudacanthicus leopardus (Fowler, 1914) GU 

Pseudacanthicus serratus (Valenciennes, 1840) SU FG 

Pseudancistrus barbatus (Valenciennes, 1840) GU SU FG 

Pseudancistrus brevispinis (Heitmans, Nijssen & Isbrücker, 1983) SU FG 

Pseudancistrus coquenani (Steindachner, 1915) BO 

Pseudancistrus corantijniensis De Chambrier & Montoya- 

Burgos, 2008 

SU 

Pseudancistrus depressus (Günther, 1868) SU 

Pseudancistrus guentheri (Regan, 1904) GU 

Pseudancistrus longispinis (Heitmans, Nijssen & Isbrücker, 1983) FG 

Pseudancistrus macrops (Lütken, 1874) SU 

Pseudancistrus megacephalus (Günther, 1868) GU SU 

Pseudancistrus niger (Norman, 1926) FG 

Pseudancistrus nigrescens Eigenmann, 1912 GU 

Pseudancistrus orinoco Isbrücker, Nijssen & Cala, 1988 CG VA BO 

Pseudancistrus pectegenitor Lujan, Armbruster & Sabaj, 2007 VA 

Pseudancistrus reus Armbruster & Taphorn, 2008 BO


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) 

Family: Pseudopimelodidae—Oscar A. Shibbata 

VA BO GU? 

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 (Flávio 

A. Bockmann & Gizelani M. Guazzeli) 

Brachyglanis frenata Eigenmann, 1912 VA BO BA GU 

Brachyglanis magoi Fernández-Yépez, 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 (Müller & 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

NUMBER 17 43 

Phenacorhamdia provenzanoi DoNascimiento & Milani, 2008 VA BO 

Pimelodella altipinnis (Steindachner, 1864) GU 

Pimelodella cristata (Müller & Troschel, 1848) GU FG 

Pimelodella cruxenti Fernández-Yépez, 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 (Müller & Troschel, 1848) RO PA GU 

Rhamdia laukidi Bleeker, 1858 CG VA BO GU 

Rhamdia muelleri (Günther, 1864) VA BO DA PA GU 

Rhamdia quelen (Quoy & Gaimard, 1824) 

Family: Pimelodidae—John G. Lundberg 

CG VA BO DA GU SU FG 

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 (Müller & 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-Suárez & Burr, 2007 BO DA 

Pseudoplatystoma orinocoense Buitrago-Suárez & Burr, 2007 VA BO DA 

Sorubim elongatus Littmann, Burr, Schmidt & Isern, 2001 CG VA BO GU


Sorubim lima (Bloch & Schneider, 1801) CG VA BO DA 

Sorubimichthys planiceps (Spix & Agassiz, 1829) CG VA BO DA 

Zungaro zungaro (Humboldt, 1821) 

Family: Doradidae—Mark H. Sabaj Pérez 

CG VA BO DA BA GU 

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 (Fernández-Yépez, 1968) CG VA BO DA? BA? 

Amblydoras gonzalezi (Fernández-Yépez, 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 Pérez & Birindelli, 2008 RO 

Hassar orestis (Steindachner, 1875) CG VA BO DA BA RO? PA? GU 

Leptodoras cataniai Sabaj, 2005 VA BA RO? 

Leptodoras copei (Fernández-Yépez, 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 (Fernández-Yépez, 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 Fernández-Yépez, 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 (Fernández-Yépez, 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) 

Family: Auchenipteridae—Carl J. Ferraris, Jr. 

CG VA BO DA? BA RO PA GU 

Ageneiosus inermis (Linnaeus, 1766) CG VA BO DA GU SU FG 

Ageneiosus marmoratus Eigenmann, 1912 GU SU 

Ageneiosus piperatus (Eigenmann, 1912) RO GU

NUMBER 17 45 

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 

Family: Sternopygidae—James S. Albert 

VA BO DA 

Archolaemus blax Korringa, 1970 RO AP? FG 

Distocyclus conirostris (Eigenmann & Allen, 1942) BO DA


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 (Fernández-Yépez, 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) 

Family: Rhamphichthyidae—Carl J. Ferraris, Jr. 

CG VA BO DA BA? RO? PA? AP? GU SU FG 

Gymnorhamphichthys hypostomus Ellis, 1912 CG VA DA 

Gymnorhamphichthys rondoni (Miranda Ribeiro, 1920) VA BO GU SU 

Iracema caiana Triques, 1996 BA RO 

Rhamphichthys apurensis (Fernández-Yépez, 1968) VA BO DA 

Rhamphichthys rostratus (Linnaeus, 1766) 

Family: Hypopomidae—James S. Albert 

GU SU 

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 Fernández-Yépez, 1968 CG VA BO BA 

Racenisia fimbriipinna Mago-Leccia, 1994 VA BO 

Steatogenys duidae (La Monte, 1929) VA BO 

Steatogenys elegans (Steindachner, 1880) 

Family: Apteronotidae—James S. Albert 

CG VA BO DA 

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 (Günther, 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 (Müller & Troschel, 1849) BO GU FG 

Sternarchorhynchus roseni Mago-Leccia, 1994 BO DA

NUMBER 17 47 

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-Alcalá, Taphorn, Lasso & León-Mata, 2006 BO 

Rivulus stagnatus Eigenmann, 1909 GU SU 

Rivulus tecminae Thomerson, Nico & Taphorn, 1992 VA BO 

Rivulus torrenticola Vermeulen & Isbrücker, 2000 GU 

Rivulus waimacui Eigenmann, 1909 GU 

Rivulus xiphidius Huber, 1979 AP FG 

Terranatos dolichopterus (Weitzman & Wourms, 1967) 

Family: Poeciliidae—Paulo H. J. Lucinda 

VA BO 

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


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 (Günther, 1866) 

Family: Hemiramphidae—Bruce B. Collette 

VA BO DA GU SU 

Hyporhamphus brederi (Fernández-Yépez, 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 (Lacpède, 1802) VA BO DA AP GU SU FG 

Pachypops pigmaeus Casatti, 2002 RO 

Pachypops trifilis (Müller & Troschel, 1849) BA RO GU SU 

Pachyurus gabrielensis Casatti, 2001 VA BO BA 

Pachyurus schomburgkii Günther, 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) 

Family: Polycentridae—Richard P. Vari 

CG DA SU FG 

Monocirrhus polyacanthus Heckel, 1840 BO RO 

Polycentrus schomburgkii Müller & Troschel, 1849 

Family: Cichlidae—Sven O. Kullander 

DA AP GU SU FG 

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

NUMBER 17 49 

Apistogramma velifera Staeck, 2003 VA 

Apistogramma viejita Kullander, 1979 CG 

Apistogramma wapisana Römer, 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 (Günther, 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, Jégu & Ferreira, 1991 PA 

Crenicichla vaillanti Pellegrin, 1903 GU FG 

Crenicichla virgatula Ploeg, 1991 RO 

Crenicichla wallacii Regan, 1905 GU 

Crenicichla zebrina Montaña, López-Fernández & Taphorn, 

2008 

VA 

Dicrossus filamentosus (Ladiges, 1958) CG VA BO 

Dicrossus gladicauda Schindler & Staeck, 2008 CG 

Geophagus abalios López-Fernández & 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 López-Fernández & Taphorn, 2004 VA BO


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 López-Fernández & Taphorn, 2004 VA 

Guianacara (Guianacara) cuyunii López-Fernández, 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 López-Fernández, 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 (Günther, 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 (Müller & Troschel, 1849) GU SU 

Satanoperca lilith Kullander & Ferreira, 1988 BA RO PA 

Satanoperca mapiritensis (Fernández-Yépez, 1950) VA BO DA 

Uaru fernandezyepezi Stawikowski, 1989 

Family: Gobiidae—Sven O. Kullander 

VA 

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 Cepède, 1800 DA FG 

Gobioides grahamae (Palmer & Wheeler, 1995) GU SU FG

NUMBER 17 51 

Gobiomorus dormitor La Cepède, 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 Cervigón, 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 (Günther, 1862) VA BO BA RO AP GU 

Order: Tetraodontiformes 

Family: Tetraodontidae—Carl J. Ferraris, Jr. & Sven O. 

Kullander 

Colomesus asellus (Müller & Troschel, 1849) BO DA GU 

Order: Lepidosireniformes 

Family: Lepidosirenidae—Based on Cloffsca account (Gloria 

Arratia) 

Lepidosiren paradoxa Fitzinger, 1837 CG? VA? BA? FG

PHOTOGRAPHIC ATLAS OF FISHES OF THE GUIANA SHIELD 

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 Bolívar 

(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 Pérez 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 ichthyological 

studies in decades to come. 

Scope 

The plates present 130 individuals representing 127 

species of 46 families. Fishes were collected in Guyana 

MARK H. SABAJ PÉREZ 

(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., Rhinosardinia 

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 highquality 

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) or 

in 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-immersion 

remains 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.


Figure 3. Author using phototank-immersion method to photograph fish streamside in Mongolia. Photo by C. Sabaj Pérez. 

The techniques described below follow a minimalist 

approach with some advice limited to the specific 

cameras and conditions involved. For a more sophisticated 

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 oneeighth 

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 compromise 

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 oneeighth 

(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

NUMBER 17 55 

Figure 4. Author photographing fish streamside in Guyana. Photo by J.W. Armbruster. 

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 3/16 th 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 illuminated 

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 5. Kyle Luckenbill photographing small alcohol-preserved specimen (above ruler) while holding polarizing filter in lab. Photo 

by author. 

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 carefully 

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 compressed 

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 achieving 

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 transparent 

fins. Black pigment in fin membranes or along 

distal fin margins disappears against dark backgrounds. 

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, particularly 

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 background 

deviates sharply from the original (i.e., black to

NUMBER 17 57 

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 horizontal 

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 apertures 

(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 compressed 

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 background. 

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 magnification 

(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


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 finetuning. 

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. 


Numerous colleagues contributed valuable assistance 

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. León-Mata, Nathan Lujan, John 

Lundberg, Jan Mol, Matthew Thomas, David Werneke, 

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, 

Raphaël Covain and Juan Montoya Burgos for 

comments on the introduction, and to Jonathan 

Armbruster, Hernán López-Fernández, 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). 


Alonso, L. E., & H. J. Berrenstein (eds.). 2006. A rapid biological 

assessment of the aquatic ecosystems of the Coppename River 

Basin, Suriname. RAP Bulletin of Biological Assessment 39. 

Conservation International, Washington, D.C., 119 pp. 

Armbruster, J. W., N. K. Lujan, & D. C. Taphorn. 2007. Four new 

Hypancistrus (Siluriformes: Loricariidae) from Amazonas, 

Venezuela.—Copeia 2007(1):62–79. 

———, M. H. Sabaj, M. Hardman, L. M. Page, & J. H. Knouft. 

2000. Catfish genus Corymbophanes (Loricariidae: Hypostominae) 

with description of one new species: Corymbophanes 

kaiei.—Copeia 2000(4):997–1006. 

Berrenstein, H. J. 2005. Field guide to the freshwater fishes of the 

Central Suriname Nature Reserve (CSNR), (Coppename 

River Basin, Suriname). Conservation International Suriname, 

Paramaribo, 96 pp. 

Cardoso, Y. P., & J. I. Montoya-Burgos. 2009. Unexpected diversity 

in the catfish Pseudancistrus brevispinis reveals dispersal routes 

in a Neotropical center of endemism: the Guyanas Region.— 

Molecular Ecology 18(5):947–964. 

de Chambrier, S., & J. I. Montoya-Burgos. 2008. Pseudancistrus 

corantijniensis, a new species from the Guyana Shield 

(Siluriformes: Loricariidae) with a molecular and morphological 

description of the Pseudancistrus barbatus group.— 

Zootaxa 1918:45–58. 

Hardman, M., L. M. Page, M. H. Sabaj, J. W. Armbruster, & J. H. 

Knouft. 2002. A comparison of fish surveys made in 1908 and 

1998 of the Potaro, Essequibo, Demerara, and coastal river 

drainages of Guyana.—Ichthyological Exploration of Freshwaters 

13:225–238. 

Jenkins, R. E., & N. M. Burkhead. 1994. Freshwater fishes of 

Virginia. American Fisheries Society, Bethesda, Maryland, 

1080 pp. 

Lasso, C. A., J. C. Señarìs, L. E. Alonso, & A. L. Flores (eds.). 2006. 

Evaluación rápida de la biodiversidad de los ecosistemas 

acuáticos en la confluencia de los ríos Orinoco y Ventuari, 

Estado Amazonas (Venezuela). Boletín RAP de Evaluación 

Biológica 30. Conservation International, Washington, D.C., 

240 pp. 

Lasso, C. A., J. Hernández-Acevedo, E. Alexander, J. C. Señaris, L. 

Mesa, H. Samudio, J. Mora-Day, C. Magalhaes, A. Shushu, 

E. Mauruwanaru, & R. Shoni. 2008. Aquatic biota: fishes, 

decapod crustaceans and mollusks of the upper Essequibo 

Basin (Konashen COCA), Southern Guyana. Pp. 43–54 in 

L. E. Alonso, J. McCullough, P. Naskrecki, E. Alexander, & 

H. E. Wright, eds., A rapid biological assessment of the 

Konashen Community-Owned Conservation Area, Southern 

Guyana. RAP Bulletin of Biological Assessment 51, Conservation 

International, Washington, D.C. 

Lord, C., Y. Fermon, F. J. Meunier, M. Jégu, & P. Keith. 2007. 

Croissance et longévité du Watau yaike, Tometes lebaili

NUMBER 17 59 

(Serrasalminae), dans le bassin du Haut Maroni (Guyane 

française). Résultats préliminaires.—Cybium, 31(3):359–367. 

Lujan, N. K., M. Arce, & J. W. Armbruster. 2009. A new black 

Baryancistrus with blue sheen from the upper Orinoco 

(Siluriformes: Loricariidae).—Copeia 2009(1):50–56. 

Machado-Allison, A., B. Chernoff, F. Provenzano, P. W. Willink, A. 

Marcano, P. Petry, B. Sidlauskas, & T. Jones. 2003. 

Inventory, relative abundance and importance of fishes in 

the Caura River basin, Bolívar State, Venezuela. Pp. 64–74 in 

B. Chernoff, A. Machado-Allison, K. Riseng, & J. R. 

Montambault, eds., A biological assessment of the aquatic 

Ecosystems of the Caura River Basin, Bolívar State, 

Venezuela. RAP Bulletin of Biological Assessment 28. 

Conservation International, Washington D.C. 284 pp. 

Planquette, P., P. Keith, & P.-Y. Le Bail. 1996. Atlas des poissons 

d’eau douce de Guyane. Volume 1, Collection du Patrimoine 

Naturel, vol. 22. Institut d’Ecologie et de Gestion de la 

Biodiversité du Muséum National d’Histoire Naturelle, 

Institut national de la Recherche Agronomique, Conseil 

Supérior de la Pêche, Paris, 429 pp. 

Sabaj Pérez, M. H, J. W. Armbruster, C. J. Ferraris, Jr., J. P. Friel, J. 

G. Lundberg, & L. M. Page (eds.). 2009. All Catfish Species 

Inventory. Internet address: http://silurus.acnatsci.org. 

Taphorn, B. D. C, H. López-Fernández, & C. R. Bernard. 2008. 



Zootaxa 1925:31–38. 

Watkins, G., W. Saul, E. Holm, C. Watson, D. Arjoon, & J. Bicknell. 

2005. The fish fauna of the Iwokrama Forest.—Proceedings of 

the Academy of Natural Sciences of Philadelphia 154(1):39–53. 

Werneke, D. C., M. H. Sabaj, N. K. Lujan, & J. W. Armbruster. 2005. 

Baryancistrus demantoides and Hemiancistrus subviridis,twonew 

uniquely colored species of catfishes from Venezuela (Siluriformes: 

Loricariidae).—Neotropical Ichthyology 3(4):533–542.

APPENDIX: PLATES


Potamotrygonidae 

Plate 1 

A. Paratrygon aiereba (live). AUM 43646 (154 mm maximum disk width). Venezuela, 

Amazonas, Río 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, Río 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, Río 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.

NUMBER 17 63


Arapaimidae 

Plate 2 

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.

NUMBER 17 65


Clupeidae 

Plate 3 

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, Río Casiquiare at 

mouth of Caño 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 km 

S-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 village 

of Jawalla, 05u41935.40N, 060u28911.80 W, 18 Apr 2008, H. López-Fernández, D. 

Taphorn, E. Liverpool, K. Kramer, & C. Thierens. Photo by D. C. Taphorn Baechle & H. 

López-Fernández. 

H. Apareiodon orinocensis (live). ANSP 185045. Venezuela, Amazonas, Río 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.

NUMBER 17 67


Curimatidae 

Plate 4 

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, 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. 

Crenuchidae 

E. Leptocharacidium omospilus (live). ANSP 189272 (60.5 mm SL). Venezuela, Amazonas, 1 

km upstream from mouth of left bank tributary of Río Siapa, mouth below Salto Oso and 

above Salto Sardinas on Río 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.

NUMBER 17 69


Anostomidae 

Plate 5 

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), 


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, Río 

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 at 

Kassi-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, Litanie 

River, 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.

NUMBER 17 71


Characidae 

Plate 6 

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, 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. 

C. Chalceus epakros (live). AUM 43073 (75 mm SL). Venezuela, Amazonas, Río Orinoco, 

inlet between 2 islands in channel, 84.5 km N of San Fernando de Atabapo, 9 km Sof 

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 



K. Wan. 

F. Myleus schomburgkii (live). ANSP 180812 (160 mm SL). Venezuela, Amazonas, Río 

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, Río 

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.

NUMBER 17 73


Acestrorhynchidae 

Plate 7 

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. 


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 of 

Konya 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, 


Lebiasinidae 

E. Copella compta (live). AUM 41327 (37 mm SL). Venezuela, Amazonas, Caño Carmen, 

tributary of Río 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, Río Ventuari 

(Orinoco drainage), large rock outcrop, 97 km NE of Macuruco, 163 km E-NE of San Fernando 

de Atabapo, 04u259100N, 066u179080W,7Apr2004,M.Sabaj,N.Lujan,D.Wernekeetal.

NUMBER 17 75


Cetopsidae 

Plate 8 

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, 


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 Linden 

bearing 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 fish 

market, 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, Río 

Casiquiare, bedrock riffle and outcrop, 74.6 km NE of San Carlos de Rio Negro, 


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 NW 

of 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.

NUMBER 17 77


Doradidae 

Plate 9 

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, Río 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, Río 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, Río Ventuari 

(Orinoco drainage), village of Marueta at landing, 91 km 

E-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, Río 

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, 


G. Doras micropoeus (alcohol). ANSP 187110 (225 mm SL). Suriname, Sipaliwini, Lawa 


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, Río 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, Río Orinoco 

near confluence with Río 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.

NUMBER 17 79


Pseudopimelodidae 

Plate 10 

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, Río 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 


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, 

Río Casiquiare, bedrock outcrop 59.5 km SW of La Esmerelda, 02u499070N, 


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.

NUMBER 17 81


Callichthyidae 

Plate 11 

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 Oct 

2002, J. Armbruster, D. Werneke, C. Allison et al. 

Trichomycteridae 

C. Ituglanis cf. metae (live). AUM 43074 (61 mm SL). Venezuela, Amazonas, Río 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 et 

al. 

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, 


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. 


J. Sarcoglanis simplex (live). ANSP 179212 (17 mm SL). Same locality as G.

NUMBER 17 83


Loricariidae 

Plate 12 

A. Leporacanthicus triactis (live). AUM 39243 (116 mm SL). Venezuela, Amazonas, Río 

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, Río 

Orinoco at Cucue Amerindian village, opposite Trapichote village, 60 km E of San 

Fernando 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 



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 Konya 

Kondre, 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, Río 

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, Río Casiquiare, bedrock in stream, 73 km NE of San Carlos de Río 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 



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.

NUMBER 17 85


Gymnotidae 

Plate 13 

A. Electrophorus electricus (live). MCNG 51982. Venezuela, Amazonas, Río 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 


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, Lawa 



K. Wan. 

Apteronotidae 

E. Apteronotus albifrons (live). AUM 40678 (109 mm SL). Venezuela, Amazonas, Río 

Manapiare (Ventuari-Orinoco drainage), at San Juan de Manapiare landing, 15 Apr 2004, 

J. Para.

NUMBER 17 87


Rivulidae 

Plate 14 

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, Río 

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 Oct 

2002, M. Sabaj, J. Armbruster, M. Thomas et al.

NUMBER 17 89


Cichlidae 

Plate 15 

A. Crenicichla multispinosa (live). ANSP 187101 (223 mm SL). Suriname, Sipaliwini, Lawa 



K. Wan. 

B. Geophagus harreri (live). ANSP 187136 (170 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. 

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, Río 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, Río Orinoco near mouth 

of Río 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, Río Orinoco, bedrock 

outcrop, 52.9 km SE of San Antonio, 102 km W of La Esmerelda, 03u069010N, 


G. Pterophyllum altum (live). AUM 40584 (59 mm SL). Same locality as E.

NUMBER 17 91


Synbranchidae 

Plate 16 

A. Synbranchus marmoratus (live). ANSP 187334 (492 mm TL). Suriname, Sipaliwini, Lawa 



K. Wan. 

B. Synbranchus marmoratus (live, ventral view). Same data as A. 

Sciaenidae 

C. Pachyurus schomburgkii (alcohol). ANSP 162800 (195 mm SL). Venezuela, Amazonas, 

Río Iguapo approximately 1 hour above its confluence with Río 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, Río 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. 

Achiridae 

F. Hypoclinemus mentalis (alcohol). ANSP 163857 (80.5 mm SL). Venezuela, Amazonas, 

caño of Río 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.

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 

Perciformes 

Pimelodidae 

Pleuronectiformes 

Poeciliidae 

Polycentridae 

Potamotrygonidae 

Pristidae 

Pristiformes 

Pristigasteridae 

Prochilodontidae 

Pseudopimelodidae 

Rhamphichthyidae 

Rivulidae 

Sciaenidae 

Serrasalminae 

Siluriformes 

Sternopygidae 

Stethaprioninae 

Synbranchidae 

Synbranchiformes 

Tetragonopterinae 

Tetraodontidae 

Tetraodontiformes 

Trichomycteridae

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