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10 September 2009 NUMBER 17

checklist of freshwater fishes of the guiana shield

BULLETIN of the

Biological Society

of Washington

ISSN 0097-0298

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CHECKLIST OF THE FRESHWATER FISHES OF THE

GUIANA SHIELD

Richard P. Vari, Carl J. Ferraris, Jr., Aleksandar Radosavljevic, and Vicki A. Funk

i

Front cover illustration: Pseudolithoxus dumus, family Loricariidae (see Plate 12, Figure G).

Illustrations facing each section:

For the Introduction, montage of radiographs of fishes from the rivers of the Guiana Shield, upper left –

Rhaphiodon vulpinis, Cynodontidae; upper right – Prochilodus mariae, Prochilodontidae; center – Serrasalmus

irritans, Characidae, Serrasalminae; lower left – Corydoras filamentosus, Callichthyidae; and lower right,Sternarchorhynchus roseni, Apteronotidae, mature male with enlarged dentary dentition. Images and plate

prepared by S. Raredon.

For the Fishes of the Guiana Shield, Acnodon oligocanthus (Serrasalminae, juvenile) from Steindachner, F. 1915.

Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen

Classe, Wien 93:15–105 (date based on release of separates of main work published in 1917).

For the Guide to the Checklist, Acroronia nassa (Cichlidae) from Steindachner, F. 1875. Sitzungsberichte der

Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen Classe, Wien 71:61–137.

For the Photographic Atlas of Fishes of the Guiana Shield, Leptodoras hasemani (Doradidae) from Steindachner,

F. 1915. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftli-

chen Classe, Wien 93:15–105 (date based on release of separates of main work published in 1917).

Preferred citations:

Vari, R. P., C. J. Ferraris, Jr., A. Radosavljevic, & V. A. Funk, eds., 2009. Checklist of the freshwater fishes of the

Guiana Shield.—Bulletin of the Biological Society of Washington, no. 17.

or, e.g.,

Vari, R. P., & C. J. Ferraris, Jr. 2009. Fishes of the Guiana Shield. In Vari, R. P., C. J. Ferraris, Jr., A.

Radosavljevic, & V. A. Funk, eds., 2009. Checklist of the freshwater fishes of the Guiana Shield.—Bulletin ofthe Biological Society of Washington, no. 17.

ii

CONTENTS

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

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

INTRODUCTION. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Vicki A. Funk and Carol L. Kelloff

FISHES OF THE GUIANA SHIELD. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

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

GUIDE TO THE CHECKLIST . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Aleksandar Radosavljevic

CHECKLIST. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25Pristiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

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

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

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

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

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

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

Gymnotiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45Cyprinodontiformes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47

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

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

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

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

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

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

PHOTOGRAPHIC ATLAS OF FISHES OF THE GUIANA SHIELD. . . . . . . . . . . . . . . . . . . . . . 53

Mark Sabaj Perez

INTRODUCTION

APPENDIX: PLATES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

INDEX TO ORDERS, FAMILIES, AND SUBFAMILIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

iii

CONTRIBUTORS

James S. Albert, Department of Biology, University of Louisiana at Lafayette, Lafayette, Louisiana 70504-2451,

U.S.A., e-mail: [email protected]

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


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

Janeiro, RJ, Brazil, e-mail: [email protected]

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

Pasteur, 458/sala 408, Urca - Rio de Janeiro, RJ, 22290-240, Brazil, e-mail: [email protected]

Marcelo R. de Carvalho, Instituto de Biociencias, Universidade de Sao Paulo, Rua do Matao, Trav. 14, no. 101,

Sao Paulo, SP, 05508-900, Brazil, e-mail: [email protected]

Lilian Cassati, Laboratorio de Ictiologia, Departamento de Zoologia e Botanica, IBILCE-UNESP, Rua Cristovao

Colombo, 2265, Sao Jose do Rio Preto, SP, 15054-000, Brazil, e-mail: [email protected]

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

e-mail: [email protected]

Bruce B. Collette, National Marine Fisheries Service, Systematics Laboratory MRC 153, National Museum of

Natural History, Washington D.C. 20560-0153, U.S.A., e-mail: [email protected]

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

Postal 68049, 21944-970 Rio de Janeiro, RJ, Brazil, e-mail: [email protected]

Fabio di Dario, Universidade Federal do Rio de Janeiro, Av. Sao Jose do Barreto, Sao Jose do Barreto, Caixa

Postal 119331, Macae, RJ, 27921-550, Brazil, e-mail: [email protected]

Carl J. Ferraris, Jr., 2944 NE Couch St., Portland, OR 97232, USA; and Division of Fishes, National Museum of

Natural History, Smithsonian Institution, Washington, D.C. 20560-0159, U.S.A., e-mail: [email protected]

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


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

Washington, D.C. 20560-0166, U.S.A., e-mail: [email protected]

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

France, e-mail: [email protected]

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


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

104 05 Stockholm, Sweden, e-mail: [email protected]

Francisco Langeani Neto, Laboratorio de Ictiologia, Departamento de Zoologia e Botanica, IBILCE-UNESP, Rua

Cristovao Colombo, 2265, Sao Jose do Rio Preto, SP, 15054-000, Brazil, e-mail: [email protected]

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

Brazil, e-mail: [email protected]

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

Francisco Portela, 794, Sao Goncalo, RJ, 24435-000, Brazil, e-mail: [email protected]

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

Porto Alegre, RS, 90619-900, Brazil, e-mail: [email protected]

iv

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

Postal 25, Porto Nacional, TO, 77500-000, Brazil, e-mail: [email protected]

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

Parkway, Philadelphia, Pennsylvania 19103, U.S.A., e-mail: [email protected]

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

Goncalves, 9500 - bloco IV - Predio 43435, Porto Alegre, RS, 91501-970, Brazil, e-mail: [email protected]

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

Station, Texas 77843-2258, U.S.A., e-mail: [email protected]

Naercio A. Menezes, Museu de Zoologia, Universidade de Sao Paulo, Caixa Postal 42594, Sao Paulo, SP, 04299-

970, Brazil, e-mail: [email protected]

Cristiano Moreira, Departamento de Ciencias Biologicas, Universidade Federal de Sao Paulo, Rua Prof. Artur

Riedel, 275, Diadema, SP, 09972-270, Brazil, e-mail: [email protected]

Carla S. Pavanelli, Fundacao Universidade Estadual de Maringa, Nupelia, Av. Colombo, 3690, Maringa, PR,

87020-900, Brazil, e-mail: [email protected]

Aleksandar Radosavljevic, The City College of New York, 526 Marshal Science Building, 160 Convent Avenue,

New York, New York 10031, U.S.A., e-mail: [email protected]

Robson T. C. Ramos, Depto de Sistematica e Ecologia, Universidade Federal da Paraıba, PB, Joao Pessoa, 58059-

900, Brazil, e-mail: [email protected]

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

Alegre, RS, 90619-900, Brazil, e-mail: [email protected]

Ricardo S. Rosa, Depto de Sistematica e Ecologia, Universidade Federal da Paraıba, Joao Pessoa, PB, 58059-900,

Brazil, e-mail: [email protected]

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

Parkway, Philadelphia, Pennsylvania 19103, U.S.A., e-mail: [email protected]

Scott A. Schaefer, Division of Vertebrate Zoology, American Museum of Natural History, Central Park West at

79th Street, New York, New York 10024-5192, U.S.A., e-mail: [email protected]

Oscar A. Shibatta, Departamento de Biologia Animal e Vegetal, Centro de Ciencias Biologicas, Universidade

Estadual de Londrina, 86051-990 Londrina, PR, Brazil, e-mail: [email protected]

Monica Toledo-Piza, Departamento de Zoologia, Instituto de Biociencias, Universidade de Sao Paulo, Caixa

Postal 11461, 05422-970 Sao Paulo, SP, Brazil, e-mail: [email protected]

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

D.C. 20560-0159, U.S.A., e-mail: [email protected]

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

1211 Geneve, Switzerland, e-mail: [email protected]

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


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


Wolmar Wosiacki, Museu Paraense Emılio Goeldi, Laboratorio de Ictiologia, Av. Magalhaes Barata 376 Caixa

Postal Box 399, Belem, PA, 66040-170, Brazil, e-mail: [email protected]

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

de Ondina, Salvador, BA, 40170-290, Brazil, e-mail: [email protected]

v

Abstract.—Distributions are provided for 1168 species of fishes that live in the

freshwater drainage systems overlying the Guiana Shield in Brazil, Colombia,

French Guiana, Guyana, and Suriname. This total includes representatives of

376 genera, 49 families, and 15 orders, with five orders (Characiformes,

Siluriformes, Perciformes, Gymnotiformes, and Cyprinodontiformes) account-

ing for 96.7% of the diversity. Drainage systems on the Guiana Shield are home

to approximately 23% of the freshwater species known from Central and SouthAmerica. A summary is provided of ichthyological collecting on the Shield along

with summaries of major publications dealing with fishes of each region on the

Shield. Factors that may account for the high species level diversity in that

region are discussed. Methods for photographing fishes are detailed, and a

photographic album of 127 species of 46 families that occur on the Shield is

included.

Key Words.—Guiana Shield Fishes, Freshwaters, Brazil, Colombia, French

Guiana, Guyana, Suriname, Venezuela.

vii

INTRODUCTION

V. A. FUNK and CAROL L. KELLOFF

In the face of the growing biodiversity crisis, we must

move to document and evaluate the biota of our

natural areas. As taxonomists, it is part of our job to

name, place into groups, and keep track of the species

that we study. Often we use the distribution of the taxa

to help with our work and to provide a broader context

for the results. In doing this job, taxonomists produce

checklists, floras and faunas, and monographs. In fact,

collecting specimens and producing these documents

are essential elements in our quest to understand the

natural world and how it evolved. Checklists are the

‘‘first pass’’ in our attempts to understand the diversity

of an area. They give us the first approximation of the

known diversity of any group or groups of organisms

and they often provide the first profile of the biodiver-

sity in relatively poorly known parts of the globe. For

many groups, they may represent the most complete

information available and since basic taxonomic data

provide the essential information for studies in sys-

tematic and evolutionary biology, we continually seek

to improve and update it. Checklists have many uses:

they are aids in the identification and correct naming of

species, they serve as essential resources for biodiver-

sity estimates and biogeographic studies, and they are

starting points for more detailed studies of an area’s

biota. When reviewed by many specialists, they often

represent the most advanced state of knowledge avail-

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

these basics they increase our knowledge of this fasci-

nating region and act as a starting point for additional

biodiversity research because they give insight into the

species richness, endemicity, and floral and faunal

affinities for the region and provide the baseline

information for analyzing species turnover rates and

migration or invasion events and many other biological

phenomena (e.g., Funk & Richardson 2002, Funk et al.

2002, 2005; Ferrier et al. 2004, Kelloff & Funk 2004).

Checklists also increase national and regional pride by

demonstrating the diversity of the area and provide

important public outreach and fundamental informa-

tion to be used by governments and conservation

organizations in addressing the biodiversity crisis. This

Checklist of the Fishes of the Guiana Shield, when

combined with the recently published Checklist of the

Terrestrial Vertebrates of the Guiana Shield (Hollowell

& Reynolds 2005), and Checklist of the Plants of the

Guiana Shield (Funk et al. 2007), provides a sound

basis for future research and conservation efforts.

This Checklist of the Fishes of the Guiana Shield is an

excellent example of effective interaction, involving

ichthyologists from around the world who contributed

in diverse ways to its completion. It began as an

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

a larger and more involved project. As with anyendeavor of this type, insights that specialists have

gained through their experience are irreplaceable in

correcting errors and updating classifications. In order

to make future editions of this checklist as current and

accurate as possible, specialists are encouraged to

contact the Smithsonian’s Biological Diversity of the

Guiana Shield Program with additions or corrections.

The Guiana Shield

The Guiana Shield region is a biologically rich area

that includes much of northeastern South America

(Fig. 1). It is strictly defined by the underlying

geological formation known as the Guiana Shield,

and in the context of this volume the term Guiana

Shield also refers to the corresponding geographic

region. That region includes the Venezuelan states ofBolıvar and Amazonas, and a portion of Delta

Amacuro; all of Guyana, Suriname, and French

Guiana; and parts of northern Brazil. Several geolog-

ical outliers of the Guiana Shield occur west of the

Orinoco River in Colombia. In Spanish and Portu-

guese speaking countries, the region is often referred to

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

Guayana, Brazilian Guayana, and VenezuelanGuayana are often used. The total area of the Guiana

Shield is approximately 1,520,000 km2. Table 1 lists the

square kilometers of political divisions that occur

within the region. See Berry et al. (1995) for a review

of definitions of and terminology related to the Guiana

Shield region.

Geology

The Guiana Shield (Fig. 1) is a distinct geologic

region that underlies Guyana, Suriname, French

Guiana, parts of Venezuela, and Brazil, and a small

area in Colombia. It is roughly bounded by the

Atlantic Ocean to the north and east, the Orinoco

River to the north and west, the Rıo Negro (a major

tributary of the Amazon River) to the southwest, and

the Amazon River to the south (Gibbs & Barron 1993).The Guiana Shield is a distinct ancient geological

formation that includes the mountain systems that

form the watershed boundary between the Amazon

and Orinoco rivers. On the Shield’s western side, the

Orinoco River and Rıo Negro are connected by the

Rıo Casiquiare, making most of the region somewhat

like an island; however, there are some areas of nearby

Colombia that have remnants of the Shield formation.The southern boundary of the Guiana Shield is

difficult to define precisely, as a broad band of outwash

materials resulting from erosion occurs between

mountains on the southern boundary of the Shield

and the Amazon and Negro rivers stretching into parts

of northern Brazil. Also, much of the Venezuelan state

of Delta Amacuro occurs over thick sedimentsdeposited primarily by the Orinoco River and is really

not strictly part of the Shield; however, some moun-

tains of the Guiana Shield occur in this state’s southern

section, and a large proportion of the sediments of the

delta are derived from outwash from the highlands of

the Shield. For more detailed discussions of the

geology of the area, readers should refer to Gibbs &

Barron (1993) and Huber (1995a).

The base of the Guiana Shield is composed of

crystalline rocks of Proterozoic origin; these are mainly

granites and gneisses formed between 3.6 and 0.8

billion years ago (Mendoza 1977, Schubert & Huber

1990). Large areas of the Shield were overlain with

sediments from 1.6 to 1 billion years ago and cemented

during thermal events (Huber 1995a). These quartzite

and sandstone rocks comprise the Roraima formation

and today remnants are scattered across the central

portion of the Shield extending west from the Potaro

Plateau of the Pakaraima Mountains in central

Guyana through parts of Venezuela and Colombia

(Arbelaez & Callejas 1999) and south into northern

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

this area, erosion has resulted in numerous vertical-

walled, frequently flat-topped mountains called ‘‘te-

puis,’’ among them are Chimantı-tepui (2550 m), Cerro

Duida (2358 m), and Auyan-tepui (2450 m). Pico de

Neblina (3014 m) is the Shield’s western-most tepui

and highest point, located on the southern-most

segment of the border between Venezuela and Brazil.

Mount Roraima (2810 m) is located at the juncture of

Guyana, Venezuela, and Brazil and includes the

highest point within Guyana. The eastern-most peaks

in Guyana reach approximately 2000 m elevation,

including Mt. Ayanganna (2041 m). Several of these

mesa-like formations are virtually inaccessible by foot

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

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

east arrangement, with abbreviations used and estimated areas.

Division Abbr. Area (km2)

*Colombian Guayana CG 120,325

Amazonas, Venezuela VA 175,750

Bolıvar, Venenzuela BO 238,000

*Delta Amacuro, Venezuela DA 40,200

*Amazonas, Brazil BA 125,550

*Roraima, Brazil RO 173,750

*Para, Brazil PA 243,280

*Amapa, Brazil AP 98,750

Guyana GU 214,970

Suriname SU 163,270

French Guiana FG 91,000

Total (km2) 1,896,845

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

the Guiana Shield region.

2 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON

and are so unusual that they inspired a fictional

scientific expedition referred to one as ‘‘The Lost

World’’ (Doyle 1912), a term sometimes applied to all

tepuis. Notable waterfalls of the region include Angel

Falls (979 m) on Auyan-tepui in Venezuela and

Kaieteur Falls (226 m), which flows year around, on

the eastern-most edge of the Roraima formation

known as the Potaro Plateau in Guyana.

Granitic dome mountains occur on the Shield in the

southern part of the three Guianas (Guyana, Sur-

iname, French Guiana), where they are known as

‘‘inselbergs,’’ as well as in the western extreme of the

Shield in the Puerto Ayacucho region in Venezuela

where they are called ‘‘lajas.’’ Deposits of low-nutrient

white sands occur inland of the coastal plain, in belts

across the Shield, and in isolated pockets. Large areas

of savanna are found in the region, particularly the

complex of savannas that includes the Rupununi

Savanna in southwestern Guyana, the Gran Sabana

in eastern Venezuela, and the savannas of northern

Roraima, Brazil. In some of these areas the sands

overlay a clay hardpan that is resistant to penetration

by tree roots and that floods during the heavy rainy

season, resulting in limited forest growth. Tertiary and

Quaternary sediments separate the southern edge of

the Guiana Shield from the Amazon River and the

eastern edge from the Atlantic Ocean.

Climate

As a whole, the Guiana Shield region has a tropical

climate characterized by a relatively high mean annual

temperature exceeding 25uC at sea level, an annual

monthly maximum temperature range of less than 5uC,

and an average daily temperature range of approxi-

mately 6uC (Snow 1976). Because of the Guiana

Shield’s location just north of the equator, its climate

varies primarily according to elevation and effects of

the trade winds that combine to affect rainfall patterns.

The trade winds blow consistently from the east and

northeast, off of the Atlantic Ocean onto northeastern

South America, with wind speeds averaging from 3–4

m per second. Due to orographic effects, the eastern-

most escarpments of the mountains of the Guiana

Shield are generally localities of increased precipitation

where these moisture-laden winds meet the slopes

(Clarke et al. 2001). Seasonal oscillations of the

Intertropical Convergence Zone (ITCZ) also bring

variations in rainfall as the locations of low pressure

zones near the equator change (Snow 1976). Varying

primarily by latitude, one or two rainy seasons result

from shifts in the ITCZ. The heaviest rains usually

occur between May and August, whereas the rainy

season running from December to January is shorter

and less intense, with rains that do not penetrate as far

inland. Even during most dry seasons, frequent storms

provide adequate moisture to allow evergreen tropical

moist forests to persist in most low elevation parts of

the region.

Biological Diversity

The variety of landscapes of the Guiana Shield

includes sandstone tepuis, granite inselbergs, white

sands, seasonally flooded tropical savannas, lowlands

with numerous rivers, isolated mountain ranges, and

coastal swamps, each supporting a characteristic

vegetation (Huber 1995b, Huber et al. 1995). This

variety accounts for a great deal of the high diversity

and endemicity of the Shield’s biota. The highlands of

the Shield have a flora and fauna with numerous

endemic species. Some tepui endemic species occur as

low as 300 m in elevation, with increasing numbers by

1500 to 1800 m, and fully developed communities

occurring by 2000 m. Few if any plant or animal

specimens have been collected from most medium to

high elevation areas of the Guiana Shield. Many parts

of the Shield are poorly explored, including parts of

Brazil north of the Amazon River, much of eastern

Colombia, and the southern parts of Venezuela,

Guyana, Suriname, and French Guiana.

Conservation

With the exceptions of the populated localities such

as Puerto Ayacucho, Ciudad Guayana, Ciudad Bolı-

var, Boa Vista, Georgetown, Paramaribo, Cayenne, the

agricultural coastal areas, and open areas like the

Rupununi Savanna, the environment of the Guiana

Shield has benefited from limited access and low

population densities, although this same isolation has

hindered biodiversity research. Estimates vary, but

much of the vegetation is still relatively undisturbed by

human activities. Recently, however, the pace of

disturbance has greatly increased. Current threats to

the environment include large-scale logging by Asian

and local companies, large- and small-scale gold and

diamond mining, oil prospecting, bauxite mining,

hydroelectric dams, wildlife trade, and population-

related pressures such as burning, grazing, agriculture,

and the expansion of towns and villages. Taken

together, these impacts have begun to take their toll,

with vast areas vulnerable to increasing disturbance a

fact easily observed by using Google Earth and

‘‘flying’’ over the area.

The status of conservation efforts varies by country.

Throughout the Guiana Shield, many areas that are

designated as protected or otherwise restricted are

often only ‘‘paper’’ parks because of a lack of

infrastructure, funds, and will to actually protect the

areas. Over the last four decades, Venezuela has

established seven national parks, 29 natural monu-

ments, and two biosphere reserves covering about

142,280 km2, more than 30% of its share of the

NUMBER 17 3

Guiana Shield (Funk & Berry 2005). In Guyana, the

progress of conservation efforts has been slower, with

the only substantial protected area being Kaieteur

National Park, its 627 km2 comprising about 3% of

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

2004), with additional reserves under consideration.

Guyana’s 3710 km2 Iwokrama forest (Clarke et al.

2001) has parts listed as reserves, but overall it is

dedicated to sustainable use; unfortunately, logging

has begun, and the section of the road from Boa Vista,

Brazil, to Georgetown, Guyana, that runs through

Iwokrama is about to be paved. Suriname’s protected

areas system includes one national park and a network

of 11 reserves, totaling almost 20,000 km2, over 12%

of its total area. This includes the recently created

16,000 km2 Central Suriname Nature Reserve, a

UNESCO World Heritage Site that joined and

expanded three existing reserves (see http://www.

stinasu.com). French Guiana has no officially desig-

nated protected areas, but 18 proposed sites total 6710

km2, about 7.5% of its area (Lindeman & Mori 1989).

The natural areas of Venezuela and Guyana are

currently under the most anthropogenic pressure,

while those of French Guiana are probably less

threatened.

The Shield encompasses part or all of six countries

with six different governments, five official languages

and many more indigenous languages. Cooperation is

sometimes hampered by border disputes, illegal cross-

border activities involving gold and wildlife, and a lack

of interest by governments that are located far away.

The implementation of conservation practices is

further complicated by many issues concerning the

indigenous peoples of the region. All of these

challenges will have to be overcome on the way to

designing and maintaining a viable reserve system for

the Guiana Shield. However, it is critical that we gain

an understanding of the flora and fauna of the Shield

area so that decisions can be made on critical areas that

have high priority for conservation and so data can be

collected from areas that might ultimately be de-

stroyed. Because it is an ancient, fairly isolated

geological area, it is rich in endemic plant and animal

taxa, with many more likely to be discovered with

additional exploration. In addition, because this area

has been long neglected by biologists, it is often an area

of ‘‘inadequate information’’ for many biodiversity

analyses.

This volume contains the fishes from the Guiana

Shield, when paired with the previously published

Checklist of the Terrestrial Vertebrates of the Guiana

Shield (Hollowell & Reynolds 2005), we can examine

the size and scope of Vertebrates, an important

monophyletic group, known to inhabit the Guiana

Shield. Table 2 lists the vertebrate groups and their

sizes. The two checklists include a total of 53 orders,

189 families, 1190 genera, and 301 species. A large

percentage of the species (38%) are contributed by the

fishes listed in this volume.

Figure 2 compares the vertebrate diversity across the

major political areas of the Guiana Shield and shows

the species turnover between different areas. Themammals and reptiles have the most similar fauna

across the Shield with a 58% and 53% overlap,

respectively, between French Guiana (the extreme east)

and Venezuela-Amazonas (the extreme west). Fish and

birds have the least (24% and 10%, respectively). With

fishes this can probably be explained by the fact that

the headwaters of the rivers in the east are widely

separated from those of the west. The rivers of theShield in the west (Venezuela) have their source in the

Venezuelan Guayana and the Andes, in the central

portion (Guyana) the Essequibo drains mainly from

the Acari Mountains which lie on the border with

Brazil as does the Corantijn River (border between

Guyana and Suriname). To the east, rivers such as the

Maroni and Oyapock drain from the Tumuk-Humak

Mountains. The bird diversity percentage of ‘turn over’was surprisingly small until one realizes that there are

very different flyways that go across the eastern and

western parts of the Shield. When the three major

avenues of vertebrate mobility are examined (land, air,

water), it seems that the land provides the most stable

species make up and the air and water provide the

least. Could this have anything to do with the resulting

high species diversity of the birds and fishes?In the wider scope of biological understanding, the

goal of checklists of this type is to understand diversity

in terms of the spatial, evolutionary, and ecological

settings of physical environments, rather than simply

by political boundaries. The assembly of these lists is a

step toward considering the fauna in terms of the

geological entity of the Guiana Shield. Future studies

will include the analyses of animal community com-position on finer landscape scales, using developing

abilities to produce customized checklists for research

and conservation with Geographic Information System

(GIS) technologies drawing upon comprehensive data-

bases that include georeferenced museum specimen

records.

Biological Diversity of the Guiana Shield (BDG)

The ‘‘Biological Diversity of the Guiana ShieldProgram’’ (BDG) is a field-oriented program of the

Table 2.—Number of vertebrate taxa at different ranks.

Orders Families Genera Species

Amphibians 2 13 59 269

Reptiles 3 22 119 295

Mammals 11 35 143 282

Birds 22 70 493 1004

Fishes 15 49 376 1168

Total 53 189 1190 3018


National Museum of Natural History that began in

1983 (federally funded since 1987). The goal of the

BDG is to ‘‘study, document and preserve the

biological diversity of the Guiana Shield.’’ Most of

the program’s field work has taken place in Guyana,

but data analyses cover the majority of the Shield. In

Guyana, the BDG operates under the auspices of the

University of Guyana (UG). The BDG program is

designed to provide specimens and data to address

questions about many groups of organisms from

locations across the Shield. Information from BDG

collections and from other herbarium collections is

used to produce checklists, vegetation maps, floristic

and faunistic studies, revisions, and monographs. The

data generated from these studies are used to ask

questions about the make up of Guiana Shield

biological diversity, such as species turnover rates,

surrogate taxa, and areas of high diversity. Finally, the

BDG is exploring practical applications of the data

that have been collected through regular collaborations

with conservation and government agencies.

In addition to collecting and research, the BDG

Program trains students and scientists in both the

U.S.A. and Guyana, assists in their research, and has

established and helped to maintain collections. Over

the years several events have been hosted in Guyana,

including two Amerindian training courses, two bird

preparation courses, two plant identification courses, a

variety of lectures at the University and public venues,

and a public scientific symposium on the biological

diversity of Guyana. We also offer training opportu-

nities; nearly every year since 1987 the Program has

hosted at least one Guyanese student or UG staff

member at the Smithsonian. Many have participated in

the Natural History Museum’s Research Training

Program or the SI/MAB training courses. BDG

worked with the University of Guyana to raise funds

from the Royal Bank of Canada to construct a new

building, the ‘‘Centre for the Study of Biological

Diversity,’’ located on the campus of UG. More

recently, we worked with UG to raise funds from

USAID to build an extension on the original building.

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

NUMBER 17 5

The Centre houses collections and research space,

provides a library, and houses a Geographic Informa-tion System (GIS) facility. The goal of the Centre is to

combine research, education and conservation in the

study of biological diversity. The Centre is funded from

outside grants, but the staff is part of the University.

Currently, the plant database maintained by BDG has

161,108 specimen records and all sheets have been

barcoded. The BDG Program is working to make its

data available to the scientific community. Thecollections are being mapped using ArcMap and then

displayed on Google Earth as place marks. The

project, Georeferencing Plants of the Guiana Shield is

available on the Department of Botany public website

(http://botany.si.edu/bdg/georeferencing.cfm).

Acknowledgments

Special thanks go to the University of Guyana andthe Guyana EPA who have consistently supported our

efforts, including Mike Tamessar, Indarjit Ramdass,

and Philip da Silva, as well as past and present staff

members of the Centre for the Study of Biological

Diversity, in particular Calvin Bernard. This is number

153 in the Smithsonian’s Biological Diversity of the

Guiana Shield Program publication series.

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Arensica de la comunidad de Monochoa (Region de

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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,

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

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Amazonas, Bolıvar, Delta Amacuro; Guyana, Surinam,

French Guiana).—Contributions from the United States

National Herbarium 55:1–584.

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

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general eds. Missouri Botanical Garden, St. Louis.

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America. Elsevier Scientific Publishing Company, Amster-

dam.


FISHES OF THE GUIANA SHIELD

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

History

A vast complex of wetlands, lakes, streams, and

rivers drains the broad savannas, dense rainforests,extensive uplands, and tepuis of the Guiana Shield.

Early European explorers and colonists were impressed

not only by the many unusual fish species dwelling in

these water systems but also by the diversity of the

ichthyofauna. Accounts of fishes from those drainage

systems commenced with descriptions by pre-Linnaean

naturalists (e.g., Gronovius 1754, 1756) based on

specimens returned to Europe, with Linnaeus (1758)formally describing a number of these species.

Much of the early descriptive activity involving

fishes of the Guiana Shield centered on the ichthyo-

fauna of British Guiana (5 Guyana). Commentaries

on fishes from that colony by Bancroft (1769) and

Hilhouse (1825) preceded the formal description of

catfish species from the Demerara by Hancock (1828).

Cuvier and Valenciennes summarized the availableinformation on the ichthyofauna of all of the Guianas

in their series entitled Histoire Naturelle des Poissons

that documented the fish species known worldwide to

science to that time; with the catfishes being the first of

the groups inhabiting the shield discussed by those

authors (Cuvier & Valenciennes 1840a, b). These and

the other treatments of that era were, however, largely

opportunistic accounts based on scattered samplesreturned to Europe rather than derivative of focused

studies on the fish fauna of any region on the shield.

Consequently, the scale of the species-level diversity of

that ichthyofauna remained unknown and underap-

preciated.

Indications of the scale of the richness of the fish

fauna inhabiting the rivers of the Guiana Shield

commenced with the expeditions of the Schomburgkbrothers, Richard and Robert. In a remarkable

endeavor for that era, Robert collected fishes from

1835 to 1839 both in the more accessible shorter

northerly flowing rivers of the Guianas and through

portions of the Rıo Orinoco and Rıo Negro and the

Rio Branco. Drawings of fishes prepared during

Schomburgk’s travels across the shield served as the

basis for 83 species accounts in Jardine’s ‘‘Naturalist’sLibrary’’ (1841, 1843), including the formal descrip-

tions of a series of species. Unfortunately, the speci-

mens that were the basis for the drawings were not

preserved, and some illustrations combined details of

more than one species. Subsequent expeditions by the

Schomburgks traversed portions of what are now

Guyana, Venezuela, Suriname, and Brazil and yielded,

what was for that time, large numbers of fish specimens.In a series of publications Muller & Troschel (1845,

1848, 1849) recognized 141 species in the Schomburgk

collections and provided the first detailed illustrations

of fishes from South American freshwaters.

Diverse factors resulted in a lag in the state of

knowledge of the fishes inhabiting many portions of

the shield, with the comparative difficulty in accessi-

bility to inland regions clearly a paramount issue for

many areas. Supplementing that impediment were a

series of misadventures that bedeviled collectors who

sampled the fish fauna of the western portions of the

shield. Alexandre Rodriques Ferreira headed an

expedition that explored a significant portion of the

Rio Negro basin, commencing with a major collecting

effort through the Rio Branco system in 1786 (Ferreira

et al. 2007:12). Confounding Ferreira’s attempts to

publish his results were a string of unfortunate events

that culminated with the 1807 invasion of Portugal by

Napoleonic forces and the seizure and shipment of

Ferreira’s collections to Paris. Ferreira’s report on

animals from the Rio Branco region remained unpub-

lished until long after his death, and even then, only

parts appeared in print (see references in Ferreira

1983). In two expeditions between 1850 and 1852,

Alfred Russel Wallace (of Natural Selection fame)

collected over 200 species of fishes throughout the Rio

Negro basin including rivers draining the shield.

Wallace’s collections were lost with the sinking of the

ship returning him to England. Nonetheless, his field

sketches (Wallace 2002) document that the lost

collection included a number of species of fishes then

unknown to science (Regan 1905a, Toledo-Piza et al.

1999, Vari & Ferraris 2006).

An accelerating pace of ichthyological collecting

across many portions of the Guiana Shield during the

latter part of the nineteenth century resulted in the

discovery and description of numerous species. These

collections also documented the presence on the Shield

of many species originally described from elsewhere in

cis-Andean South America. Notwithstanding those

advances, the information was dispersed through

revisionary (e.g., Regan 1905b, c) and monographic

studies (e.g., Eigenmann & Eigenmann 1890), general

ichthyofaunal summaries of regions on the shield (e.g.,

Pellegrin 1908), and species descriptions in multiple

languages.

Exceptions to this pattern of scattered publication

were limited to a handful of papers focused on subsets

of the ichthyofauna from comparatively small regions.

Among the more notable of these were the analysis of

the catfishes of Suriname (Bleeker 1862), discussions of

the fishes present in portions of French Guiana

(Vaillant 1899, 1900), and a semi-popular overview of

the fishes of French Guiana (Pellegrin 1908). Compen-

dia of the freshwater fish species known from

individual colonies, countries, or regions were not

developed, let alone summaries of the fishes inhabiting

in the numerous streams, rivers, and lakes across the

shield. The dispersed literature prevented an appreci-

ation of the scale of the diversity of the shield

ichthyofauna.

The first overview of the freshwater fishes of

northeastern South America, including the Guiana

Shield, was Eigenmann’s (1912) treatise on the

freshwater fishes of British Guiana. Although Eigen-

mann sampled the fish fauna of only a comparatively

small section of British Guiana, his collections were

extensive for that era. In a series of papers, he and his

students described 128 new species from those collec-

tions (Eigenmann 1912:133). Eigenmann’s monograph

included data from his own collections, information

from the literature, and records of fishes that

originated on the Shield in various museums. Summary

tables (Eigenmann 1912:64) detailed the fish species

known from ten subunits that fall, at least in part,

within the boundaries of the Guiana Shield (Rıo

Orinoco basin, ‘‘West Coast’’ of British Guiana

[5 Barima River basin], Rio Branco basin, Rupununi

River, Lower Essequibo River, Lower Potaro River,

Demerara River, Dutch Guiana [5 Suriname], and

French Guiana [5 Guyane Francaise]).

Eigenmann (1912) reported 493 species from those

ten geographic units; a total that was in excess of the

species reported to that time from the rivers of the

Shield. These additional species were a function of two

factors. His total included all of the fish species then

known to inhabit the Rıo Orinoco; however, that vast

river system extends far beyond the Shield boundaries

with approximately only 40% of that watershed

overlying the Shield. Many of the fish species known

at the end of the first decade of the twentieth century

from the Rıo Orinoco basin originated in the llanos

(savannas) of the north central and western portions of

the basin. Aquatic habitats and the fish faunas in these

floodplain savanna settings differ dramatically from

the ecosystems and fish communities of the more

rapidly flowing rivers that drain the forested northern

slope of the Guiana Shield. Further inflating Eigen-

mann’s species total was his inclusion of some

primarily marine forms. Such species penetrate the

lower reaches of the rivers draining the Guianas during

periods of low river flow and consequent increased

estuarine salinity. Few, if any, of these species are likely

to range upriver onto the Shield even during the height

of the dry season.

The decades since Eigenmann’s monograph have

seen numerous ichthyological collecting expeditions in

many systems on the Shield. Two wide-ranging and

productive collecting endeavors through that region

during the first half of the twentieth century remain

relatively poorly known. John Haseman, who collected

throughout the Rio Branco basin and the southern-

most portion of the Rupununi River system in 1912

and 1913, made the first of these. Haseman deposited

these extensive collections in the Naturhistorisches

Museum in Vienna where he studied them for a year in

collaboration with Franz Steindachner. Nonetheless,

only one major publication based on those collections

was published (Steindachner 1915), most likely because

of the onset of World War I, disruptions during and

immediately after the conflict, and the death of

Steindachner soon after the cessation of hostilities.

Various revisionary studies in recent decades incorpo-

rated subsets of Haseman’s collection; nonetheless,

much of the material is yet-to-be analyzed critically.

The second collector, Carl Ternetz, sampled fishes

through the Rio Negro, Rıo Casiquiare, and Rıo

Orinoco basins during 1924 and 1925. Myers

(1927:107) remarked that the collection was ‘‘a

magnificent series of fishes, most of them hitherto

unexplored systematically by an ichthyologist.’’ Not-

withstanding the description of some new species

collected by Ternetz in rivers of the shield by Myers

(1927) and other authors and the use of portions of

that collection in some studies (e.g., Myers & Weitz-

man 1960), most of the material remains unstudied,

even after its transfer from Indiana University to the

California Academy of Sciences.

The 1960s brought a resurgence of major ichthyo-

logical collecting efforts in many of the river systems

on the shield (e.g., the Brokopondo Project; Boeseman

1968:4), with the pace of these endeavors accelerating

during recent decades. A compendium of these

collecting efforts lies beyond the purpose and scope

of this paper; however, as summarized in the next

section many of these expeditions were integral to

checklists, regional revisionary studies, and summaries

of the ichthyofauna in river basins or regions of the

Shield.

State of Knowledge of the Shield Fish Fauna

The nearly ten decades since the preparation of

Eigenmann’s 1912 magnum opus saw numerous

publications on fishes of the Guiana Shield. Many

were revisionary studies of genera or families whose

ranges extend far beyond the limits of the Shield, often

across major portions of cis-Andean South America

and in some instances into trans-Andean regions or

occasionally Central America. Other publications were

restricted to the members of a genus, subfamily, or

family from a country within the Shield region (e.g.,

Suriname: Boeseman 1968, Nijssen 1970, Kullander &

Nijssen 1989) or across a major portion of that area

(e.g., Boeseman 1982). Relatively few of these papers

involved broad surveys of an entire ichthyofauna in a

river system or country on the Shield with those that


did so summarized below arranged by the geographic

subdivisions in the checklist. Many include associated

ecological and life history information for fish com-

munities and individual species.

Brazil, Para (PA). The single publication of note

from this region is Ferreira (1993) that summarized the

results of intensive collecting efforts at sites within the

Rio Trombetas, one of the major northern tributaries

of the Amazon River east of the Rio Negro.

Brazil, Roraima (RO). Ferreira et al. (1988) summa-

rized the ichthyofauna at several closely situated

localities in the Rio Mucajai, a tributary of the Rio

Branco. More recently, Ferreira et al. (2007) provided

detailed information on the ichthyofauna across the

expanse of the Rio Branco basin, supplemented by

numerous color photographs, discussions of habitats,

and comments on the anthropogenic impact on the

aquatic systems within the basin.

French Guiana (FG). French Guiana has the most

intensely studied ichthyofauna of any portion of the

Guyana Shield. The first attempt to summarize

information on the fishes of the entire department

was that of Puyo (1949). Gery (1972) followed up with

studies of the characiforms (his characoids) from the

Guianas with a particular focus on French Guiana.

Planquette et al. (1996), Keith et al. (2000), and Le Bail

et al. (2000), in a groundbreaking series of publica-

tions, brought together information on the spectrum of

the freshwater fish fauna in that department. Each

species account includes a description, illustration, and

comments on its biology. Distributions within and

beyond French Guiana are discussed, and the sites of

occurrences of the species in the department are

plotted.

Guyana (GU). Notwithstanding the title of the

publication, Eigenmann’s (1912) monographic study

was based primarily on collections from the northern

portions of Guyana, in particular the Potaro River and

lower courses of the Essequibo and Demerara rivers,

albeit with that data supplemented with information

from the literature. Hardman et al. (2002) reported on

the fish fauna captured at Eigenmann’s collecting

localities nine decades after his expedition and provid-

ed a checklist of the 272 species collected in that survey.

Lowe (McConnell) (1964) included lists of fishes from

the southern most reaches of the Essequibo River

system along with observations on their ecology and on

the movements of various species during the yearly

flood and drought cycles. Watkins et al. (2004)

provided a summary of the fishes of the Iwokrama

Forest Reserve.

Suriname (SU). The ichthyofauna of Suriname

remains relatively poorly documented, with the listing

of the freshwater fishes in the country by Eigenmann

(1912:64–73) largely derived from literature informa-

tion. Boeseman (1952, 1953, 1954) supplemented

Eigenmann’s listing. Ouboter & Mol (1993) presented

the most comprehensive published list of the freshwater

fishes of Suriname. Kullander & Nijssen (1989)

published a detailed analysis of the cichlids of

Suriname.

Venezuela, Amazonas (VA). The state of Amazonas,

Venezuela, includes portions of the south-flowing Rıo

Negro of the Amazon basin, the north-draining Rıo

Orinoco and the entirety of the intervening Rıo

Casiquiare. Mago-Leccia (1971) produced the first

summary of the fishes of the Rıo Casiquiare. Lasso

(1992), Royero et al. (1992), and Lasso et al. (2004a,

2004b) provided information on the fish faunas of

various river systems within the state.

Venezuela, Bolivar (BO). A series of studies treated

the fish fauna of several right bank tributaries of the

Rıo Orinoco that drain the northern slopes of the

Guyana Shield. These included summaries of species in

various basins, with those listings supplemented in

some instances by information on fish biology and

distribution. Significant publications on the ichthyo-

fauna of the Rıo Caroni were published by Lasso

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

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

al. (2003), and Vispo et al. (2003). The Rıo Cuyuni, a

western tributary of the Essequibo River, drains the

eastern portions of the Shield in the state of Bolivar.

Machado-Allison et al. (2000) summarized the fish

fauna of the Venezuelan portions of that river system.

Lasso et al. (2004a) and Girardo et al. (2007) provide

supplemental information on the ichthyofauna of that

drainage basin.

Ichthyofaunal Richness

This checklist of fishes known from the water

systems of the Guiana Shield includes 1168 species.

Included in that total are representatives of 376 genera,

49 families, and 15 orders. Five orders are dominant in

terms of number of species living on the shield and

account for 96.7% of the species (Characiformes, 478

species and 41.0%; Siluriformes, 425 species and 36.4%;

Perciformes, 126 species and 10.8%; Gymnotiformes,

52 species and 4.5%; Cyprinodontiformes, 47 species

and 4.0%). This sum of 1168 species attests to the

dramatic improvement of our knowledge of the

freshwater fish fauna on the Shield in slightly less than

a century since Eigenmann (1912) documented fewer

than 500 species from that region. The 1168 species are

approximately 4.1% of the 28,400 fish species recently

estimated to be present in all marine and freshwater

systems worldwide (Nelson 2006), a percentage that

amply testifies to the striking diversity of the ichthyo-

fauna within that region. All the more noteworthy is

the species-level richness of the ichthyofauna within the

context of the overall Neotropical freshwater fish

fauna. According to a recent summary, approximately

5000 species of freshwater fishes occur across the

NUMBER 17 11

entirety of Central and South America (Reis et al.

2003). Thus, the drainage systems of the Guiana Shield

are home to approximately 23% of the freshwater fish

species that occur across the vast expanse between

southern South America and the southern border of

Mexico. Many factors contributed to the Shield region

being a repository of freshwater ichthyological diver-

sity, with a few particularly worthy of comment.

Physiography

The Guiana Shield is the ancient Precambrian

Guianan formation resulting from the uplift of the

underlying craton (Gibbs & Barron 1993) and demon-

strates attributes that generally lead to high levels of

biodiversity: geological diversity, a topographically

variable landscape, and transitions between ecosystems

(Killeen et al. 2002). Overall the region has a primarily

low to somewhat hilly physiography, albeit with some

abrupt changes in topography in the regions proximate

to the tepui formations that extend across much of the

region in an approximately east to west alignment.

Some river valleys have marked shifts in topography

with resultant waterfalls, rapids, and riffles that

increase the complexity of drainage system structure.

These topographic factors result in multiple aquatic

habitats with differing levels of physical complexity. At

one extreme are the lentic waters of swamps, wetlands,

channels, and lowland rivers. With increasing gradient,

the drainage systems progress through variably flowing

waters interrupted by higher energy settings such as

riffles and isolated lower scale rapids. Finally, there are

regions of greater gradients with rapidly flowing waters

and major repetitive rapids and waterfalls. To the

degree that differences in stream structure directly

correlate with elevational gradients, there also occur

differences in water temperatures.

Water Chemistry

Physical river system attributes are the most obvious

manifestation of variation in aquatic systems across the

Shield but represent a distinct subset of the spectrum of

factors that contribute to shifts in the composition and

relative biomass of fish communities across that region.

Complementing diversity in river structure are varia-

tions in water chemistry that occur not only between

but also within drainage basins across the Shield. Three

major water types occur in the tropics. The first of

these are white waters carrying nutrient-rich sediment

loads for at least part of the year and which, despite

their name, are actually brown (e.g., Rio Branco;

Goulding et al. 2003:42). A second major group are

clear water streams and rivers, including those draining

many regions of the eastern portion of the Amazonian

portion of the Shield (Para and Amapa; Goulding et al.

2003:43). Finally there are acidic black water rivers

that drain heavily leached soils where decomposing

plant matter produces high levels of fluvic and humic

acids but with the water poor in dissolved solids and

nutrients (e.g., Rio Negro; Goulding et al. 2003:44;

Savanna Belt rivers in Suriname, Ouboter & Mol

1993:134). Such water type differences occur both at

the level of major river systems and at a much smaller

scale within some river basins (Arbelaez et al. 2008).

Admixtures of water types resulting from within basin

water type differences yield conjoined drainages with

variably intermediate chemistries (e.g., downriver of

where the black water Rio Negro empties into the

white water Rio Solimoes at Manaus, Goulding et al.

2003:44; or where the white water Rupununi River

empties into the black water upper Essequibo River,

Watkins et al. 2005:40).

Species of freshwater fishes demonstrate physiolog-

ical, morphological, and behavioral adaptations that

often allow them to specialize for life in particular

water types but often simultaneously exclude them

from other water types. Some species or genera are,

therefore, more common in, or effectively limited to

waters with particular chemical characteristics (Lowe-

McConnell 1995). Exemplifying this situation are

acidic black waters such as those in the Rio Negro

that appear to be the primary, if not exclusive, habitat

for some fish species (Goulding et al. 1988, table 2).

These acidic waters are at the same time apparently

inimical to other species and some genera notwith-

standing the presence of these taxa in adjoining rivers

of different water types (Goulding et al. 1988:98). The

different water types also differ in degrees of primary

productivity and dependence on detritus-based energy

systems (De Jesus & Kohler 2004), factors that further

impact fish diversity and community composition.

Allochthonous Influences

Terrestrial habitats further influence freshwater

systems via the shift of nutrients and organic matter,

with the often-substantial input into water bodies

mediated by both water and wind. Terrestrial to

freshwater inputs and their impacts on aquatic systems

span a broad spectrum of scale. At one extreme, both

continuing upland erosion and periodic floods trans-

port dissolved and particulate matter into water bodies

(Sioli 1975, Polis et al. 1997). Alternative forms of

riparian vegetation also affect the amount and types of

allochthonous materials, including detritus, seeds and

fruits, and animals (primarily terrestrial insects) input

into the aquatic food web via runoff and wind. The

input of allochthonous detritus into Neotropical water

systems supports particularly large populations of

detritivorous fishes (Flecker 1996). Most notably,

many species and groups of fishes specialize on

exploiting allochthonous seeds, fruits, and insects, with

variation in the input of these items impacting the


composition of ichthyofaunal communities (see Gould-

ing et al. 1988, Boujard et al. 1990).

At the other end of the scale spectrum, alternate soil

types in conjunction with factors such as rainfall

regimes and temperature also support dramatically

different plant communities ranging from dense rain-

forests through open savannas. Physical attributes of

differing marginal plant communities directly influence

fish community composition and structure in the water

bodies that they border. Most obvious of these effects

is differential shading by riparian forests, an influence

that is particularly significant in terms of the species

dwelling in streams and smaller rivers. Marginal

vegetation and submerged macrophytes affect the

physical complexity of water bodies and thus the

composition of the resident ichthyofaunas in various

fashions. Most noteworthy among these are differential

inputs to the aquatic systems in the amount and type of

woody and leafy debris, variation in the submerged

portions of overhanging terrestrial plants along water

margins, and differing amounts and types of sub-

merged emergent vegetation. Synergy of drainage

structure and energy, differences in water types,

variation in associated riparian animals and plants,

and differences in input of nutrients yield dramatically

different fish communities. That variation at the local

level is a major contributor to the overall species-level

richness of the ichthyofauna across the totality of the

Shield.

Drainage System Interconnections

Physiological and physical factors closely tie fresh-

water fishes to drainage patterns. Historical separa-

tions of, and associations between, drainage systems

thereby contributed to the present day distributions of

many species on the Shield and the richness of that

fauna. Notwithstanding the tectonic quiescence of the

Guiana Shield for over 550 million years, the highlands

resulting from the uplift of the craton underwent

progressive erosion of the sedimentary layers overlying

that base (Gibbs & Barron 1993). The pronounced

degree of endemicity in many of the basins across the

Shield is indicative of their long isolation, with factors

including differences in water types being influential in

this regard. Nonetheless, there have been changes in

the water flow patterns on and along the margins of the

Shield that influenced the present composition of the

ichthyofauna in those systems.

At the large scale, tectonic events resulted in the

broader details of the present Orinoco and Amazon

basins, both of which contribute to the Shield

ichthyofauna. A large paleo-drainage encompassing

much of the present Orinoco and Amazon basin

drained north into the Caribbean Sea approximately

at the present location of Lago Maracaibo. Tectonic

events at the end of the Miocene resulted in separation

of the Amazon from Orinoco basin (Hoorn 1994).

Another consequence of these changes was the shift

eastward of the mainstream Orinoco along the

northern boundary of the Shield to its present mouth

slightly north of the northeastern margin of the Shield.

This dramatic realignment led to its capture of

drainages flowing from the northern slopes of the

Shield. The shift of the mainstream Amazon to it

present mouth similarly resulted in the capture of the

southern draining rivers of the Shield. Disrupting the

continuity between many of those freshwater systems

for varying periods were subsequent marine transgres-

sions into the eastern portions of the Amazon valley.

Superimposed on these large-scale drainage pattern

changes were long-term, often pronounced, climate

changes through the region (Baker et al. 2001) with

resultant sequential contraction and expansion of

suitable aquatic habitats. This combination of hydro-

graphic and climatic changes resulted in disruptions, in

some instances repeated disruptions, of previously

continuous species ranges, thereby setting the stage

for subsequent speciation.

The complex hydrological history of the drainages

on the Shield involved not only division of previously

continuous drainages but also in new connections

between various river systems. Connectivity resulted

from landscape tilting during uplift events and via

headwater stream capture. Those events permitted, and

continue to permit, movements of fishes between what

have been isolated basins on the Shield. Subsequent

disruptions of connectivity provided an opportunity

for the evolution of species. Such past connections may

account for unusual present day distribution patterns

previously highlighted by some authors (e.g., Armbrus-

ter 2005). Prominent among these possible connections

was the river hypothesized to have drained directly

from the south slope of the Guiana Shield into the

Atlantic Ocean through what is now northeastern

Guyana (Hammond 2005:137). Faunal similarities and

close phylogenetic relationships among included fish

species also point to possible past associations between

the upper Rıo Caroni of the Rıo Orinoco basin and the

Rıo Cuyuni, a western tributary of the Essequibo River

(Lasso et al. 1991a, Sabaj Perez & Birindelli 2008). An

interconnection between the Amazon and the upper

portions of the Maroni River was proposed by

Cardoso & Montoya-Burgos (2009) who also proposed

that temporary connections between adjoining river

systems during periods of lower sea levels permitted

dispersal of freshwater fishes along the coasts of the

Guianas.

The paramount example of an extant interconnec-

tion between major river systems on the Shield, or

indeed across the continent, is the Rio Casiquiare. This

over 300 km long natural canal connects the Rıo Negro

of the Amazon River basin with the upper portions of

the Rıo Orinoco. This unusual drainage begins as a

NUMBER 17 13

bifurcation of the upper portion of the Rıo Orinoco

and represents an ongoing capture of the upper portion

of the latter river system by the Rio Negro (Sternberg

1975, Winemiller at al. 2008). Despite its substantial

size at the divergence, where it is approximately 100 m

wide, and the fact that it carries a significant portion of

the total flow of that portion of the Rıo Orinoco, the

Rıo Casiquiare does not provide unimpeded transit for

all species of fishes between those basins. Lack of

interbasin species panmixis is, in part, a function of the

fact that the Rıo Casiquiare conjoins headwaters

habitats inhabited by a subset of the species resident

across the entirety of each basin. Equally, or perhaps

more significantly, the gradient in water types along

the course of the Rıo Casiquiare acts as a partial filter

that impedes movement of many fish species (Wine-

miller et al. 2008), thereby maintaining differences in

ichthyofaunal composition between the upper Rıo

Negro and upper Rıo Orinoco.

Although the Rıo Casiquiare is the most notable

connection between major rivers on the Guiana Shield

and the only year-round continuity, some degree of

seasonal connectivity occurs between the upper reaches

of the Rio Branco and the southern most portions of

the Essequibo River in the Rupununi Savannas of

southwestern Guyana. That region is an expansive

floodplain where the headwaters of the Rupununi

River (Essequibo basin) and the Takutu and Ireng

rivers (both components of the Rio Branco of the

Amazon River basin) come into close proximity. This

proximity and varying degree of continuity of the

headwaters of these rivers across a vast flooded plan

during high water periods [Lowe (McConnell) 1964,

Watkins et al. 2005] facilitate movement of at least

some fish species between the headwaters of the Rio

Branco and Essequibo River basins. Although such

movements potentially enrich the ichthyofaunas of

each of those river systems, they do not add to the

overall richness of the fish fauna of the shield.

Future Directions

Notwithstanding the series of papers listed under

‘‘State of Knowledge of the Shield Fish Fauna’’ and

many other publications, the continuing discovery and

description of freshwater species from water systems on

the Shield testifies to the incomplete state of our

knowledge of that fish fauna. The primary impediment

is the lack of exhaustive ichthyological collecting across

that vast region, with many river systems still

effectively unsampled (e.g., the upper Mazaruni River

in Guyana; Taphorn et al. 2008). Headwater tributary

streams, deep mainstream channels, and difficult-to-

sample habitats such as swamps and rapids remain

unsampled or poorly sampled even in those drainage

systems that have been the subject of ichthyological

collecting efforts. Those habitats and some largely

ichthyologically unexplored drainage systems hold the

greatest promise as sources of undescribed species;

however, we must not lose sight of the fact that areas

that have been long the foci of ichthyological sampling

continue to yield new species and are deserving of

continued attention. Recent monographic studies of

speciose genera and families of Neotropical freshwater

fishes demonstrate that collections in museums, uni-

versities, and research institutions of North and South

America and Europe house numerous species as of yet

unknown to science. Indeed in many groups, the vast

majority, if not all, of recently described species were

already represented in collections and awaited discov-

ery, often for many decades.

Thorough sampling of the freshwater fish fauna is

vital as is the thorough examination of materials in

collections, but equally or perhaps more important for

furthering our knowledge of the fishes on the Shield are

comprehensive revisionary studies of all groups of

fishes represented in that ichthyofauna. Such in-depth

studies are critical given our inadequate understanding

of the species-level diversity of many Neotropical

freshwater groups (Vari & Malabarba 1998). Inclusive

revisions of complex groups of Neotropical freshwater

fishes have repeatedly demonstrated that the sum of

long recognized species within a genus typically

underestimates the actual number of species in a taxon,

sometimes to a pronounced degree. An example is the

67 species now recognized in Creagrutus; a total three

and one-half times the number of species (19)

recognized in the genus prior to 1994 (Harold & Vari

1994, Vari & Harold 2001, Vari & Lima 2003, Ribeiro

et al. 2004, Torres-Mejia & Vari 2005). Comprehensive

revisions similarly further the subsequent identification

of additional previously unrecognized species by other

researchers (Reis 2004).

It is impossible to estimate the degree to which the

total number of freshwater fish species summarized in

this checklist falls short of the actual count of species

dwelling on the Shield. Nonetheless, it is clear that the

rate of continued additions to this speciose fish fauna,

the many regions and habitats that have not yet been

thoroughly explored ichthyologically, and the large

numbers of groups of fishes in the region that have not

yet been exhaustively studied portend a significant

increase in the species total.

Conservation Challenges

Deleterious anthropogenic activities impact freshwa-

ter ichthyofaunas across the Neotropics (Killeen 2007),

with many affecting various portions of the Guiana

Shield and adversely influencing fish communities in

the region. Adverse impacts are pervasive across

freshwater ichthyofaunas worldwide (Millennium Eco-

system Assessment 2005, Revenga et al. 2005), with

freshwater fishes consequently the most threatened


groups of vertebrates across the world in terms of

affected species (Dudgeon et al. 2006, Chapman et al.

2008). Major impacts on freshwater ichthyofaunas of

the Guiana Shield cover the spectrum of human

activities. These include overfishing for human con-

sumption and the aquarium trade, pollution from

agricultural, domestic, industrial and mining sources,

diversion of water for agricultural, domestic, and

industrial purposes, mining within river channels,

introductions of exotic species, transplanting of native

species between separate drainage systems, deforesta-

tion within drainage basins with consequent changes in

water flow patterns and quality, increased erosion and

siltation as a consequence of development, agriculture,

and mining operations, and impoundments for hydro-

electric and irrigation systems with disruption of

migration routes for fishes.

Major advances are necessary before we approach a

definitive understanding of the species-level diversity

for the fishes inhabiting the rivers, streams, lakes, and

other water bodies on the Guiana Shield. Nonetheless,

the following Checklist can serve as a foundation for

future studies, leading to a better appreciation of the

diversity of that ichthyofauna. Such information is

vital to inform decisions by resource managers,

government agencies, and members of the public

interested in protecting both the fish fauna and the

broader aquatic communities, both of which provide

essential and important ecosystem services across the

Shield.

Species of the Guiana Shield

The Checklist includes species recognized at the time

that contributors completed their accounts (mid-2008)

with these supplemented whenever possible by infor-

mation on new species described from the Shield

through early 2009. Readers interested in further

information on the families and species included in

the listing can refer to CLOFFSCA (Reis et al. 2003).

That listing includes bibliographic information for all

fish species in Central and South America, including

those known to occur on the Guiana Shield, through

the end of 2002. References to the original descriptions

of species published post that date are listed under the

following Guide to the Checklist. The regions utilized

in the checklist correspond to those used for terrestrial

vertebrates in Hollowell & Reynolds (2005). Abell et al.

(2008) recently proposed a hydrographically delimited

series of zones for South America. The more fine-

scaled resolution of that system is potentially more

informative in terms of areas of regional endemism for

aquatic organisms. We defer, however, from applying

it to the freshwater fishes of the Guiana Shield given

the large degree of uncertainty as to distributional

limits for most species in that fish fauna.

Acknowledgments

Completion of this Bulletin was facilitated by S.

Raredon who prepared the various images that face

each section. Numerous individuals assisted our fieldefforts and studies of the fishes on the Shield over the

years, with particular thanks to L. Aguana, A.

Machado-Allison, O. Castillo, J. Fernandez, S. Jewett,

C. Lasso, H. Madarie, F. Mago-Leccia, L. Parenti, F.

Provenzano, and D. Taphorn.

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GUIDE TO THE CHECKLIST

ALEKSANDAR RADOSAVLJEVIC

The cornerstone of this project was the exhaustive

Checklist of the freshwater fishes of South and Central

America (abbreviated hereafter as CLOFFSCA; Reis et

al. 2003). Only those species with confirmed collections

within the geographic bounds of the Guiana Shield

(after Gibbs & Barron 1993; excluding outlier forma-

tions in Colombia) were extracted from the

CLOFFSCA accounts. Records from the main channel

of the Amazon River that lies at a distance from Shield

boundaries were excluded, but those from the main

channel of the Rıo Orinoco that runs closer to the

northern limit of the Shield were included. This

preliminary assemblage was updated from the litera-

ture published since CLOFFSCA’s release and the

resultant summaries sent to relevant authorities for

review. Whenever possible, the families and/or sub-

families were reviewed by the original CLOFFSCA

authors. References include papers describing new

species from the area of interest that appeared after

the completion of CLOFFSCA.

Classification in the checklist follows CLOFFSCA.

Orders and families are listed in systematic order with

genera and species listed alphabetically within families/

subfamilies and genera, respectively. Subfamilies are

limited to the Characidae (Characiformes) and Lori-

cariidae (Siluriformes). Synonyms and common names

were not included, but that information is available in

CLOFFSCA.

The Guiana Shield was divided into 11 regions (see

Table 3 for abbreviations and Fig. 1 for map) to

illustrate distributions. The four regions in Brazil

include only those parts of each state (Amapa,

Amazonas, Para, Roraima) falling within the Shield’s

boundaries. Species from rivers forming borders

between two regions were listed as occurring in both

regions. Records from river systems not totally

contained within the Guiana Shield were only included

if it could be confirmed that a particular species was

collected in the portion of the drainage basin overlying

Shield. Regional abbreviations in the checklist are

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

uncertain or questionable.

Literature Cited

(Including papers describing new species published after completion

of CLOFFSCA.)

Armbruster, J. W. 2003a. The species of the Hypostomus cochliodon

group (Siluriformes: Loricariidae).—Zootaxa 249:1–60.

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Shield (Siluriformes: Loricariidae).—Zootaxa 344:1–12.

———. 2004. Pseudancistrus sidereus, a new species from southern

Venezuela (Siluriformes: Loricariidae) with a redescription of

Pseudancistrus.—Zootaxa 628:1–15.

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

with descriptions of two new species.—Neotropical Ichthyol-

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———. 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: Loricar-

iidae) from Guyana.—Zootaxa 920:1–12.

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

from the Rıo Caronı, Venezuela (Siluriformes: Loricarii-

dae).—Zootaxa 1731:33–41.

———, & D. C. Werneke. 2005. Peckoltia cavatica, a new loricariid

catfish from Guyana and a redescription of P. braueri

(Eigenmann 1912) (Siluriformes).—Zootaxa 882:1–14.

———, N. K. Lujan, & D. C. Taphorn. 2007. Four new Hypancistrus

(Siluriformes: Loricariidae) from Amazonas, Venezuela.—

Copeia 2007(1):62–79.

———, L. A. Tansey, & N. K. Lujan. 2007. Hypostomus rhantos

(Siluriformes: Loricariidae), a new species from southern

Venezuela.—Zootaxa 1553:59–68.

Benine, R. C., & G. A. M. Lopes. 2007. A new species of

Hemigrammus Gill, 1858 (Characiformes: Characidae) from

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

———, G. Z. Pelicao, & R. P. Vari. 2004. Tetragonopterus

lemniscatus (Characiformes: Characidae), a new species from

the Corantijn River basin in Suriname.—Proceedings of the

Biological Society of Washington 117:339–345.

Birindelli, J. L. O., M. H. Sabaj, & D. C. Taphorn. 2007. New species

of Rhynchodoras from the Rıo Orinoco, Venezuela, with

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

2007(3):672–684.

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

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

Cheirodontinae), and description of two new species from

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

Buitrago-Suarez, U. A., & B. M. Burr. 2007. Taxonomy of the catfish

genus Pseudoplatystoma Bleeker (Siluriformes: Pimelodidae)

with recognition of eight species.—Zootaxa 1512:1–38.

Chernoff, B., & A. Machado-Allison. 2005. Bryconops magoi and

Bryconops collettei (Characiformes: Characidae), two new

freshwater fish species from Venezuela, with comments on B.

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

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

corantijniensis, a new species from the Guyana Shield

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

approximately west to east order.

CG Colombian Guayana

VA Venezuela—Amazonas

BO Venezuela—Bolıvar

DA Venezuela—Delta Amacuro

BA Brazil—Amazonas

RO Brazil—Roraima

PA Brazil—Para

AP Brazil—Amapa

GU Guyana

SU Suriname

FG French Guiana

(Siluriformes: Loricariidae) with a molecular and morpholog-

ical description of the Pseudancistrus barbatus group.—

Zootaxa 1918:45–58.

Deynat, P. 2006. Potamotrygon marinae n. sp., a new species of

freshwater stingrays from French Guiana (Myliobatiformes:

Potamotrygonidae).—Comptes Rendus Biologies 329(7):483–

493.

DoNascimiento, C., & J. G. Lundberg. 2005. Myoglanis aspredi-

noides (Siluriformes: Heptapteridae), a new catfish from the

Rıo Ventuari, Venezuela.—Zootaxa 1009:37–49.

Friel, J. P. 2008. Pseudobunocephalus, a new genus of banjo catfish

with the description of a new species from the Orinoco River

system of Colombia and Venezuela (Siluriformes: Aspredini-

dae).—Neotropical Ichthyology 6(3):293–300.

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

(Teleostei, Characidae) e descricao de duas especies novas para

o genero.—Papeis Avulsos de Zoologia (Sao Paulo) 43(1):1–9.


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Hrbek, T., D. C. Taphorn, & J. E. Thomerson. 2005. Molecular

phylogeny of Austrofundulus Myers (Cyprinodontiformes:

Rivulidae), with revision of the genus and the description of

four new species.—Zootaxa 825:1–39.

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

(Teleostei: Characidae) une nouvelle espece de grand Serra-

salminae phytophage du bouclier guyanais.—Revue Suisse de

Zoologie 1024:833–853.

Keith, P., L. Nandrin, & P.-Y. Le Bail. 2006. Rivulus gaucheri, a new

species of rivuline (Cyprinodontiformes: Rivulidae) from

French Guiana.—Cybium 30(2):133–137.

Kullander, S. O., & E. J. G. Ferreira. 2005. Two new species of

Apistogramma Regan (Teleostei: Cichlidae) from the rio

Trombetas, Para State, Brazil.—Neotropical Ichthyology

3(3):361–371.

———, & ———. 2006. A review of the South American cichlid

genus Cichla, with descriptions of nine new species (Teleostei:

Cichlidae).—Ichthyological Exploration of Freshwaters 17(4):

289–398.

Lasso, C. A., & F. Provenzano. 2002. Dos nuevas especies de bagres

del genero Trichomycterus (Siluriformes: Trichomycteridae)

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

Revista de Biologıa Tropical 50(3/4):1139–1149.

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

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

Alcala, G. Galvis, L. Vasquez, M. Lugo, A. Machado Allison,

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

cuenca del rıo Orinoco. Parte I: lista de especies y distribucion

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

Lasso-Alcala, O. M., D. C. Taphorn B., C. A. Lasso, & O. Leon-

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dontiformes: Rivulidae) from the Paragua River system,

Caronı River drainage, Guyana Shield, Venezuela.—Zootaxa

1275:21–29.

Lehman, A. P., & R. E. Reis. 2004. Callichthys serralabium: a new

species of neotropical catfish from the upper Orinoco and

Negro Rivers (Siluriformes: Callichthyidae).—Copeia 2004(2):

336–343.

Lopez-Fernandez, H., & D. C. Taphorn. 2004. Geophagus abalios, G.

dicrozoster and G. winemilleri (Perciformes: Cichlidae), three

new species from Venezuela.—Zootaxa 439:1–27.

———, D. C. Taphorn Baechle, & S. O. Kullander. 2006. Two new

species of Guianacara from the Guiana Shield of eastern

Venezuela (Perciformes: Cichlidae).—Copeia 2006(3):384–

395.

Lujan, N. K. 2008. Description of a new Lithoxus (Siluriformes:

Loricariidae) from the Guayana Highlands with a discussion

of Guiana Shield biogeography.—Neotropical Ichthyology

6(3):413–418.

———, M. Arce, & J. W. Armbruster. 2009. A new black

Baryancistrus with blue sheen from the upper Orinoco

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

———, J. W. Armbruster, & M. H. Sabaj. 2007. Two new species of

Pseudancistrus from southern Venezuela (Siluriformes: Lor-

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163–174.

Lundberg, J. G., & W. M. Dahdul. 2008. Two new cis-Andean

species of the South American catfish genus Megalonema

allied to trans-Andean Megalonema xanthum, with description

of a new subgenus (Siluriformes: Pimelodidae).—Neotropical

Ichthyology 6(3):439–454.

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

Triportheus Cope, 1872 (Characiformes: Characidae).—Neo-

tropical Ichthyology 2(4):167–204.

Montana, C. G., H. Lopez-Fernandez, & D. C. Taphorn. 2008. A

new species of Crenicichla (Perciformes: Cichlidae) from the

Ventuari River, Upper Orinoco River Basin, Amazonas State,


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 (Pleuronecti-

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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.


Roman-Valencia, C., D. C. Taphorn B., & R. I. Ruiz-C. 2008. Two

new Bryconamericus: B. cinarucoense n. sp. and B. singularis n.

sp. (Characiformes, Characidae) from the Cinaruco River,

Orinoco Basin, with keys to all Venezuelan species.—Animal

Biodiversity and Conservation 31(1):15–27.

Romer, U., I. Hahn, & A. Conrad. 2006. Apistogramma wapisana

sp. n. Description of a dwarf cichlid from northern Brazil.

Cichlid Atlas 2. Mergus Verlag, Melle, Germany, pp. 748–

763.

Rosa, R. S., M. R. de Carvalho, & C. de Almeida Wanderley. 2008.

Potamotrygon boesemani (Chondrichthyes: Myliobatiformes:

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stingray from Surinam.—Neotropical Ichthyology 6(1):1–8.

Sabaj, M. H. 2005. Taxonomic assessment of Leptodoras (Siluri-

formes: Doradidae) with descriptions of three new species.—

Neotropical Ichthyology 3(4):637–678.

———, D. C. Taphorn, & O. E. Castillo G. 2008. Two new species of

thicklip thornycats, genus Rhinodoras (Teleostei: Siluriformes:

Doradidae).—Copeia 2008(1):209–226.

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

extant Doras Lacepede, 1803 (Siluriformes: Doradidae) with

descriptions of three new species.—Proceedings of the

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de Santana, C. D., & D. C. Taphorn. 2006. Sternarchorhynchus

gnomus, a new species of electric knifefish from the Lower Rıo

Caroni, Venezuela (Gymnotiformes: Apteronotidae).—Ich-

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

Platyurosternarchus (Gymnotiformes: Apteronotidae).—Co-

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Sarmento-Soares, L. M., & R. F. Martins-Pinheiro. 2008. A

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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

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Trichomycteridae), with discussion of their biogeography and

psammophily.—American Museum Novitates 3496:1–27.

Schindler, I., & W. Staeck. 2006. Geophagus gottwaldi sp. n. – a new

species of cichlid fish (Teleostei: Perciformes: Cichlidae) from

the drainage of the upper rıo Orinoco in Venezuela.—

Zoologische Abhandlungen (Dresden) 56:91–97.

———, & ———. 2008. Dicrossus gladicauda sp. n. – a new species of

crenicarine dwarf cichlids (Teleostei: Perciformes: Cichlidae)

from Colombia, South-America.—Vertebrate Zoology 58(1):

67–73.

Sidlauskas, B. L., & G. M. dos Santos. 2005. Pseudanos winterbot-

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Anostomidae) from Venezuela and Brazil, and comments on

its phylogenetic relationships.—Copeia 2005(1):109–123.

———, J. C. Garavello, & J. Jellen. 2007. A new Schizodon

(Characiformes: Anostomidae) from the Rıo Orinoco system,

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system.—Copeia 2007(3):711–725.

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

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

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

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Suijker, W. H., & G. E. Collier. 2006. Rivulus mahdiaensis, a new

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Taphorn, B. D. C., H. Lopez-Fernandez, & C. R. Bernard. 2008.



Zootaxa 1925:31–38.

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

longum (Characiformes: Crenuchidae), a new fish from


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the armored catfish genus Loricaria (Siluriformes: Loricar-

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

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———, ———, & M. C. C. de Pinna. 2005. The Neotropical whale

catfishes (Siluriformes: Cetopsidae: Cetopsinae), a revisionary

study.—Neotropical Ichthyology 3(2):127–238.

Vermeulen, F. B. M., & T. Hrbek. 2005 Kryptolebias sepia n. sp.

(Actinopterygii: Cyprinodontiformes: Rivulidae), a new killi-

fish from the Tapanahony River drainage in southeast

Surinam.—Zootaxa 928:1–20.

Werneke, D. C., J. W. Armbruster, N. K. Lujan, & D. C. Taphorn.

2005. Hemiancistrus guahiborum, a new suckermouth armored

catfish from Southern Venezuela (Siluriformes: Loricarii-

dae).—Neotropical Ichthyology 3(4):543–548.

———, M. H. Sabaj, N. K. Lujan, & J. W. Armbruster. 2005.

Baryancistrus demantoides and Hemiancistrus subviridis, two

new uniquely colored species of catfishes from Venezuela

(Siluriformes: Loricariidae).—Neotropical Ichthyology 3(4):

533–542.

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

P. Petry. 2003. Aphyocharax yekwanae, a new species of

bloodfin tetra (Teleostei: Characiformes: Characidae) from

the Guyana Shield of Venezuela.—Ichthyological Exploration

of Freshwaters 14(1):1–8.

Zanata, A. M., & M. Toledo-Piza. 2004. Taxonomic revision of the

South American fish genus Chalceus Cuvier (Teleostei:

Ostariophysi: Characiformes) with the description of three

new species.—Zoological Journal of the Linnean Society

140(1):103–135.

Zarske, A., P.-Y. Le Bail, & J. Gery. 2006. New and poorly known

Characiform fishes from French Guiana. 1. Two new Tetras

of the genera Hemigrammus and Hyphessobrycon (Teleostei:

Characiformes: Characidae).—Zoologische Abhandlungen

(Dresden) 55:17–30.

———, J. Gery, & I. Isbrucker. 2004. Moenkhausia rara sp. n. – eine

neue, bereits bestandsgefahrdete Salmler-Art (Teleostei: Char-

aciformes: Characidae) aus Surinam und Franzosisch Guayana

mit einer erganzenden Beschreibung von M. simulata (Eigen-

mann in Pearson, 1924).—Zoologische Abhandlungen (Dres-

den) 54:19–30.

NUMBER 17 23

CHECKLIST OF THE FISHES OF THE GUIANA SHIELD

Order: Pristiformes

Family: Pristidae—John D. McEachran & M. R. de Carvalho

Pristis pectinata Latham, 1794 DA GU?

Pristis pristis (Linnaeus, 1758) DA

Order: Myliobatiformes

Family: Potamotrygonidae—Marcelo R. de Carvalho & Ricardo

S. Rosa

Paratrygon aiereba (Muller & Henle, 1841) BO DA RO

Potamotrygon boesemani Rosa, Carvalho & Almeida, 2008 GU SU

Potamotrygon marinae Deynat, 2006 FG

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

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

Potamotrygon schroederi Fernandez-Yepez, 1957 VA BO DA

Potamotrygon scobina Garman, 1913 RO PA

Order: Osteoglossiformes

Family: Osteoglossidae—Carl J. Ferraris, Jr.

Osteoglossum bicirrhosum (Cuvier, 1829) CG RO AP GU FG

Family: Arapaimidae—Carl J. Ferraris, Jr.

Arapaima gigas (Schinz, 1822) GU

Order: Anguilliformes

Family: Ophichthidae—Sven O. Kullander

Stictorhinus potamius (Bohlke & McCosker, 1975) BO DA

Order: Clupeiformes

Family: Clupeidae—Carl J. Ferraris, Jr.

Rhinosardinia bahiensis (Steindachner, 1879) BO DA AP GU SU FG

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

Anchoa spinifer (Valenciennes, 1848) DA GU SU FG

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

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

Anchoviella cayennensis (Puyo, 1946) SU FG

Anchoviella guianensis (Eigenmann, 1912) BO DA GU SU FG

Anchoviella jamesi (Jordan & Seale, 1926) VA BO BA

Anchoviella lepidentostole (Fowler 1911) GU SU FG

Lycengraulis batesii (Gunther, 1868) BO DA GU SU FG

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

Pterengraulis atherinoides (Linnaeus, 1766) BO DA SU FG

Family: Pristigasteridae—Fabio di Dario

Pellona castelnaeana (Valenciennes, 1847) CG VA BO DA

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

Order: Characiformes

Family: Parodontidae—Carla S. Pavanelli

Apareiodon agmatos Taphorn, Lopez-Fernandez & Bernard,

2008

GU

Apareiodon gransabana Starnes & Schindler, 1993 BO GU SU FG

Apareiodon orinocensis Bonilla et al., 1999 VA BO

Parodon apolinari Myers, 1930 CG VA BO

Parodon bifasciatus Eigenmann, 1912 RO GU

Parodon guyanensis Gery, 1959 VA BO GU SU FG

Parodon suborbitalis Valenciennes, 1850 BO

Family: Curimatidae—Richard P. Vari

Curimata cerasina Vari, 1984 VA BO DA

Curimata cisandina (Allen, 1942) BA

Curimata cyprinoides (Linnaeus, 1766) DA GU SU FG

Curimata incompta Vari, 1984 VA BO DA

Curimata ocellata (Eigenmann & Eigenmann, 1889) CG VA BO BA

Curimata roseni Vari, 1989 VA BO DA GU

Curimata vittata (Kner, 1858) CG VA BO GU

Curimatella alburna (Muller & Troschel, 1844) RO GU

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

Curimatella immaculata (Fernandez-Yepez, 1948) CG VA BO DA RO GU

Curimatopsis crypticus Vari, 1982 AP GU SU FG

Curimatopsis evelynae Gery, 1964 CG VA BA?

Curimatopsis macrolepis (Steindachner, 1876) CG VA BO DA BA

Cyphocharax abramoides (Kner, 1859) CG VA BO PA

Cyphocharax festivus Vari, 1992 VA BO GU

Cyphocharax helleri (Steindachner, 1910) BO AP GU SU FG

Cyphocharax leucostictus (Eigenmann & Eigenmann, 1889) VA RO

Cyphocharax meniscaprorus Vari, 1992 VA BO

Cyphocharax mestomyllon Vari, 1992 BA

Cyphocharax microcephalus (Eigenmann & Eigenmann, 1889) GU SU FG?

Cyphocharax multilineatus (Myers, 1927) VA BO BA

Cyphocharax notatus (Steindachner, 1908) VA

Cyphocharax oenas Vari, 1992 VA BO DA

Cyphocharax punctatus (Vari & Nijssen, 1986) SU FG

Cyphocharax spilurus (Gunther, 1864) CG VA BO BA RO GU SU FG

Potamorhina altamazonica (Cope, 1878) CG VA BO DA

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

Psectrogaster essequibensis (Gunther, 1864) GU

Steindachnerina argentea (Gill, 1858) CG VA BO DA

Steindachnerina bimaculata (Steindachner, 1876) BO DA

Steindachnerina guentheri (Eigenmann & Eigenmann, 1889) BO DA GU

Steindachnerina planiventris Vari & Vari, 1989 RO

Steindachnerina pupula Vari, 1991 BO

Steindachnerina varii Gery, Planquette & Le Bail, 1991 AP SU FG

Family: Prochilodontidae—Richard P. Vari

Prochilodus mariae Eigenmann, 1922 CG VA BO DA

Prochilodus rubrotaeniatus Jardine, 1841 VA BO RO GU SU FG

Semaprochilodus insignis (Jardine, 1841) CG GU

Semaprochilodus kneri (Pellegrin, 1909) VA BO DA

Semaprochilodus laticeps (Steindachner, 1879) CG VA BO DA

Semaprochilodus varii Castro, 1988 SU FG

Family: Anostomidae—Richard P. Vari

Abramites hypselonotus (Gunther, 1868) VA BO DA GU


Anostomoides atrianalis Pellegrin, 1908 VA BO

Anostomoides laticeps (Eigenmann, 1912) VA BO GU

Anostomus anostomus (Linnaeus, 1758) GU

Anostomus brevior Gery, 1961 FG

Anostomus plicatus Eigenmann, 1912 GU SU

Anostomus spiloclistron Winterbottom, 1974 SU

Anostomus ternetzi Fernandez-Yepez, 1949 VA BO DA GU SU FG

Gnathodolus bidens Myers, 1927 VA BO

Laemolyta fernandezi Myers, 1950 VA BO DA

Laemolyta orinocensis (Steindachner, 1879) VA BO DA

Laemolyta proximate (Garman, 1890) BA GU

Laemolyta taeniata (Kner, 1859) CG VA BO

Leporinus acutidens (Valenciennes, 1836) FG

Leporinus affinis Gunther, 1864 BO

Leporinus agassizi Steindachner, 1876 VA BO

Leporinus alternus Eigenmann, 1912 DA GU

Leporinus arcus Eigenmann, 1912 VA BO GU SU FG

Leporinus badueli Puyo, 1948 FG

Leporinus brunneus Myers, 1950 VA BO BA

Leporinus desmotes Fowler, 1914 GU SU

Leporinus despaxi Puyo, 1943 RO FG

Leporinus fasciatus (Bloch, 1794) SU FG

Leporinus friderici (Bloch, 1794) GU SU FG

Leporinus gossei Gery, Planquette & Le Bail, 1991 SU FG

Leporinus granti Eigenmann, 1912 GU FG

Leporinus latofasciatus Steindachner, 1910 VA BO DA

Leporinus lebaili Gery & Planquette, 1983 SU FG

Leporinus leschenaulti Valenciennes, 1850 FG

Leporinus maculatus Muller & Troschel, 1844 GU SU FG

Leporinus megalepis Gunther, 1863 GU

Leporinus melanostictus Norman, 1926 AP FG

Leporinus nigrotaeniatus (Jardine, 1841) GU

Leporinus nijsseni Garavello, 1990 SU FG

Leporinus ortomaculatus Garavello, 2000 VA BO

Leporinus paralternus Fowler, 1914 GU

Leporinus pellegrinii Steindachner, 1910 GU SU FG

Leporinus pitingai Santos & Jegu, 1996 BA

Leporinus punctatus Garavello, 2000 VA BO

Leporinus spilopleura Norman, 1926 AP FG

Leporinus steyermarki Inger, 1956 VA BO DA

Leporinus uatumaensis Santos & Jegu, 1996 BA

Leporinus yophorus Eigenmann, 1922 VA BO

Pseudanos gracilis (Kner, 1858) VA BO BA

Pseudanos irinae Winterbottom, 1980 VA BO

Pseudanos trimaculatus (Kner, 1858) GU

Pseudanos winterbottomi Sidlauskas & Santos, 2005 VA BO

Sartor elongatus Santos & Jegu, 1987 PA

Schizodon fasciatus Spix & Agassiz, 1829 BO FG

NUMBER 17 27

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

Synaptolaemus cingulatus Myers & Fernandez-Yepez, 1950 VA BO

Family: Chilodontidae—Richard P. Vari

Caenotropus labyrinthicus (Kner, 1858) CG VA BO DA GU SU

Caenotropus maculosus (Eigenmann, 1912) BO GU SU FG

Caenotropus mestomorgmatos Vari, Castro & Raredon, 1995 VA BO BA

Chilodus punctatus Muller & Troschel, 1844 CG VA BO DA GU SU

Chilodus zunevei Puyo, 1946 FG

Family: Crenuchidae—Paulo A. Buckup

Ammocryptocharax elegans Weitzman & Kanazawa, 1976 CG VA BO

Ammocryptocharax lateralis (Eigenmann, 1909) GU

Ammocryptocharax minutus Buckup, 1993 CG VA BA

Ammocryptocharax vintonae (Eigenmann, 1909) BO GU

Characidium boaevistae Steindachner, 1915 VA BO RO

Characidium chupa Schultz, 1944 BO

Characidium declivirostre Steindachner, 1915 VA BO BA

Characidium hasemani Steindachner, 1915 BO RO

Characidium longum Taphorn, Montana & Buckup, 2006 VA BO BA

Characidium pellucidum Eigenmann, 1909 GU SU FG

Characidium pteroides Eigenmann, 1909 GU

Characidium steindachneri Cope, 1878 CG VA BO BA RO PA GU

Characidium zebra Eigenmann, 1909 CG VA BO DA BA RO PA AP GU SU FG

Crenuchus spilurus Gunther, 1863 CG VA BO GU SU FG

Elachocharax geryi Weitzman & Kanazawa, 1978 CG VA BO BA

Elachocharax mitopterus Weitzman, 1986 VA BA

Elachocharax pulcher Myers, 1927 CG VA BO BA PA

Leptocharacidium omospilus Buckup, 1993 VA BO BA RO

Melanocharacidium blennioides (Eigenmann, 1909) BO GU

Melanocharacidium compressus Buckup, 1993 VA BO

Melanocharacidium depressum Buckup, 1993 VA BA

Melanocharacidium dispilomma Buckup, 1993 CG VA BO BA RO PA FG

Melanocharacidium melanopteron Buckup, 1993 VA BO

Melanocharacidium nigrum Buckup, 1993 BA RO

Melanocharacidium pectorale Buckup, 1993 VA BO BA RO PA

Microcharacidium eleotrioides (Gery, 1960) SU FG

Microcharacidium gnomus Buckup, 1993 CG VA BO BA

Microcharacidium weitzmani Buckup, 1993 CG VA BO BA PA

Odontocharacidium aphanes (Weitzman & Kanazawa, 1977) CG VA BA

Poecilocharax bovalii Eigenmann, 1909 GU

Poecilocharax weitzmani Gery, 1965 CG VA BO

Skiotocharax meizon Presswell, Weitzman & Bergquist, 2000 GU

Family: Hemiodontidae—Francisco Langeani Neto

Anodus elongatus Agassiz, 1829 PA

Anodus orinocensis (Steindachner, 1887) CG VA BO DA

Argonectes longiceps (Kner, 1858) PA SU FG

Bivibranchia bimaculata Vari, 1985 GU SU FG

Bivibranchia fowleri (Steindachner, 1908) CG VA BO DA BA RO GU

Bivibranchia simulata Gery, Planquette & Le Bail, 1991 AP FG


Bivibranchia velox (Eigenmann & Myers, 1927) AP

Hemiodus amazonum (Humboldt, 1821) GU

Hemiodus argenteus Pellegrin, 1908 CG VA BO DA GU SU

Hemiodus atranalis (Fowler, 1940) RO PA GU

Hemiodus goeldii Steindachner, 1908 AP

Hemiodus gracilis Gunther, 1864 VA BO

Hemiodus huraulti (Gery, 1964) SU FG

Hemiodus immaculatus Kner, 1858 CG VA BO DA PA

Hemiodus microlepis Kner, 1858 VA BO

Hemiodus quadrimaculatus Pellegrin, 1908 RO AP GU SU FG

Hemiodus semitaeniatus Kner, 1858 CG VA BO GU

Hemiodus thayeria Bohlke, 1955 CG VA BO BA

Hemiodus unimaculatus (Bloch, 1794) AP GU SU FG

Hemiodus vorderwinkleri (Gery, 1964) BO GU

Family: Gasteropelecidae—Stanley H. Weitzman

Carnegiella marthae Myers, 1927 CG VA BO DA BA

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

Gasteropelecus sternicla (Linnaeus, 1758) BO DA GU FG

Thoracocharax stellatus (Kner, 1858) CG VA BO DA

Family: Characidae

Genera Incerta Sedis—Flavio C. T. Lima

Aphyocharacidium melandetum (Eigenmann, 1912) GU

Aphyodite grammica Eigenmann, 1912 GU

Astyanax bimaculatus (Linnaeus, 1758) CG VA BO DA GU SU FG

Astyanax clavitaeniatus Garutti, 2003 RO

Astyanax fasciatus (Cuvier, 1819) BO

Astyanax guianensis Eigenmann, 1909 VA BO GU

Astyanax leopoldi Gery, Planquette & Le Bail, 1988 AP FG

Astyanax metae Eigenmann, 1914 VA BO

Astyanax mutator Eigenmann, 1909 GU

Astyanax myersi (Fernandez-Yepez, 1950) VA BO

Astyanax rupununi Garutti, 2003 GU

Astyanax scintillans Myers, 1928 VA BO

Astyanax siapae Garutti, 2003 VA

Astyanax validus Gery, Planquette & Le Bail, 1991 FG

Astyanax venezuelae Schultz, 1944 VA BO

Aulixidens eugeniae Bohlke, 1952 VA BO

Bryconamericus alpha Eigenmann, 1914 CG VA BO

Bryconamericus beta Eigenmann, 1914 VA BO

Bryconamericus cinarucoense Roman-Valencia, Taphorn & Ruiz-

C., 2008

GU

Bryconamericus cismontanus Eigenmann, 1914 VA BO

Bryconamericus cristiani Roman-Valencia, 1998 CG

Bryconamericus deuterodonoides Eigenmann, 1914 VA BO

Bryconamericus hyphesson Eigenmann, 1909 GU

Bryconamericus macropthalmus Roman-Valencia, 2003 VA

Bryconamericus orinocoense Roman-Valencia, 2003 VA

Bryconamericus subtilisform Roman-Valencia, 2003 BO

Bryconexodon trombetasi Jegu, Santos & Ferreira, 1991 PA

NUMBER 17 29

Bryconops affinis (Gunther, 1864) GU SU FG

Bryconops alburnoides Kner, 1858 CG VA BO

Bryconops caudomaculatus (Gunther, 1864) VA BO DA GU SU FG

Bryconops colanegra Chernoff & Machado-Allison, 1999 VA BO

Bryconops colaroja Chernoff & Machado-Allison, 1999 BO

Bryconops collettei Chernoff & Machado-Allison, 2005 VA BO

Bryconops cyrtogaster (Norman, 1926) AP FG

Bryconops disruptus Machado-Allison & Chernoff, 1997 VA BA

Bryconops giacopinii (Fernandez-Yepez, 1950) VA BO

Bryconops humeralis Machado-Allison, Chernoff & Buckup,

1996

VA BO

Bryconops imitator Chernoff & Machado-Allison, 2002 VA BO

Bryconops inpai Knoppel, Junk & Gery, 1968 VA BA

Bryconops melanurus (Bloch, 1794) GU SU FG

Bryconops vibex Machado-Allison, Chernoff & Buckup, 1996 VA BO

Ceratobranchia joanae Chernoff & Machado-Allison, 1990 VA BO

Chalceus epakros Zanata & Toleda-Piza, 2004 VA BO RO PA GU

Chalceus macrolepidotus Cuvier, 1816 CG VA BO DA GU SU FG

Creagrutus bolivari Schultz, 1944 CG VA BO DA

Creagrutus ephippiatus Vari & Harold, 2001 VA BO DA

Creagrutus gyrospilus Vari & Harold, 2001 VA BO

Creagrutus machadoi Vari & Harold, 2001 BO

Creagrutus magoi Vari & Harold, 2001 VA BO

Creagrutus maxillaris (Myers, 1927) CG VA BO BA

Creagrutus melanzonus Eigenmann, 1909 BO GU FG

Creagrutus melasma Vari, Harold & Taphorn, 1994 VA BO

Creagrutus menezesi Vari & Harold, 2001 RO

Creagrutus phasma Myers, 1927 CG VA BO BA

Creagrutus planquettei Gery & Renno, 1989 FG

Creagrutus provenzanoi Vari & Harold, 2001 VA BO

Creagrutus runa Vari & Harold, 2001 CG VA BO BA

Creagrutus veruina Vari & Harold, 2001 VA

Creagrutus vexillapinnus Vari & Harold, 2001 CG VA BA?

Creagrutus xiphos Vari & Harold, 2001 BO

Creagrutus zephyrus Vari & Harold, 2001 CG VA BA?

Ctenobrycon spilurus (Valenciennes, 1850) VA BO DA GU SU FG

Deuterodon potaroensis Eigenmann, 1909 GU

Exodon paradoxus Muller & Troschel, 1844 GU

Gymnocorymbus thayeri Eigenmann, 1908 CG VA BO DA GU

Gymnotichthys hildae Fernandez-Yepez, 1950 VA BO

Hemibrycon surinamensis Gery, 1962 SU FG

Hemigrammus aereus Gery, 1959 FG

Hemigrammus analis Durbin, 1909 CG VA BO BA GU

Hemigrammus barrigonae Eigenmann & Henn, 1914 CG VA BO

Hemigrammus belottii (Steindachner, 1882) FG

Hemigrammus boesemani Gery, 1959 SU FG

Hemigrammus cylindricus Durbin, 1909 GU

Hemigrammus erythrozonus Durbin, 1909 GU

Hemigrammus guyanensis Gery, 1959 FG


Hemigrammus iota Durbin, 1909 GU

Hemigrammus lunatus Durbin, 1918 SU

Hemigrammus marginatus Ellis, 1911 VA BO DA

Hemigrammus micropterus Meek, 1907 VA BO DA

Hemigrammus microstomus Durbin, 1918 VA BO

Hemigrammus mimus Bohlke, 1955 VA BO

Hemigrammus newboldi (Fernandez-Yepez, 1949) VA BO DA

Hemigrammus ocellifer (Steindachner, 1882) GU SU FG

Hemigrammus ora Zarske, Le Bail & Gery, 2006 FG

Hemigrammus orthus Durbin, 1909 GU

Hemigrammus rhodostomus Ahl, 1924 CG VA BO

Hemigrammus rodwayi Durbin, 1909 GU SU FG

Hemigrammus schmardae (Steindachner, 1882) CG VA BO BA

Hemigrammus stictus (Durbin, 1909) CG VA BO GU

Hemigrammus taphorni Benine & Lopes, 2007 BO

Hemigrammus unilineatus (Gill, 1858) GU SU FG

Hyphessobrycon albolineatum Fernandez-Yepez, 1950 VA BO

Hyphessobrycon borealis Zarske, Le Bail & Gery, 2006 AP SU FG

Hyphessobrycon catableptus (Durbin, 1909) GU

Hyphessobrycon copelandi Durbin, 1908 FG

Hyphessobrycon diancistrus Weitzman, 1977 CG VA BO

Hyphessobrycon eos Durbin, 1909 GU

Hyphessobrycon epicharis Weitzman & Palmer, 1997 CG VA BA

Hyphessobrycon eques (Steindachner, 1882) FG

Hyphessobrycon georgettae Gery, 1961 SU

Hyphessobrycon hildae Fernandez-Yepez, 1950 VA BO

Hyphessobrycon metae Eigenmann & Henn, 1914 CG VA BO

Hyphessobrycon minimus Durbin, 1909 GU

Hyphessobrycon minor Durbin, 1909 GU

Hyphessobrycon rosaceus Durbin, 1909 GU SU

Hyphessobrycon roseus (Gery, 1960) FG

Hyphessobrycon simulatus (Gery, 1960) FG

Hyphessobrycon sweglesi (Gery, 1961) CG VA BO

Hyphessobrycon takasei Gery, 1964 AP FG

Jupiaba abramoides (Eigenmann, 1909) CG VA BO GU SU FG

Jupiaba atypindi Zanata, 1997 RO

Jupiaba essequibensis (Eigenmann, 1909) GU

Jupiaba keithi (Gery, Planquette & Le Bail, 1996) FG

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

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

Jupiaba mucronata (Eigenmann, 1909) GU

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

Jupiaba pinnata (Eigenmann, 1909) GU SU

Jupiaba polylepis (Gunther, 1864) PA GU SU

Jupiaba potaroensis (Eigenmann, 1909) GU

Jupiaba scologaster (Weitzman & Vari, 1986) CG VA BO BA

Knodus heteresthes (Eigenmann, 1908) VA BO

Microschemobrycon callops Bohlke, 1953 VA BO PA

NUMBER 17 31

Microschemobrycon casiquiare Bohlke, 1953 VA BA

Microschemobrycon melanotus (Eigenmann, 1912) GU

Microschemobrycon meyburgi Meinken, 1975 RO

Moenkhausia browni Eigenmann, 1909 GU

Moenkhausia chrysargyrea (Gunther, 1864) VA BO GU FG

Moenkhausia collettii (Steindachner, 1882) CG VA BO DA GU SU FG

Moenkhausia copei (Steindachner, 1882) CG VA BO

Moenkhausia cotinho Eigenmann, 1908 CG VA BO

Moenkhausia dichroura (Kner, 1858) VA BO DA

Moenkhausia georgiae Gery, 1965 SU FG

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

Moenkhausia hemigrammoides Gery, 1965 SU FG

Moenkhausia inrai Gery, 1992 FG

Moenkhausia intermedia Eigenmann, 1908 CG VA BO FG

Moenkhausia lata Eigenmann, 1908 AP FG

Moenkhausia lepidura (Kner, 1858) VA BO DA GU SU

Moenkhausia megalops (Eigenmann, 1907) FG

Moenkhausia miangi Steindachner, 1915 BO RO

Moenkhausia moisae Gery, Planquette & Le Bail, 1995 FG

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

Moenkhausia rara Zarske, Gery & Isbrucker, 2004 PA SU FG

Moenkhausia shideleri Eigenmann, 1909 GU

Moenkhausia surinamensis Gery, 1965 AP SU FG

Paracheirodon axelrodi (Schultz, 1956) CG VA BO BA

Paracheirodon simulans (Gery, 1963) CG VA BO BA

Paragoniates alburnus Steindachner, 1876 VA BO

Parapristella aubynei (Eigenmann, 1909) GU

Pristella maxillaris (Ulrey, 1894) VA BO GU FG

Salminus hilarii Valenciennes, 1850 CG VA BO RO

Scissor macrocephalus Gunther, 1864 SU

Serrabrycon magoi Vari, 1986 CG VA BO BA

Thayeria ifati Gery, 1959 FG

Triportheus angulatus (Spix & Agassiz, 1829) GU?

Triportheus auritus Malabarba, 2004 VA BO DA GU?

Triportheus brachipomus Malabarba, 2004 VA AP? GU SU FG

Triportheus elongatus (Gunther, 1864) VA BO

Triportheus venezuelensis Malabarba, 2004 BO

Xenagoniates bondi Myers, 1942 CG VA BO DA

Subfamily: Agoniatinae—Angela M. Zanata

Agoniates anchovia Eigenmann, 1914 PA

Agoniates halecinus Muller & Troschel, 1845 VA BO RO PA AP? GU

Subfamily: Iguanodectinae—Cristiano R. Moreira

Iguanodectes adujai Gery, 1970 VA BA

Iguanodectes geisleri Gery, 1970 VA BA

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

Piabucus dentatus (Koelreuter, 1763) VA DA GU SU FG

Subfamily: Bryconinae—Flavio C. T. Lima

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


Brycon bicolor Pellegrin, 1909 VA BO DA

Brycon coquenani Steindachner, 1915 VA BO

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

Brycon pesu Muller & Troschel, 1845 CG VA BO DA RO PA AP GU SU FG

Subfamily: Serrasalminae—Michel Jegu

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

Acnodon senai Jegu & Santos, 1990 PA AP

Catoprion mento (Cuvier, 1819) CG VA BO DA BA RO PA AP GU

Colossoma macropomum (Cuvier, 1818) CG VA BO DA

Metynnis altidorsalis Ahl, 1923 GU SU

Metynnis argenteus Ahl, 1924 VA BO DA GU

Metynnis hypsauchen (Muller & Troschel, 1844) VA BO RO GU

Metynnis lippincottianus (Cope, 1870) CG VA AP? FG

Metynnis luna (Cope, 1878) CG VA BO GU

Metynnis orinocensis (Steindachner, 1908) DA

Mylesinus paraschomburgkii Jegu, Santos & Ferreira, 1989 BA PA AP

Mylesinus schomburgkii Valenciennes, 1850 GU

Myleus knerii (Steindachner, 1881) FG

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

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

Myloplus lobatus (Valenciennes, 1850) PA

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

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

Myloplus rubripinnis (Muller & Troschel, 1844) CG VA BO RO AP GU SU FG

Myloplus schomburgkii (Jardine & Schomburgk, 1841) CG VA BO BA PA

Myloplus ternetzi (Norman, 1929) AP SU FG

Myloplus torquatus (Kner, 1858) VA BO BA RO GU

Mylossoma aureum (Agassiz, 1829) BO DA

Mylossoma duriventre (Cuvier, 1818) VA BO DA AP

Piaractus brachypomus (Cuvier, 1818) VA BO DA GU

Pristobrycon aureus (Spix & Agassiz, 1829) GU

Pristobrycon calmoni (Steindachner, 1908) BO DA BA AP GU

Pristobrycon careospinus Fink & Machado-Allison, 1992 CG VA

Pristobrycon eigenmanni (Norman, 1929) VA BO DA BA RO PA AP GU SU FG

Pristobrycon maculipinnis Fink & Machado-Allison, 1992 VA

Pristobrycon striolatus (Steindachner, 1908) VA BO DA BA PA AP GU SU FG

Pygocentrus cariba (Humboldt & Valenciennes, 1821) CG VA BO DA

Pygocentrus nattereri Kner, 1858 AP GU

Pygopristis denticulata (Cuvier, 1819) VA BO BA PA AP GU SU FG

Serrasalmus altispinis Merckx, Jegu & Santos, 2000 BA PA

Serrasalmus altuvei Ramırez, 1965 VA BO DA

Serrasalmus elongatus Kner, 1858 BO DA BA PA

Serrasalmus gouldingi Fink & Machado-Allison, 1992 VA BO BA PA

Serrasalmus hastatus Fink & Machado-Allison, 2001 BA RO

Serrasalmus irritans Peters, 1877 VA BO DA

Serrasalmus maculatus Kner, 1858 AP

Serrasalmus manueli (Fernandez-Yepez & Ramırez, 1967) CG VA BA AP

Serrasalmus medinai Ramırez, 1965 BO DA

NUMBER 17 33

Serrasalmus nalseni Fernandez-Yepez, 1969 VA BO

Serrasalmus rhombeus (Linnaeus, 1766) CG VA BO DA BA RO PA AP GU SU FG

Serrasalmus serrulatus (Valenciennes, 1850) GU?

Tometes lebaili Jegu, Keith & Belmont-Jegu, 2002 SU FG

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

Tometes trilobatus Valenciennes, 1850 AP FG

Subfamily: Aphyocharacinae—Rosana S. Lima

Aphyocharax avary Souza-Lima, 2003 VA BO GU

Aphyocharax colifax Taphorn & Thomerson, 1991 VA BO

Aphyocharax erythrurus Eigenmann, 1912 GU

Aphyocharax yekwanae Willink, Chernoff, Machado-Allison,

Provenzano & Petry, 2003

BO

Subfamily: Characinae—Naercio A. Menezes & Carlos A.S.

Lucena

Acanthocharax microlepis Eigenmann, 1912 GU

Acestrocephalus sardina (Fowler, 1913) VA BO BA GU

Charax apurensis Lucena, 1987 VA BO DA

Charax gibbosus (Linnaeus, 1758) GU SU

Charax hemigrammus (Eigenmann, 1912) GU

Charax metae Eigenmann, 1922 BO

Charax michaeli Lucena, 1989 RO

Charax notulatus Lucena, 1987 VA BO DA

Charax rupununi Eigenmann, 1912 PA GU

Cynopotamus bipunctatus Pellegrin, 1909 VA BO

Cynopotamus essequibensis Eigenmann, 1912 GU SU FG

Gnathocharax steindachneri Fowler, 1913 CG VA BO GU

Heterocharax leptogrammus Toledo-Piza, 2000 VA BA

Heterocharax macrolepis Eigenmann, 1912 CG VA BO GU

Heterocharax virgulatus Toledo-Piza, 2000 VA BA

Lonchogenys ilisha Myers, 1927 CG VA BO BA

Phenacogaster apletostigma Lucena & Gama, 2007 AP

Phenacogaster carteri (Norman, 1934) GU

Phenacogaster megalostictus Eigenmann, 1909 GU

Phenacogaster microstictus Eigenmann, 1909 GU SU

Priocharax ariel Weitzman & Vari, 1987 VA BO

Roeboides affinis (Gunther, 1868) VA BO

Roeboides araguaito Lucena, 2003 CG

Roeboides dientonito Schultz, 1944 VA BO DA GU

Roeboides myersii Gill, 1870 VA BO DA

Roeboides numerosus Lucena, 2000 VA BO

Roeboides oligistos Lucena, 2000 RO PA

Roeboides thurni Eigenmann, 1912 GU SU FG

Subfamily: Stethaprioninae—Roberto E. Reis

Brachychalcinus orbicularis (Valenciennes, 1850) GU SU

Poptella brevispina Reis, 1989 RO PA GU SU

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

Poptella longipinnis (Popta, 1901) CG VA BO DA SU

Subfamily: Tetragonopterinae—Roberto E. Reis

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


Tetragonopterus lemniscatus Benine, Pelicao & Vari, 2004 SU

Subfamily: Cheirodontinae—Luiz R. Malabarba

Cheirodontops geayi Schultz, 1944 VA BO DA

Odontostilbe gracilis (Gery, 1960) FG

Odontostilbe littoris (Gery, 1960) FG

Odontostilbe pulchra (Gill, 1858) VA BO DA RO

Odontostilbe splendida Buhrnheim & Malabarba, 2007 BO

Subfamily: Glandulocaudinae—Stanley H. Weitzman

Ptychocharax rhyacophila Weitzman, Fink, Machado-Allison &

Royero, 1994

VA

Family: Acestrorhynchidae—Naercio A. Menezes

Acestrorhynchus falcatus (Bloch, 1794) VA GU SU FG

Acestrorhynchus falcirostris (Cuvier, 1819) CG VA BA GU

Acestrorhynchus grandoculis Menezes & Gery, 1983 VA BO BA

Acestrorhynchus heterolepis (Cope, 1878) VA

Acestrorhynchus microlepis (Schomburgk, 1841) CG VA BO DA BA GU SU FG

Acestrorhynchus minimus Menezes, 1969 VA BO PA

Acestrorhynchus nasutus Eigenmann, 1912 VA GU

Family: Cynodontidae—Monica Toledo-Piza

Cynodon gibbus Spix & Agassiz, 1829 CG BO DA GU

Cynodon meionactis Gery, Le Bail & Keith, 1999 SU FG

Cynodon septenarius Toledo-Piza, 2000 VA BO GU

Hydrolycus armatus (Jardine, 1841) CG VA BO DA GU

Hydrolycus tatauaia Toledo-Piza, Menezes & Santos, 1999 VA BO GU

Hydrolycus wallacei Toledo-Piza, Menezes & Santos, 1999 VA BO

Rhaphiodon vulpinus Spix & Agassiz, 1829 VA BO DA GU

Roestes ogilviei (Fowler, 1914) RO GU

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

Oyakawa)

Erythrinus erythrinus (Bloch & Schneider, 1801) CG VA BO DA GU SU FG

Hoplerythrinus gronovii (Valenciennes, 1847) FG

Hoplerythrinus unitaeniatus (Agassiz, 1829) CG VA BO DA AP GU SU FG

Hoplias aimara (Valenciennes, 1847) VA BO FG

Hoplias macrophthalmus (Pellegrin, 1907) CG VA BO DA GU SU FG

Hoplias malabaricus (Bloch, 1794) CG VA BO DA GU SU FG

Hoplias patana (Valenciennes, 1847) FG

Family: Lebiasinidae—Marilyn Weitzman & Stanley H.

Weitzman

Copella arnoldi (Regan, 1912) PA GU SU FG

Copella carsevennensis (Regan, 1912) AP SU FG

Copella compta (Myers, 1927) CG VA BA

Copella eigenmanni (Regan, 1912) BO DA PA AP GU SU FG

Copella metae (Eigenmann, 1914) CG VA BO DA? BA

Copella nattereri (Steindachner, 1876) CG VA BO BA RO GU

Derhamia hoffmannorum Gery & Zarske, 2002 GU

Lebiasina provenzanoi Ardila Rodrıguez, 1999 BO

Lebiasina taphorni Ardila Rodrıguez, 2004 BO

Lebiasina uruyensis Fernandez-Yepez, 1967 BO

Lebiasina yuruaniensis Ardila Rodrıguez, 2000 BO

NUMBER 17 35

Nannostomus anduzei Fernandez & Weitzman, 1987 VA BO

Nannostomus beckfordi Gunther, 1872 AP GU SU FG

Nannostomus bifasciatus Hoedeman, 1954 SU FG

Nannostomus digrammus (Fowler, 1913) BA RO GU

Nannostomus espei (Meinken, 1956) GU

Nannostomus harrisoni (Eigenmann, 1909) GU

Nannostomus marginatus Eigenmann, 1909 CG VA BO DA GU SU

Nannostomus marilynae Weitzman & Cobb, 1975 CG VA BA

Nannostomus minimus Eigenmann, 1909 GU

Nannostomus trifasciatus Steindachner, 1876 RO? GU

Nannostomus unifasciatus Steindachner, 1876 CG VA BO DA GU GU

Piabucina unitaeniata Gunther, 1864 BO GU

Pyrrhulina filamentosa Valenciennes, 1847 VA BO DA GU SU FG

Pyrrhulina lugubris Eigenmann, 1922 CG VA BO DA

Pyrrhulina semifasciata Steindachner, 1876 RO GU

Pyrrhulina stoli Boeseman, 1953 GU SU

Family: Ctenoluciidae—Richard P. Vari

Boulengerella cuvieri (Agassiz, 1829) CG VA BO DA BA AP FG

Boulengerella lateristriga (Boulenger, 1895) VA BA

Boulengerella lucius (Cuvier, 1816) CG VA BO BA PA AP

Boulengerella maculata (Valenciennes, 1850) CG VA BO DA

Boulengerella xyrekes Vari, 1995 VA BO

Order: Siluriformes

Family: Cetopsidae—Richard P. Vari

Cetopsidium ferreirai Vari, Ferraris & de Pinna, 2005 PA

Cetopsidium minutum Vari, Ferraris & de Pinna, 2005 GU SU FG

Cetopsidium morenoi Vari, Ferraris & de Pinna, 2005 VA BO

Cetopsidium orientale Vari, Ferraris & de Pinna, 2005 AP GU SU FG

Cetopsidium pemon Vari, Ferraris & de Pinna, 2005 BO RO GU

Cetopsidium roae Vari, Ferraris & de Pinna, 2005 GU

Cetopsis coecutiens (Lichtenstein, 1819) CG VA BO

Cetopsis orinoco Vari, Ferraris & de Pinna, 2005 BO

Denticetopsis iwokrama Vari, Ferraris & de Pinna, 2005 GU

Denticetopsis macilenta Vari, Ferraris & de Pinna, 2005 GU

Denticetopsis praecox Vari, Ferraris & de Pinna, 2005 VA

Denticetopsis royeroi Ferraris, 1996 VA

Denticetopsis sauli Ferraris, 1996 VA

Helogenes castaneus (Dahl, 1960) CG

Helogenes marmoratus Gunther, 1863 VA BA GU SU FG

Helogenes uruyensis Fernandez-Yepez, 1967 BO

Family: Aspredinidae—John P. Friel

Acanthobunocephalus nicoi Friel, 1995 VA BO

Amaralia hypsiura (Kner, 1855) GU

Aspredinichthys filamentosus (Valenciennes, 1840) DA GU SU FG

Aspredinichthys tibicen (Valenciennes, 1840) DA GU SU FG

Aspredo aspredo (Linnaeus, 1758) DA GU SU FG

Bunocephalus aleuropsis Cope, 1870 VA BO

Bunocephalus amaurus Eigenmann, 1912 VA BO GU SU FG


Bunocephalus chamaizelus Eigenmann, 1912 GU

Bunocephalus verrucosus (Walbaum, 1792) GU SU

Ernstichthys anduzei Fernandez-Yepez, 1953 VA BO

Hoplomyzon sexpapilostoma Taphorn & Marrero, 1990 VA BO

Platystacus cotylephorus Bloch, 1794 DA GU SU FG

Pseudobunocephalus lundbergi Friel, 2008 BO

Pterobunocephalus depressus (Haseman, 1911) VA BO

Family: Trichomycteridae—Wolmar Wosiacki

Ammoglanis pulex de Pinna & Winemiller, 2000 VA BO

Glanapteryx anguilla Myers, 1927 VA BO BA

Glanapteryx niobium de Pinna, 1998 BA

Haemomaster venezuelae Myers, 1927 VA BO

Henonemus taxistigmus (Fowler, 1914) GU

Henonemus triacanthopomus DoNascimiento & Provenzano, 2006 DA

Ituglanis amazonicus (Steindachner, 1882) FG

Ituglanis gracilior (Eigenmann, 1912) GU

Ituglanis metae (Eigenmann, 1917) CG VA? BO?

Ituglanis nebulosus de Pinna & Keith, 2003 FG

Megalocentor echthrus de Pinna & Britski, 1991 VA BO

Ochmacanthus alternus Myers, 1927 CG VA BO DA

Ochmacanthus flabelliferus Eigenmann, 1912 BO? GU

Ochmacanthus orinoco Myers, 1927 VA BO

Ochmacanthus reinhardtii (Steindachner, 1882) FG

Paracanthopoma parva Giltay, 1935 RO

Pseudostegophilus haemomyzon (Myers, 1942) VA BO

Pygidianops cuao Schaefer, Provenzano, de Pinna & Baskin,

2005

VA

Pygidianops magoi Schaefer, Provenzano, de Pinna & Baskin,

2005

BO DA

Schultzichthys bondi (Myers, 1942) VA BO

Stauroglanis gouldingi de Pinna, 1989 BA

Stegophilus septentrionalis Myers, 1927 VA BO

Trichomycterus celsae Lasso & Provenzano, 2002 BO

Trichomycterus conradi (Eigenmann, 1912) BO? GU

Trichomycterus guianensis (Eigenmann, 1909) VA BO GU FG

Trichomycterus lewi Lasso & Provenzano, 2002 BO

Trichomycterus santanderensis Castellanos-Morales, 2007 CG

Typhlobelus guacamaya Schaefer, Provenzano, de Pinna &

Baskin, 2005

VA

Typhlobelus lundbergi Schaefer, Provenzano, de Pinna & Baskin,

2005

BO DA

Vandellia beccarii Di Caporiacco, 1935 CG VA BO DA GU

Vandellia sanguinea Eigenmann, 1917 VA BO GU

Family: Callichthyidae—Roberto E. Reis

Callichthys callichthys (Linnaeus, 1758) CG VA BO DA GU SU FG

Callichthys serralabium Lehmann & Reis, 2004 VA BA

Corydoras aeneus (Gill, 1858) CG VA BO GU SU FG

Corydoras amapaensis Nijssen, 1972 AP FG

Corydoras approuaguensis Nijssen & Isbrucker, 1983 FG

Corydoras axelrodi Rossel, 1962 CG VA BO

NUMBER 17 37

Corydoras baderi Geisler, 1969 PA SU

Corydoras bicolor Nijssen & Isbrucker, 1967 SU

Corydoras blochi Nijssen, 1971 VA BO RO GU

Corydoras boehlkei Nijssen & Isbrucker, 1982 VA BO

Corydoras boesemani Nijssen & Isbrucker, 1967 SU

Corydoras bondi Gosline, 1940 VA BO GU SU

Corydoras breei Isbrucker & Nijssen, 1992 SU

Corydoras brevirostris Fraser-Brunner, 1947 VA BO SU

Corydoras concolor Weitzman, 1961 VA BO

Corydoras condiscipulus Nijssen & Isbrucker, 1980 AP FG

Corydoras coppenamensis Nijssen, 1970 SU

Corydoras delphax Nijssen & Isbrucker, 1983 CG VA

Corydoras ephippifer Nijssen, 1972 AP

Corydoras filamentosus Nijssen & Isbrucker, 1983 SU

Corydoras geoffroy La Cepede, 1803 SU FG

Corydoras griseus Holly, 1940 GU

Corydoras guianensis Nijssen, 1970 SU FG

Corydoras habrosus Weitzman, 1960 CG? VA BO

Corydoras heteromorphus Nijssen, 1970 SU

Corydoras melanistius Regan, 1912 GU SU FG

Corydoras melini Lonnberg & Rendahl, 1930 CG VA? BO?

Corydoras metae Eigenmann, 1914 CG VA BO

Corydoras nanus Nijssen & Isbrucker, 1967 SU FG

Corydoras oiapoquensis Nijssen, 1972 AP FG

Corydoras osteocarus Bohlke, 1951 VA BO SU

Corydoras oxyrhynchus Nijssen & Isbrucker, 1967 SU

Corydoras potaroensis Myers, 1927 GU

Corydoras punctatus (Bloch, 1794) SU FG

Corydoras sanchesi Nijssen & Isbrucker, 1967 SU

Corydoras saramaccensis Nijssen, 1970 SU

Corydoras septentrionalis Gosline, 1940 CG? VA BO

Corydoras sipaliwini Hoedeman, 1965 GU SU

Corydoras solox Nijssen & Isbrucker, 1983 AP FG

Corydoras spilurus Norman, 1926 SU FG

Corydoras surinamensis Nijssen, 1970 SU

Corydoras trilineatus Cope, 1872 SU

Hoplosternum littorale (Hancock, 1828) CG VA BO DA GU SU FG

Megalechis picta (Muller & Troschel, 1848) VA BO DA BA GU

Megalechis thoracata (Valenciennes, 1840) VA BO GU SU FG

Family: Loricariidae

Subfamily: Lithogeninae—Carl J. Ferraris, Jr.

Lithogenes villosus Eigenmann, 1909 GU

Lithogenes wahari Schaefer & Provenzano, 2008 VA

Subfamily: Hypoptopomatinae—Scott A. Schaefer

Acestridium dichromum Retzer, Nico & Provenzano, 1999 VA

Acestridium martini Retzer, Nico & Provenzano, 1999 VA BO

Hypoptopoma guianense Boeseman, 1974 GU SU

Nannoptopoma spectabile (Eigenmann, 1914) CG? VA? BO


Niobichthys ferrarisi Schaefer & Provenzano, 1998 VA

Otocinclus huaorani Schaefer, 1997 CG

Otocinclus mariae Fowler, 1940 BA

Otocinclus vittatus Regan, 1904 CG? VA BO

Oxyropsis acutirostra Miranda Ribeiro, 1951 CG VA BO

Parotocinclus britskii Boeseman, 1974 VA BO GU SU

Parotocinclus collinsae Schmidt & Ferraris, 1985 GU

Parotocinclus eppleyi Schaefer & Provenzano, 1993 VA BO

Parotocinclus polyochrus Schaefer, 1988 VA

Subfamily: Loricariinae—Carl J. Ferraris, Jr.

Cteniloricaria platystoma (Gunther, 1868) SU

Farlowella nattereri Steindachner, 1910 GU

Farlowella oxyrryncha (Kner, 1853) VA BO

Farlowella reticulata Boeseman, 1971 GU SU FG

Farlowella rugosa Boeseman, 1971 GU SU FG

Farlowella vittata Myers, 1942 CG VA BO DA

Harttia fowleri (Pellegrin, 1908) AP FG

Harttia guianensis Rapp Py-Daniel & Oliveira, 2001 SU FG

Harttia maculata (Boeseman, 1971) SU FG

Harttia merevari Provenzano, Machado-Allison, Chernoff,

Willink & Petry, 2005

BO

Harttia surinamensis Boeseman, 1971 SU

Harttia trombetensis Rapp Py-Daniel & Oliveira, 2001 PA

Harttiella crassicauda (Boeseman, 1953) SU

Hemiloricaria castroi Isbrucker & Nijssen, 1984 PA

Hemiloricaria eigenmanni (Pellegrin, 1908) VA BO

Hemiloricaria fallax (Steindachner, 1915) RO GU

Hemiloricaria formosa Isbrucker & Njissen, 1979 CG VA BO DA

Hemiloricaria platyura (Muller & Troschel, 1848) GU FG

Hemiloricaria stewarti (Eigenmann, 1909) GU SU FG

Hemiodontichthys acipenserinus (Kner, 1853) AP GU FG

Limatulichthys griseus (Eigenmann, 1909) CG VA BO DA GU

Loricaria cataphracta Linnaeus, 1758 GU SU FG

Loricaria lundbergi Thomas & Rapp Py-Daniel, 2008 VA

Loricaria nickeriensis Isbrucker, 1979 SU FG

Loricaria parnahybae Steindachner, 1907 AP? FG

Loricaria simillima Regan, 1904 VA BO

Loricaria spinulifera Thomas & Rapp Py-Daniel, 2008 RO

Loricariichthys maculatus (Bloch, 1794) SU

Loricariichthys microdon (Eigenmann, 1909) GU

Metaloricaria nijsseni (Boeseman, 1976) SU

Metaloricaria paucidens Isbrucker, 1975 SU FG

Pseudoloricaria laeviuscula (Valenciennes, 1840) RO

Reganella depressa (Kner, 1853) RO

Sturisoma monopelte Fowler, 1914 GU

Subfamily: Hypostominae—Jonathan W. Armbruster & Claude

Weber

Acanthicus hystrix Spix & Agassiz, 1829 VA BO DA

Ancistrus hoplogenys (Gunther, 1864) GU

NUMBER 17 39

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

Ancistrus lithurgicus Eigenmann, 1912 GU

Ancistrus macrophthalmus (Pellegrin, 1912) VA BO

Ancistrus nudiceps (Muller & Troschel, 1848) GU

Ancistrus temminckii (Valenciennes, 1840) SU

Ancistrus triradiatus Eigenmann, 1918 CG VA BO DA

Baryancistrus beggini Lujan, Arce & Armbruster, 2009 CG VA

Baryancistrus demantoides Werneke, Sabaj, Lujan &

Armbruster, 2005

VA

Baryancistrus niveatus (Castelnau, 1855) PA

Chaetostoma jegui Rapp Py-Daniel, 1991 RO

Chaetostoma vasquezi Lasso & Provenzano, 1998 VA BO

Corymbophanes andersoni Eigenmann, 1909 GU

Corymbophanes kaiei Armbruster & Sabaj, 2000 GU

Dekeyseria niveata (La Monte, 1929) VA

Dekeyseria pulcher (Steindachner, 1915) CG VA BA?

Dekeyseria scaphirhyncha (Kner, 1854) BA

Exastilithoxus fimbriatus (Steindachner, 1915) BO

Exastilithoxus hoedemani Isbrucker & Nijssen, 1985 BA

Hemiancistrus guahiborum Werneke, Armbruster, Lujan &

Taphorn, 2005

VA BO

Hemiancistrus medians (Kner, 1854) SU FG

Hemiancistrus sabaji Armbruster, 2003 VA GU

Hemiancistrus subviridis Werneke, Sabaj, Lujan & Armbruster,

2005

VA

Hypancistrus contradens Armbruster, Lujan & Taphorn, 2007 VA

Hypancistrus debilittera Armbruster, Lujan & Taphorn, 2007 VA

Hypancistrus furunculus Armbruster, Lujan & Taphorn, 2007 VA

Hypancistrus inspector Armbruster, 2002 VA BO

Hypancistrus lunaorum Armbruster, Lujan & Taphorn, 2007 VA

Hypostomus coppenamensis Boeseman, 1969 SU

Hypostomus corantijni Boeseman, 1968 SU

Hypostomus crassicauda Boeseman, 1968 SU

Hypostomus gymnorhynchus (Norman, 1926) SU FG

Hypostomus hemicochliodon Armbruster, 2003 VA BO

Hypostomus hemiurus (Eigenmann, 1912) GU

Hypostomus macrophthalmus Boeseman, 1968 SU

Hypostomus macushi Armbruster & de Souza, 2005 GU

Hypostomus micromaculatus Boeseman, 1968 SU

Hypostomus nematopterus Isbrucker & Nijssen, 1984 FG

Hypostomus nickeriensis Boeseman, 1969 SU

Hypostomus occidentalis Boeseman, 1968 SU

Hypostomus paucimaculatus Boeseman, 1968 SU

Hypostomus plecostomoides (Eigenmann, 1922) BO

Hypostomus plecostomus (Linnaeus, 1758) GU SU

Hypostomus pseudohemiurus Boeseman, 1968 SU

Hypostomus rhantos Armbruster, Tansey & Lujan, 2007 VA

Hypostomus saramaccensis Boeseman, 1968 SU

Hypostomus sculpodon Armbruster, 2003 VA


Hypostomus sipaliwini Boeseman, 1968 SU

Hypostomus squalinus (Jardine, 1841) RO GU

Hypostomus surinamensis Boeseman, 1968 SU

Hypostomus tapanahoniensis Boeseman, 1969 SU

Hypostomus taphorni (Lilyestrom, 1984) BO GU

Hypostomus tenuis (Boeseman, 1968) SU?

Hypostomus ventromaculatus Boeseman, 1968 SU FG

Hypostomus villarsi (Lutken, 1874) VA BO

Hypostomus watwata Hancock, 1828 DA GU SU FG

Lasiancistrus schomburgkii (Gunther, 1864) GU

Lasiancistrus tentaculatus Armbruster, 2005 VA BO

Leporacanthicus galaxias Isbrucker & Nijssen, 1989 VA BO

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

Lithoxus boujardi Muller & Isbrucker, 1993 FG

Lithoxus bovallii (Regan, 1906) GU

Lithoxus jantjae Lujan, 2008 VA

Lithoxus lithoides Eigenmann, 1910 GU SU

Lithoxus pallidimaculatus Boeseman, 1982 SU

Lithoxus planquettei Boeseman, 1982 FG

Lithoxus stocki Nijssen & Isbrucker, 1990 FG

Lithoxus surinamensis Boeseman, 1982 SU

Neblinichthys pilosus Ferraris, Isbrucker & Nijssen, 1986 VA

Neblinichthys roraima Provenzano, Lasso & Ponte, 1995 BO

Neblinichthys yaravi (Steindachner, 1915) BO

Panaque maccus Schaefer & Stewart, 1993 VA BO

Panaque nigrolineatus (Peters, 1877) CG VA BO

Peckoltia braueri (Eigenmann, 1912) GU

Peckoltia brevis (La Monte, 1935) CG?

Peckoltia cavatica Armbruster & Werneke, 2005 GU

Peckoltia lineola Armbruster, 2008 CG VA

Peckoltia vittata (Steindachner, 1881) CG? VA PA

Pseudacanthicus fordii (Gunther, 1868) SU

Pseudacanthicus leopardus (Fowler, 1914) GU

Pseudacanthicus serratus (Valenciennes, 1840) SU FG

Pseudancistrus barbatus (Valenciennes, 1840) GU SU FG

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

Pseudancistrus coquenani (Steindachner, 1915) BO

Pseudancistrus corantijniensis De Chambrier & Montoya-

Burgos, 2008

SU

Pseudancistrus depressus (Gunther, 1868) SU

Pseudancistrus guentheri (Regan, 1904) GU

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

Pseudancistrus macrops (Lutken, 1874) SU

Pseudancistrus megacephalus (Gunther, 1868) GU SU

Pseudancistrus niger (Norman, 1926) FG

Pseudancistrus nigrescens Eigenmann, 1912 GU

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

Pseudancistrus pectegenitor Lujan, Armbruster & Sabaj, 2007 VA

Pseudancistrus reus Armbruster & Taphorn, 2008 BO

NUMBER 17 41

Pseudancistrus sidereus Armbruster, 2004 VA

Pseudancistrus yekuana Lujan, Armbruster & Sabaj, 2007 VA

Pseudolithoxus anthrax (Armbruster & Provenzano, 2000) VA BO

Pseudolithoxus dumus (Armbruster & Provenzano, 2000) VA BO

Pseudolithoxus nicoi (Armbruster & Provenzano, 2000) VA

Pseudolithoxus tigris (Armbruster & Provenzano, 2000) VA BO

Pterygoplichthys gibbiceps (Kner, 1854) CG VA BO

Pterygoplichthys multiradiatus (Hancock, 1828) VA BO GU?

Family: Pseudopimelodidae—Oscar A. Shibbata

Batrochoglanis raninus (Valenciennes, 1840) GU SU FG

Batrochoglanis villosus (Eigenmann, 1912) VA BO GU SU

Cephalosilurus albomarginatus (Eigenmann, 1912) GU

Cephalosilurus nigricaudus (Mees, 1974) SU

Microglanis poecilus Eigenmann, 1912 GU FG

Microglanis secundus Mees, 1974 CG? VA BO GU SU

Pseudopimelodus bufonius (Valenciennes, 1840) CG? VA BO GU SU FG

Family: Heptapteridae—Updated from Cloffsca account (Flavio

A. Bockmann & Gizelani M. Guazzeli)

Brachyglanis frenata Eigenmann, 1912 VA BO BA GU

Brachyglanis magoi Fernandez-Yepez, 1967 VA BO

Brachyglanis melas Eigenmann, 1912 GU

Brachyglanis microphthalmus Bizerril, 1991 PA

Brachyglanis phalacra Eigenmann, 1912 GU

Brachyrhamdia heteropleura (Eigenmann, 1912) GU SU

Brachyrhamdia imitator Myers, 1927 VA BO

Cetopsorhamdia insidiosa (Steindachner, 1917) RO

Cetopsorhamdia orinoco Schultz, 1944 BO

Chasmocranus brevior Eigenmann, 1912 GU SU FG

Chasmocranus chimantanus Inger, 1956 BO

Chasmocranus longior Eigenmann, 1912 VA BO BA GU SU

Chasmocranus surinamensis (Bleeker, 1862) SU

Gladioglanis machadoi Ferraris & Mago-Leccia, 1989 VA BO BA?

Goeldiella eques (Muller & Troschel, 1848) CG VA BO BA? GU

Heptapterus bleekeri Boeseman, 1953 AP? SU FG

Heptapterus tapanahoniensis Mees, 1967 SU FG

Heptapterus tenuis Mees, 1986 FG

Imparfinis hasemani Steindachner, 1917 RO

Imparfinis pijpersi (Hoedeman, 1961) SU

Imparfinis pristos Mees & Cala, 1989 CG VA BO

Imparfinis pseudonemacheir Mees & Cala, 1989 CG VA BO

Leptorhamdia essequibensis (Eigenmann, 1912) GU

Leptorhamdia marmorata Myers, 1928 CG? VA BO BA

Mastiglanis asopos Bockmann, 1994 VA BO PA

Medemichthys guayaberensis (Dahl, 1961) CG

Myoglanis aspredinoides DoNascimiento & Lundberg, 2005 VA

Myoglanis potaroensis Eigenmann, 1912 GU

Nemuroglanis pauciradiatus Ferraris, 1988 CG VA BO BA

Phenacorhamdia anisura (Mees, 1987) VA BO

Phenacorhamdia macarenensis Dahl, 1961 CG


Phenacorhamdia provenzanoi DoNascimiento & Milani, 2008 VA BO

Pimelodella altipinnis (Steindachner, 1864) GU

Pimelodella cristata (Muller & Troschel, 1848) GU FG

Pimelodella cruxenti Fernandez-Yepez, 1950 VA BO

Pimelodella figueroai Dahl, 1961 CG

Pimelodella geryi Hoedeman, 1961 SU

Pimelodella linami Schultz, 1944 VA BO

Pimelodella macturki Eigenmann, 1912 GU SU FG

Pimelodella megalops Eigenmann, 1912 GU FG

Pimelodella pallida Dahl, 1961 CG

Pimelodella procera Mees, 1983 FG

Pimelodella wesselii (Steindachner, 1877) GU

Rhamdella leptosoma Fowler, 1914 GU

Rhamdia foina (Muller & Troschel, 1848) RO PA GU

Rhamdia laukidi Bleeker, 1858 CG VA BO GU

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

Rhamdia quelen (Quoy & Gaimard, 1824) CG VA BO DA GU SU FG

Family: Pimelodidae—John G. Lundberg

Brachyplatystoma filamentosum (Lichtenstein, 1819) CG VA BO DA GU SU FG

Brachyplatystoma juruense (Boulenger, 1898) CG? VA BO DA

Brachyplatystoma platynemum (Boulenger, 1898) CG VA BO DA

Brachyplatystoma rousseauxii (Castelnau, 1855) BO DA FG

Brachyplatystoma vaillantii (Valenciennes, 1840) CG VA BO DA GU SU FG

Calophysus macropterus (Lichtenstein, 1819) CG VA BO DA PA

Duopalatinus peruanus Eigenmann & Allen, 1942 BO

Exallodontus aguanai Lundberg, Mago-Leccia & Nass, 1991 BO DA

Hemisorubim platyrhynchos (Valenciennes, 1840) CG VA BO DA GU SU FG

Hypophthalmus edentatus Spix & Agassiz, 1829 VA BO DA GU SU

Hypophthalmus fimbriatus Kner, 1858 BO

Hypophthalmus marginatus Valenciennes, 1840 SU FG

Leiarius marmoratus (Gill, 1870) CG VA BO

Leiarius pictus (Muller & Troschel, 1849) CG? VA BO GU

Megalonema amaxanthum Lundberg & Dahdul, 2008 GU

Megalonema platycephalum Eigenmann, 1912 CG VA BO GU

Phractocephalus hemioliopterus (Bloch & Schneider, 1801) CG VA BO DA GU

Pimelodina flavipinnis Steindachner, 1877 BO DA

Pimelodus albofasciatus Mees, 1974 CG VA BO BA SU

Pimelodus blochii Valenciennes, 1840 CG VA BO DA GU SU FG

Pimelodus ornatus Kner, 1858 CG VA BO GU SU FG

Pimelodus pictus Steindachner, 1877 BO

Pinirampus pirinampu (Spix & Agassiz, 1829) CG VA BO DA GU

Platynematichthys notatus (Jardine, 1841) CG VA BO DA

Platysilurus mucosus (Vaillant, 1880) CG VA BO DA

Propimelodus eigenmanni (van der Stigchel, 1946) AP FG

Pseudoplatystoma fasciatum (Linnaeus, 1766) GU SU FG

Pseudoplatystoma metaense Buitrago-Suarez & Burr, 2007 BO DA

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

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

NUMBER 17 43

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

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

Zungaro zungaro (Humboldt, 1821) CG VA BO DA BA GU

Family: Doradidae—Mark H. Sabaj Perez

Acanthodoras cataphractus (Linnaeus, 1758) CG VA BO? BA RO PA AP GU SU FG

Acanthodoras spinosissimus (Eigenmann & Eigenmann, 1888) CG VA BO? DA? BA RO GU

Agamyxis albomaculatus (Peters, 1877) CG VA BO DA BA?

Amblydoras affinis (Kner, 1855) RO GU

Amblydoras bolivarensis (Fernandez-Yepez, 1968) CG VA BO DA? BA?

Amblydoras gonzalezi (Fernandez-Yepez, 1968) CG VA BO DA? BA?

Anadoras regani (Steindachner, 1908) PA AP FG

Anduzedoras oxyrhynchus (Valenciennes, 1821) CG VA BA

Centrodoras hasemani (Steindachner, 1915) CG? VA BA

Doras carinatus (Linnaeus, 1766) BO AP GU SU FG

Doras micropoeus (Eigenmann, 1912) GU SU FG

Doras phlyzakion Sabaj Perez & Birindelli, 2008 RO

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

Leptodoras cataniai Sabaj, 2005 VA BA RO?

Leptodoras copei (Fernandez-Yepez, 1968) CG VA BO BA RO? PA?

Leptodoras hasemani (Steindachner, 1915) CG VA BO DA BA? RO GU

Leptodoras linnelli Eigenmann, 1912 VA BO BA? RO GU

Leptodoras praelongus (Myers & Weitzman, 1956) CG VA BA RO PA?

Leptodoras rogersae Sabaj, 2005 CG VA BO DA

Megalodoras guayoensis (Fernandez-Yepez, 1968) CG? VA? BO DA

Megalodoras uranoscopus (Eigenmann & Eigenmann, 1888) CG? VA? BA? RO? PA? GU

Nemadoras leporhinus (Eigenmann, 1912) CG VA BO BA? RO PA GU

Nemadoras trimaculatus (Boulenger, 1898) CG VA BO BA RO GU

Opsodoras morei (Steindachner, 1881) CG VA BO? BA RO? PA? GU

Opsodoras ternetzi Eigenmann, 1925 CG VA BO DA BA RO PA GU

Orinocodoras eigenmanni Myers, 1927 BO DA

Oxydoras niger (Valenciennes, 1821) CG? VA? BA? RO? GU

Oxydoras sifontesi Fernandez-Yepez, 1968 CG? VA? BO DA

Physopyxis ananas Sousa & Rapp Py-Daniel 2005 CG VA BA RO PA GU

Physopyxis cristata Sousa & Rapp Py-Daniel 2005 BA RO?

Platydoras armatulus (Valenciennes 1840) BO DA

Platydoras costatus (Linnaeus, 1758) SU FG

Platydoras hancockii (Valenciennes 1840) CG VA BO? BA RO GU

Pterodoras granulosus (Valenciennes, 1821) CG? VA? BA? RO? PA? GU SU

Pterodoras rivasi (Fernandez-Yepez, 1950) CG? VA? BO DA

Rhinodoras armbrusteri Sabaj, Taphorn & Castillo, 2008 GU

Rhynchodoras woodsi Glodek, 1976 RO GU

Scorpiodoras heckelii (Kner, 1855) CG VA BO BA

Trachydoras brevis (Kner, 1853) BA RO GU

Trachydoras microstomus (Eigenmann, 1912) CG VA BO DA? BA RO PA GU

Family: Auchenipteridae—Carl J. Ferraris, Jr.

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

Ageneiosus marmoratus Eigenmann, 1912 GU SU

Ageneiosus piperatus (Eigenmann, 1912) RO GU


Ageneiosus polystictus Steindachner, 1915 RO

Ageneiosus ucayalensis Castelnau, 1855 VA BO DA GU SU

Ageneiosus vittatus Steindachner, 1908 BO DA

Asterophysus batrachus Kner, 1858 VA BO BA?

Auchenipterus ambyiacus Fowler, 1915 CG? VA BO DA GU

Auchenipterus brevior Eigenmann, 1912 GU

Auchenipterus demerarae Eigenmann, 1912 BO GU

Auchenipterus dentatus Valenciennes, 1840 SU FG

Auchenipterus nuchalis (Spix & Agassiz, 1829) RO? FG

Centromochlus concolor (Mees, 1974) SU

Centromochlus existimatus Mees, 1974 VA BO

Centromochlus punctatus (Mees, 1974) SU

Centromochlus reticulatus (Mees, 1974) GU

Entomocorus gameroi Mago-Leccia, 1984 VA BO DA

Gelanoglanis nanonocticolus Soares-Porto, Walsh, Nico & Netto,

1999

VA BO

Glanidium leopardum (Hoedeman, 1961) GU SU FG

Liosomadoras oncinus (Jardine, 1841) RO

Pseudepapterus gracilis Ferraris & Vari, 2000 BO

Tatia brunnea Mees, 1974 SU FG

Tatia creutzbergi (Boeseman, 1953) SU

Tatia galaxias Mees, 1974 VA BO DA

Tatia gyrina (Eigenmann & Allen, 1942) SU

Tatia intermedia (Steindachner, 1877) GU SU FG

Tatia meesi Sarmento-Soares & Martins-Pinheiro, 2008 GU

Tatia musaica Royero, 1992 VA BO

Tatia nigra Sarmento-Soares & Martins-Pinheiro, 2008 PA

Tatia strigata Soares-Porto, 1995 VA BO BA

Tetranematichthys wallacei Vari & Ferraris, 2006 VA BO BA

Trachelyichthys decaradiatus Mees, 1974 GU

Trachelyopterichthys anduzei Ferraris & Fernandez, 1987 VA BO

Trachelyopterichthys taeniatus (Kner, 1858) CG VA BO BA?

Trachelyopterus ceratophysus (Kner, 1858) RO

Trachelyopterus coriaceus Valenciennes, 1840 FG

Trachelyopterus galeatus (Linnaeus, 1766) SU FG

Trachycorystes trachycorystes Valenciennes, 1840 GU

Order: Gymnotiformes

Family: Gymnotidae—Ricardo Campos-da-Paz

Electrophorus electricus (Linnaeus, 1766) CG VA BO DA GU SU FG

Gymnotus anguillaris Hoedeman, 1962 SU FG

Gymnotus carapo Linnaeus, 1758 VA BO DA FG

Gymnotus cataniapo Mago-Leccia, 1994 BO SU

Gymnotus coropinae Hoedeman, 1962 GU SU

Gymnotus pedanopterus Mago-Leccia, 1994 VA BO BA

Gymnotus stenoleucus Mago-Leccia, 1994 VA BO DA

Family: Sternopygidae—James S. Albert

Archolaemus blax Korringa, 1970 RO AP? FG

Distocyclus conirostris (Eigenmann & Allen, 1942) BO DA

NUMBER 17 45

Eigenmannia limbata (Schreiner & Miranda Ribeiro, 1903) BO

Eigenmannia macrops (Boulenger, 1897) BA GU

Eigenmannia nigra Mago-Leccia, 1994 CG VA BO BA GU

Eigenmannia virescens (Valenciennes, 1842) CG VA BO DA BA AP GU SU FG

Rhabdolichops caviceps (Fernandez-Yepez, 1968) BO

Rhabdolichops eastwardi Lundberg & Mago-Leccia, 1986 BO DA

Rhabdolichops electrogrammus Lundberg & Mago-Leccia, 1986 BO DA BA RO

Rhabdolichops jegui Keith & Meunier, 2000 FG

Rhabdolichops stewarti Lundberg & Mago-Leccia, 1986 VA BO

Rhabdolichops troscheli (Kaup, 1856) BO DA

Rhabdolichops zareti Lundberg & Mago-Leccia, 1986 VA BO DA

Sternopygus astrabes Mago-Leccia, 1994 CG? VA BO BA

Sternopygus macrurus (Bloch & Schneider, 1801) CG VA BO DA BA? RO? PA? AP? GU SU FG

Family: Rhamphichthyidae—Carl J. Ferraris, Jr.

Gymnorhamphichthys hypostomus Ellis, 1912 CG VA DA

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

Iracema caiana Triques, 1996 BA RO

Rhamphichthys apurensis (Fernandez-Yepez, 1968) VA BO DA

Rhamphichthys rostratus (Linnaeus, 1766) GU SU

Family: Hypopomidae—James S. Albert

Brachyhypopomus beebei (Schultz, 1944) CG VA BO DA GU SU

Brachyhypopomus brevirostris (Steindachner, 1868) CG VA BO DA GU SU

Brachyhypopomus pinnicaudatus (Hopkins, 1991) BO GU SU

Hypopomus artedi (Kaup, 1856) PA? AP? GU SU FG

Hypopygus lepturus Hoedeman, 1962 CG VA BO DA BA? SU FG

Hypopygus neblinae Mago-Leccia, 1994 CG VA BO DA

Microsternarchus bilineatus Fernandez-Yepez, 1968 CG VA BO BA

Racenisia fimbriipinna Mago-Leccia, 1994 VA BO

Steatogenys duidae (La Monte, 1929) VA BO

Steatogenys elegans (Steindachner, 1880) CG VA BO DA

Family: Apteronotidae—James S. Albert

Adontosternarchus clarkae Mago-Leccia, Lundberg & Baskin,

1985

CG VA BA RO

Adontosternarchus devenanzii Mago-Leccia, Lundberg & Baskin,

1985

BO DA

Adontosternarchus sachsi (Peters, 1877) BO DA

Apteronotus albifrons (Linnaeus, 1766) VA BO DA GU SU FG

Apteronotus leptorhynchus (Ellis, 1912) VA BO DA BA GU SU FG

Compsaraia compsus (Mago-Leccia, 1994) BO DA

Megadontognathus cuyuniense Mago-Leccia, 1994 BO

Platyurosternarchus crypticus de Santana &Vari, 2009 RO GU

Platyurosternarchus macrostomus (Gunther, 1870) VA BO DA GU

Porotergus gymnotus Ellis, 1912 GU FG

Sternarchella orthos Mago-Leccia, 1994 VA BO DA

Sternarchella sima Starks, 1913 BO DA

Sternarchorhamphus muelleri (Steindachner, 1881) BO DA

Sternarchorhynchus gnomus de Santana & Taphorn, 2006 BO

Sternarchorhynchus oxyrhynchus (Muller & Troschel, 1849) BO GU FG

Sternarchorhynchus roseni Mago-Leccia, 1994 BO DA


Order: Cyprinodontiformes

Family: Rivulidae—Wilson J. E. M. Costa

Austrofundulus rupununi Hrbek, Taphorn &Thomerson, 2005 GU

Austrofundulus transilis Myers, 1932 VA BO

Gnatholebias hoignei (Thomerson, 1974) VA BO

Gnatholebias zonatus (Myers, 1935) VA BO

Kryptolebias sepia Vermeulen & Hrbek, 2005 SU

Micromoema xiphophora (Thomerson & Taphorn, 1992) VA BO

Rachovia maculipinnis (Radda, 1964) BO DA

Rachovia stellifer (Thomerson & Turner, 1973) VA BO

Renova oscari Thomerson & Taphorn, 1995 VA BO

Rivulus agilae Hoedeman, 1954 GU SU FG

Rivulus altivelis Huber, 1992 CG

Rivulus amphoreus Huber, 1979 SU

Rivulus breviceps Eigenmann, 1909 GU

Rivulus caurae Radda, 2004 BO

Rivulus cladophorus Huber, 1991 FG

Rivulus corpulentus Thomerson & Taphorn, 1993 CG

Rivulus deltaphilus Seegers, 1983 VA BO DA

Rivulus frenatus Eigenmann, 1912 GU SU

Rivulus gaucheri Keith, Nandrin & Le-Bail 2006 FG

Rivulus geayi Vaillant, 1899 AP FG

Rivulus gransabanae Lasso, Taphorn & Thomerson, 1992 BO

Rivulus holmiae Eigenmann, 1909 GU

Rivulus igneus Huber, 1991 AP FG

Rivulus immaculatus Thomerson, Nico & Taphorn, 1991 BO

Rivulus lanceolatus Eigenmann, 1909 GU

Rivulus lungi Berkenkamp, 1984 FG

Rivulus lyricauda Thomerson, Berkenkamp & Taphorn, 1991 VA BO

Rivulus mahdiaensis Suijker & Collier, 2006 GU

Rivulus manaensis Hoedeman, 1961 FG

Rivulus mazaruni Myers, 1924 GU

Rivulus nicoi Thomerson & Taphorn, 1992 VA BO

Rivulus sape Lasso-Alcala, Taphorn, Lasso & Leon-Mata, 2006 BO

Rivulus stagnatus Eigenmann, 1909 GU SU

Rivulus tecminae Thomerson, Nico & Taphorn, 1992 VA BO

Rivulus torrenticola Vermeulen & Isbrucker, 2000 GU

Rivulus waimacui Eigenmann, 1909 GU

Rivulus xiphidius Huber, 1979 AP FG

Terranatos dolichopterus (Weitzman & Wourms, 1967) VA BO

Family: Poeciliidae—Paulo H. J. Lucinda

Fluviphylax palikur Costa & Le Bail, 1999 AP FG

Fluviphylax pygmaeus (Myers & Carvalho, 1955) CG VA BA PA

Fluviphylax simplex Costa, 1996 PA

Micropoecilia bifurca (Eigenmann, 1909) GU SU FG

Micropoecilia parae (Eigenmann, 1894) AP? GU SU FG

Micropoecilia picta (Regan, 1913) GU SU FG

Poecilia reticulata Peters, 1859 BO DA AP? GU SU FG

NUMBER 17 47

Poecilia vivipara Bloch & Schneider, 1801 DA GU SU FG

Tomeurus gracilis Eigenmann, 1909 DA AP GU SU FG

Order: Beloniformes

Family: Belonidae—Bruce B. Collette

Belonion dibranchodon Collette, 1966 VA BA

Potamorrhaphis guianensis (Jardine, 1843) CG VA BA RO PA GU SU FG

Potamorrhaphis petersi Collette, 1974 CG VA BO BA

Pseudotylosurus microps (Gunther, 1866) VA BO DA GU SU

Family: Hemiramphidae—Bruce B. Collette

Hyporhamphus brederi (Fernandez-Yepez, 1948) VA BO DA

Order: Synbranchiformes

Family: Synbranchidae—Sven O. Kullander

Synbranchus marmoratus Bloch, 1795 CG VA BO DA BA RO AP GU SU FG

Order: Perciformes

Family: Sciaenidae—Ning Labbish Chao & Lilian Casatti

Pachypops fourcroi (Lacpede, 1802) VA BO DA AP GU SU FG

Pachypops pigmaeus Casatti, 2002 RO

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

Pachyurus gabrielensis Casatti, 2001 VA BO BA

Pachyurus schomburgkii Gunther, 1860 VA BO

Petilipinnis grunniens (Jardine, 1843) BO GU

Plagioscion auratus (Castelnau, 1855) BO DA GU SU FG

Plagioscion squamosissimus (Heckel, 1840) CG VA BO DA SU FG

Plagioscion surinamensis (Bleeker, 1973) CG DA SU FG

Family: Polycentridae—Richard P. Vari

Monocirrhus polyacanthus Heckel, 1840 BO RO

Polycentrus schomburgkii Muller & Troschel, 1849 DA AP GU SU FG

Family: Cichlidae—Sven O. Kullander

Acaronia nassa (Heckel, 1840) RO AP GU FG

Acaronia vultuosa Kullander, 1989 CG VA BO BA

Aequidens chimantanus Inger, 1956 BO

Aequidens diadema (Heckel, 1840) CG VA BO BA

Aequidens paloemeuensis Kullander & Nijssen, 1989 SU

Aequidens potaroensis Eigenmann, 1912 GU

Aequidens tetramerus (Heckel, 1840) CG VA BO DA PA AP GU SU FG

Aequidens tubicen Kullander & Ferreira, 1991 PA

Apistogramma angayuara Kullander & Ferreira, 2005 PA

Apistogramma diplotaenia Kullander, 1987 VA BO BA

Apistogramma gibbiceps Meinken, 1969 RO

Apistogramma gossei Kullander, 1982 AP FG

Apistogramma hoignei Meinken, 1965 VA BO

Apistogramma hongsloi Kullander, 1979 VA

Apistogramma iniridae Kullander, 1979 CG

Apistogramma inornata Staeck, 2003 BO

Apistogramma ortmanni (Eigenmann, 1912) BO? GU SU

Apistogramma rupununi Fowler, 1914 RO GU

Apistogramma salpinction Kullander & Ferreira, 2005 PA

Apistogramma steindachneri (Regan, 1908) GU SU


Apistogramma velifera Staeck, 2003 VA

Apistogramma viejita Kullander, 1979 CG

Apistogramma wapisana Romer, Hahn & Conrad, 2006 RO

Astronotus ocellatus (Agassiz, 1831) FG

Biotodoma cupido (Heckel, 1840) GU

Biotodoma wavrini (Gosse, 1963) CG VA BO BA

Biotoecus dicentrarchus Kullander, 1989 CG VA BO DA

Caquetaia spectabilis (Steindachner, 1875) BA RO AP GU

Chaetobranchus flavescens Heckel, 1840 CG VA BO AP GU SU FG

Cichla intermedia Machado-Allison, 1971 VA BO

Cichla jariina Kullander & Ferreira, 2006 PA

Cichla monoculus Spix & Agassiz, 1831 PA AP FG

Cichla nigromaculata Jardine, 1843 VA

Cichla ocellaris Schneider, 1801 RO GU SU FG

Cichla orinocensis Humboldt, 1821 CG VA BO DA

Cichla temensis Humboldt, 1821 CG VA BO DA BA

Cichla thyrorus Kullander & Ferreira, 2006 PA

Cichlasoma amazonarum Kullander, 1983 AP FG

Cichlasoma bimaculatum (Linnaeus, 1758) BO RO GU SU FG

Cichlasoma orinocense Kullander, 1983 CG VA BO

Cleithracara maronii (Steindachner, 1881) DA GU SU FG

Crenicara punctulatum (Gunther, 1863) GU

Crenicichla albopunctata Pellegrin, 1904 GU SU FG

Crenicichla alta Eigenmann, 1912 RO GU

Crenicichla coppenamensis Ploeg, 1987 SU

Crenicichla heckeli Ploeg, 1989 PA

Crenicichla hummelincki Ploeg, 1991 PA

Crenicichla johanna Heckel, 1840 CG VA BO AP GU FG

Crenicichla lugubris Heckel, 1840 BA RO GU SU

Crenicichla multispinosa Pellegrin, 1903 SU FG

Crenicichla nickeriensis Ploeg, 1987 SU

Crenicichla pydanielae Ploeg, 1991 PA

Crenicichla reticulata (Heckel, 1840) GU

Crenicichla saxatilis (Linnaeus, 1758) DA GU SU FG

Crenicichla sipaliwini Ploeg, 1987 SU

Crenicichla ternetzi Norman, 1926 FG

Crenicichla tigrina Ploeg, Jegu & Ferreira, 1991 PA

Crenicichla vaillanti Pellegrin, 1903 GU FG

Crenicichla virgatula Ploeg, 1991 RO

Crenicichla wallacii Regan, 1905 GU

Crenicichla zebrina Montana, Lopez-Fernandez & Taphorn,

2008

VA

Dicrossus filamentosus (Ladiges, 1958) CG VA BO

Dicrossus gladicauda Schindler & Staeck, 2008 CG

Geophagus abalios Lopez-Fernandez & Taphorn, 2004 VA BO

Geophagus brachybranchus Kullander & Nijssen, 1989 GU SU

Geophagus brokopondo Kullander & Nijssen, 1989 SU

Geophagus camopiensis Pellegrin, 1903 AP FG

Geophagus dicrozoster Lopez-Fernandez & Taphorn, 2004 VA BO

NUMBER 17 49

Geophagus gottwaldi Schindler & Staeck, 2006 VA

Geophagus grammepareius Kullander & Taphorn, 1992 VA BO

Geophagus harreri Gosse, 1976 SU FG

Geophagus surinamensis (Bloch, 1791) SU FG

Geophagus taeniopareius Kullander & Royero, 1992 VA BO

Geophagus winemilleri Lopez-Fernandez & Taphorn, 2004 VA

Guianacara (Guianacara) cuyunii Lopez-Fernandez, Taphorn, &

Kullander, 2006

BO

Guianacara (Guianacara) geayi (Pellegrin, 1902) AP FG

Guianacara (Guianacara) owroewefi Kullander & Nijssen, 1989 SU FG

Guianacara (Guianacara) sphenozona Kullander & Nijssen, 1989 GU SU

Guianacara (Guianacara) stergiosi Lopez-Fernandez, Taphorn,

& Kullander, 2006

BO

Guianacara (Oelemaria) oelemariensis Kullander & Nijssen,

1989

SU

Heros severus Heckel, 1840 CG VA BO BA

Hoplarchus psittacus (Heckel, 1840) CG VA BO BA

Hypselecara coryphaenoides (Heckel, 1840) CG VA BO BA

Hypselecara temporalis (Gunther, 1862) AP

Ivanacara adoketa (Kullander & Prada-Pedreros, 1993) BA

Ivanacara bimaculata Eigenmann, 1912 GU

Krobia guianensis (Regan, 1905) GU SU

Krobia itanyi (Puyo, 1943) SU FG

Laetacara fulvipinnis Staeck & Schindler, 2007 CG VA BA

Mazarunia mazarunii Kullander, 1990 GU

Mesonauta egregius Kullander & Silfvergrip, 1991 CG

Mesonauta guyanae Schindler, 1998 RO GU

Mesonauta insignis (Heckel, 1840) CG VA BO BA

Nannacara anomala Regan, 1905 GU SU

Nannacara aureocephalus Allgayer, 1983 FG

Pterophyllum altum Pellegrin, 1903 CG VA

Pterophyllum leopoldi (Gosse, 1963) GU

Pterophyllum scalare (Schultze, 1823) BA AP GU SU FG

Retroculus septentrionalis Gosse, 1971 AP FG

Satanoperca acuticeps (Heckel, 1840) RO

Satanoperca daemon (Heckel, 1840) CG VA BO BA

Satanoperca jurupari (Heckel, 1840) BA PA AP

Satanoperca leucosticta (Muller & Troschel, 1849) GU SU

Satanoperca lilith Kullander & Ferreira, 1988 BA RO PA

Satanoperca mapiritensis (Fernandez-Yepez, 1950) VA BO DA

Uaru fernandezyepezi Stawikowski, 1989 VA

Family: Gobiidae—Sven O. Kullander

Awaous flavus Valenciennes, 1837 DA AP GU SU FG

Ctenogobius claytonii (Meek, 1902) DA

Dormitator maculatus (Bloch, 1792) GU SU FG

Eleotris amplyopsis (Cope, 1871) DA GU SU FG

Eleotris pisonis (Gmelin, 1789) DA GU SU FG

Gobioides broussonnetii La Cepede, 1800 DA FG

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


Gobiomorus dormitor La Cepede, 1800 GU SU

Microphilypnus ternetzi Myers, 1927 VA BO DA

Order: Pleuronectiformes

Family: Achiridae—Robson T. C. Ramos

Achirus achirus (Linnaeus, 1758) GU FG

Achirus novoae Cervigon, 1982 BO DA

Apionichthys dumerili Kaup, 1858 DA PA AP GU SU FG

Apionichthys finis Eigenmann, 1912 RO GU

Apionichthys menezesi Ramos, 2003 VA BO

Hypoclinemus mentalis (Gunther, 1862) VA BO BA RO AP GU

Order: Tetraodontiformes

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

Kullander

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

Order: Lepidosireniformes

Family: Lepidosirenidae—Based on Cloffsca account (Gloria

Arratia)

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

NUMBER 17 51

PHOTOGRAPHIC ATLAS OF FISHES OF THE GUIANA SHIELD

MARK H. SABAJ PEREZ

Introduction

The last decade or so has witnessed a surge in

expeditions to both ichthyologically familiar and virgin

waters in southeastern Venezuela, Guyana, Suriname, and

French Guiana. Included are surveys of the Iwokrama

Forest in west-central Guyana (Watkins et al. 2005),

retracing Carl Eigenmann’s 1908 collecting route up the

Essequibo to the Potaro River above Kaiteur Falls

(Hardman et al. 2002), and rapid assessments targeting

species-rich waters such as the upper Essequibo Basin,

Guyana (Lasso et al. 2008), Coppename Basin, Suriname

(Berrenstein 2005, Alonso & Berrenstein 2006, and

references therein), and Venezuelan states of Amazonas

(Lasso et al. 2006, and references therein), and Bolıvar

(Machado-Allison et al. 2003). Systematic fish inventories

of French Guiana began over 50 years ago (see references

in Vari & Ferraris, this volume), and have been recently

expanded by French and Swiss ichthyologists to include

ecological (e.g., Lord et al. 2007) and molecular data, the

latter to investigate the origins of the Guianas’ highly

diversified fish fauna (Cardoso & Montoya-Burgos 2009).

Explorations of remote Shield regions in search of

undescribed catfishes (Sabaj Perez et al. 2009) have

assembled a parade of new taxa led by the sucker-mouth

armored siluriforms in the family Loricariidae. Fifteen

new loricariid species from Guyana, Suriname, and

Amazonas, Venezuela, have been described in the last

five years (e.g., Werneke et al. 2005, Armbruster et al.

2007, de Chambrier & Montoya-Burgos 2008, Lujan et al.

2009) with many more discoveries awaiting description.

This impressive amount of fieldwork has significantly

advanced our taxonomic understanding of fishes in the

Guianas; nevertheless, much must still be accomplished.

Expeditions to remote, previously unsampled waters,

particularly headwater systems above waterfalls or large

cataracts, routinely yield new and sometimes enigmatic

ichthyofaunas (Taphorn et al. 2008; Lujan, pers. comm.;

pers. obs.). More comprehensive collecting efforts (e.g.,

night sampling) in relatively well-sampled waters have

uncovered new species that escaped prior efforts (e.g.,

Armbruster et al. 2000; pers. obs.). Fieldwork aside,

there exists in museums a wealth of specimens of

Guianas fishes that require critical evaluation. The rich

and complex diversity of fishes in the Guianas, and their

systematic placement in the greater context of the

Neotropical fauna, will remain a lodestone for ichthy-

ological studies in decades to come.

Scope

The plates present 130 individuals representing 127

species of 46 families. Fishes were collected in Guyana

(53 species), Suriname (36) and Amazonas State,

Venezuela (38) from 1985 to 2008. Most of the species

occur on or immediately peripheral to the Guiana

Shield, with a few species restricted to lowland, coastal

habitats in fresh and/or estuarine waters (i.e., Rhino-

sardinia amazonica, Sciades parkeri, Tomeurus gracilis,

Anableps anableps, Polycentrus schomburgkii).

Fishes were imaged live or shortly after death (89

species), or from specimens purchased at market (2),preserved in formalin (2), or stored in alcohol (34).

Each image is identified in the plate description by

taxon, condition of specimen at time of photo, museum

and catalog number, size and sex (if so determined),

current status of voucher if other than preserved whole

in alcohol, and complete locality data. Depositories are

The Academy of Natural Sciences, Philadelphia

(ANSP), Auburn University Natural History Museum(AUM), Field Museum of Natural History (FMNH),

Illinois Natural History Survey (INHS), Museo de

Ciencias Naturales de la UNELLEZ, Guanare

(MCNG), National Zoological Collection of Suriname

(NZCS), and University of Guyana, Center for the

Study of Biological Diversity (UG/CSBD). Photos are

by author unless credited otherwise. Abbreviations in

the text are: LEA – length to end of anal fin; SL –standard length; and TL – total length. Scale bars are

presented only for those species in which that indicator

was included in the original photograph.

Fish Photography

There is a variety of techniques for capturing high-

quality color images of fishes, all of which have been

vastly simplified and in many ways improved by the

advent of digital technology. Most of the images

presented here are of live (or recently so) and alcohol

preserved specimens immersed in water in a glass

phototank. Materials and methods are largely the same

whether taken streamside of live specimens (Figs. 3, 4) orin the lab of preserved specimens (Fig. 5), except for the

light source: ambient sunlight in the field vs. incandescent

light in-doors. Other photographers have used electronic

flashes (e.g., Jenkins & Burkhead 1994:129, Planquette et

al. 1996:17) to produce stunning photos of live fishes in

phototanks. I have not tried such techniques, but consider

a cooperative sun to be equally effective and in some ways

less burdensome. In any event, phototank-immersionremains the gold standard for ex-situ fish photography.

Phototank-immersion Method

This method involves three stages: equipment set up,specimen preparation, and image capture and editing.

The techniques described below follow a minimalist

approach with some advice limited to the specific

cameras and conditions involved. For a more sophis-

ticated system and additional tips on fish photography

see Jenkins & Burkhead (1994:127–130).

Equipment set up.—The phototank is made of

ordinary plate glass bonded together with clear silicone

adhesive. Outside dimensions (in inches) of the tanks

used for the photos in this section are: 13.5 length 3

10.25 height 3 2.75 width (field and lab) and 15.75

length 3 12.25 height 3 3.5 width (lab only). Both are

made from one-quarter inch thick glass, except one-

eighth inch glass is used for the front plate of smaller

tank. These dimensions are well suited for lateral and

often dorsal/ventral views of small to medium-sized

fishes up to about 300 mm total length and 63 mm

width for smaller tank, and 370 mm total length and 75

mm width for larger. Two important factors compro-

mise field utility of larger phototanks: the volume of

water necessary to fill it and size of carrying case (see

below). Each tank requires a separate glass plate to

immobilize the subject. The free plate can be one-

eighth (smaller tank) or one-quarter (larger) inch thick

and is slightly shorter and deeper than the inside

dimensions of the tanks (e.g., 13 3 10.25 and 15 3 12

inches for smaller and larger tank, respectively).

Having smooth edges of all plates is recommended.

The tank should be filled with clear bottled or

filtered/deionized tap water to minimize formation of

air bubbles on specimen and glass. Stream or lake

water is unsuitable because it lacks the desired clarity

and suspended debris is a significant distraction in an

otherwise good photo. Any water will accumulate

debris over an extended photo session, and an ample

supply of clean photo water must accompany long

forays to remote locations.

In the lab the phototank is stationed between two

pairs of incandescent bulbs positioned to the side and

slightly above the top of the tank (Fig. 5). Polarizing

filters are useful for reducing glare or overexposed hot

spots on the specimen, particularly on the snout. When

using sunlight, the tank is oriented to maximize the

even distribution of light and minimize glare and

shadows on the subject.

Selection of a camera is important, but the rapid

pace of digital technology soon outdistances specific

recommendations on make or model. By current

standards a digital camera with a good optical zoom

(6X and higher) that records images at or above

resolutions of 12 Megapixels (MP) is generally a safe

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


choice. Most of the photos herein were taken with a

Nikon Coolpix E8700 (8 MP); others with this model’s

predecessors, the E4500 (4 MP) and older E995 (3.1

MP). The most recent photos, all of alcohol preserved

specimens, were taken in the lab with a Nikon D90 D-

SLR (12.3 MP) fitted with a 60 mm f/2.8G micro lens.

Images taken with the E8700 contain a high level of

sharp detail that is slightly exceeded by the D90 (or

other cameras offering greater MP), particularly for

small specimens. The differences, however, are only

visible at high magnification or extremely large print

sizes. The greatest advantage of the D-SLR design and

micro lens is the enhanced ability to reliably focus on

very small specimens. Any camera and lens should be

thoroughly vetted by comparing published reviews

(many available on-line), and then personally tested

with the phototank-immersion method. A few digital

cameras apparently have difficulties rendering a sharp

specimen image through glass and water.

Additional essentials for basic set up are a tripod

(mini-tripods are handy in the field; Figs. 3–5), 4-ply

mat board in several background colors (e.g., flat

black, dull light blue) and 3/16th inch foam board with

flat black surface for camera blind (sizes of all boards

ideally fitted to carrying case), glass cleaner, and paper

towels or lint-free cloth, both long and small forceps,

large metal binder clips, 12-inch plastic metal rulers,

stiff wire, an assortment of needles and insect pins,

calipers, a system for tagging individual specimens

(e.g., dymo-tags in pre-punched number series tied to

strong twine), extra camera batteries and charger,

memory cards and reader, and laptop computer for

image storage. These essentials are best stored with the

phototank in a crushproof and watertight carrying

case. The smaller tank is ideal for field use as it requires

less water and allows for co-storage of accessories and

laptop in a small case suitable for carry-on luggage (see

Fig. 3). Cameras are better stored separately to

facilitate other uses and avoid residual moisture in

the phototank.

Specimen preparation.—The overarching strategy

when photographing fishes for identification purposes

is to maximize the content and accuracy of information

in the image. This aim determines which among

multiple specimens is photographed, how it is illumi-

nated and arranged for display, and which color

background is used. Most striking are photographs of

the most impressive specimens (i.e., in peak coloration

and with fins and scales intact), but even the image of

an impressive fish may be rendered less informative if

the photograph is poorly composed.

Once a live or alcohol specimen is selected it is

carefully inspected and cleaned of foreign debris.

Mucous-laden skin and fins often attract distracting grit

or other suspended particles, and cheesecloth fibers may

adhere to preserved specimens. An anesthetized fish (e.g.,

with a few drops of clove oil) is quickly euthanized in a

container of strong (30–50%) formalin. This often causes

the body to straighten and fins to become completely

erect. Otherwise the anesthetized specimen may be

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

NUMBER 17 55

removed to a tray of shallow formalin wherein small

forceps are carefully used to hold the fins erect without

damaging them. The most important consideration

when photographing live specimens is time; bright colors

and iridescences are soon lost in formalin. Fatty skin, as

in pseudopimelodid catfishes, also becomes opaque in

formalin, obscuring any underlying color.

Once the specimen is flat with fins erect, it is care-

fully wedged between the front plate of the phototank

and free plate of glass, the latter set at an angle and

braced against metal binder clips either attached to the

sides of the tank or loosely set between the free plate

and back of tank (Fig. 3). Positioning laterally com-

pressed fishes in this manner is easy. Dorsoventrally

depressed specimens, particularly those with pectoral

spines, require more attention to achieve a vertical

lateral view. Maintaining pectoral spines folded against

the body as one wedges the specimen between the two

glass plates requires practice and patience. Long

forceps, a metal ruler and stiff wire are useful tools

for fine-tuning a specimen’s posture, arranging long

delicate features such as barbels, and dislodging air

bubbles that form on the fish. Information content of a

fish photo is diminished when the specimen is tilted or

otherwise poorly positioned.

Preserved specimens offer fewer options for achiev-

ing an ideal photo-friendly posture. Laterally contorted

specimens often can be made to appear more linear

when tightly wedged between the two plates of glass.

Issues that are more difficult arise with partial or

complete folding of fins. In some cases insect pins

(carefully inserted in the body opposite the side to be

imaged) may be used to prop up the anterior most

portions of fins. This technique, however, may cause

small tears in the fin membranes.

Next is selection of an appropriate background.

Many specimens, particularly dark ones with opaque

fins, often render best with more dramatic effect

against flat black backgrounds. This may pose a

serious drawback for specimens with relatively trans-

parent fins. Black pigment in fin membranes or along

distal fin margins disappears against dark back-

grounds. In such cases, a light blue background

provides better contrast and will highlight dark

pigmentation in fins. Conversely, transparent fins

lacking pigmentation and with clear margins, particu-

larly in live specimens, are often lost against light

backgrounds. This can be alleviated to a certain degree

by adjusting the tank relative to light source to achieve

a small measure of direct side or back lighting. While it

is true that graphics editing software (e.g., Vertus Fluid

Mask) can virtually affect any color background,

specimens may not appear natural if the new back-

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

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

by author.


white and vice versa). Choice of background color

often involves trade-offs, and is ultimately a reflection

of personal taste determined via trial and error.

The final step is placement of a scale bar. This is

accomplished by cutting out a 10+ mm portion of a

plastic ruler, dipping it in water and adhering it to the

outside front of the phototank beneath the specimen

and within the photographic field.

Image capture and editing.—The camera is mounted

on a tripod, as most exposures are too long to permit

hand-held use, and positioned behind a black foam

board with central circular aperture fitted to lens. The

blind prevents the phototank glass from reflecting the

images of camera and photographer. Whether hori-

zontal or angled the specimen should occupy about

90% of the length of the digital image recorded. To

preserve detail in extremely long and slender fishes

(e.g., belonids; Pl. 14, Fig. E) the specimen is imaged in

two aligned and overlapping parts (anterior and

posterior halves) that are digitally combined. The

shutter is placed on a timer delay and white balance

set appropriately (e.g., sunlight vs. incandescent).

Digital photography frees one from limits imposed

by the amount of available film and developing costs.

In the field, particularly while the sun is dodging

clouds, it is advisable to take multiple photos for

each of several combinations of exposures and aper-

tures (f-stops). Full sunlight often highlights fine

structures (e.g., odontodes in loricariids), but at the

same time may wash out bright colors or result in

overexposed hot spots on the snout or dorsum. The

phototank should be carefully oriented with respect

to the light source, and extra mat boards used to

shadow harsh sunlight and maintain vibrant colors

(Fig. 3).

For the Coolpix E8700 in manual mode, the shutter

speed is set such that the target aperture (i.e., lower

f-stops) lies between f-stops 5 and 7; larger apertures

reduce depth of field, and smaller apertures tend to

reduce resolution. The Nikon D90 D-SLR better

accommodates smaller apertures (f-stop fixed at 16

with ISO set to 200), and the shutter speed is manually

adjusted for the best exposure. Autofocus generally

works fine as long as the active area of focus includes

important features on the fish, not the scale bar or

background. Digital cameras typically have a setting

whereby the user determines the active area of

autofocus. Depending on specimen size, the camera

may need to be manually set to macro mode, and some

cameras (e.g., Nikon CoolPix) also require one to

slightly zoom in on subject for sharp autofocus. Nikon

images presented here are of Fine quality (recorded as

JPEGs with compression ratio of roughly 1:4) and

maximum size (3264 3 2448 and 4288 3 2848 pixels for

E8700 and D90, respectively). Higher quality settings

record either uncompressed TIFF or RAW (NEF)

images, the latter requiring extra software and com-

puting time for conversion to TIFF files (Nikon D90

allows one to record NEF and Fine JPEG images

concurrently). TIFF and RAW files retain the full

quality of the image and the latter maximizes allowable

post exposure processing, whereas JPEGs are com-

pressed often with some visual quality permanently lost

in the process (the loss, however, is barely perceptible).

Larger image files (NEF, RAW, TIFF) do offer

slightly higher resolution, but the improvement is often

negligible, except at high magnification. For any

camera, there is no substitute for testing a variety of

settings and image qualities to optimize the desired

effect and protocol.

While photographing a specimen it is difficult to

know which image will optimize the desired effect; so,

it is best to have ample images from which to choose.

The number of images I generally take is directly

proportional to the impressive and unique nature of

the specimen added to the amount of time expended to

pose it properly in the phototank. It is easy to

accumulate many photos of numerous species, thus it

is critical to have a system for later identification and

management of images. Failure to do so guarantees

extra time and often frustration when attempting to

match images to specimens long after capture. The best

field solution is to take a final photo of the specimen

together with a uniquely numbered tag that is then

secured to the fish. In the case of museum specimens,

the jar label is photographed immediately after imaging

the fish. A photo-log is useful for recording the

standard length of the specimen. Such practices greatly

facilitate subsequent annotation of images with catalog

and measurement data. A new and much welcomed

trend in digital cameras is a built in or accessory global

positioning system (GPS) receiver that records and

embeds latitude, longitude, altitude and universal time

as image metadata.

The final step is image editing, all of which was

performed on the photos in this section using Adobe

Photoshop. This program offers a seemingly endless

myriad of simple to advanced tools for graphics

manipulation. Only a few of the more basic tools and

techniques are mentioned briefly here.

Once an image is selected the background (original)

layer is immediately duplicated and subsequent edits

are made to the duplicate layer. A third blank layer is

added to mask the specimen with a uniform back-

ground. Masking color (e.g., solid black or white, or a

color shade taken from the original background using

the eyedropper tool) is first added as a rough outline

using a large diameter pencil tool, and then completed

with a fine-tipped brush (1–10 pixels) under magnifi-

cation (e.g., $300%) to carefully trace the specimen’s

precise contours. The magic wand and/or magnetic

lasso are more expedient, yet less precise, tools for

masking the specimen with a uniform background.

Next, the duplicate layer is automatically and manually

NUMBER 17 57

adjusted for levels (tonal range and color balance),

brightness/contrast, and hue/saturation. The auto

options often render extreme values that are manually

faded to desired opacity before additional manual fine-

tuning. The cloning stamp tool is useful for removing

small bubbles or debris on the specimen, while the

dodge and burn tools help lighten or darken localized

regions (brush size/shape and exposure/opacity of such

tools are manually adjusted). Under the Filter menu

‘Sharpen.Unsharp Mask’ can sharpen an image with

soft focus, and ‘Noise.Despeckle’ removes graininess,

particularly for images scanned from 35 mm slides.

One final trick is to render a solid scale bar by: 1)

rotating the entire image so the ruler piece in the

photograph is horizontal, 2) using the rectangular

marquee tool to select and copy a 5 or 10 mm long

portion, 3) rotating the entire image back to its final

intended position, 4) pasting the copied selection,

thereby creating a new layer, and 5) adjusting the

brightness/contrast of this layer to extreme values to

render a black or white bar that is then labeled

accordingly with the text tool.

The final edited version of the specimen image (i.e.,

duplicate layer) can be quickly compared to the

original by using the layers window and clicking the

‘eye’ icon to hide or display the corresponding layer.

Creation of additional layers requires the new image to

be saved as an uncompressed TIFF file that is suitable

for archiving and any additional post processing. A

copy of the final edited image is flattened to a single

layer and resaved as a TIFF for print publication (with

Image.Mode set to Grayscale, RGB, or CMYK color

based upon printer specifications) and separately as a

JPEG (with Mode set to 8 bits/channel) for use in

presentations or easy transmission and accession via

the Internet.

In sum, a high-quality fish photo is the product

of preparation, practice, and patience all committed

with keen attention to detail. Additional factors in

the field are perseverance under suboptimal conditions

and a bit of luck with respect to weather and finding

the ideal specimen. The amount and quality of the

images presented herein had more to do with will than

skill.

Acknowledgments

Numerous colleagues contributed valuable assis-

tance both before and after the shutter click. For

assistance in the field I wish to thank in particular:

Mariangeles Arce, Jonathan Armbruster, Michael

Hardman, Oscar J. Leon-Mata, Nathan Lujan, John

Lundberg, Jan Mol, Matthew Thomas, David Wer-

neke, and Philip Willink. For loans of specimens I

thank Michael Retzer, and for help tracking down

specimens and taking measurements I thank Osvaldo

Oyakawa and especially Nathan Lujan. For help

cleaning backgrounds I thank Kyle Luckenbill and

Rebecca Meyer. Special thanks to Pierre-Yves le Bail,

Raphael Covain and Juan Montoya Burgos for

comments on the introduction, and to Jonathan

Armbruster, Hernan Lopez-Fernandez, Nathan Lujan,

Donald Stewart, and Donald Taphorn for graciously

contributing their images to this atlas. Finally I thank

my ichthyological friends for years of help identifying

many of the fishes presented here. Support for this

project was received in part from the All Catfish

Species Inventory (NSF DEB-0315963).

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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.

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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: Hyposto-

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

Central Suriname Nature Reserve (CSNR), (Coppename

River Basin, Suriname). Conservation International Suri-

name, 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 morpholog-

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Zootaxa 1918:45–58.

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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 Fresh-

waters 13:225–238.

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

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acuaticos en la confluencia de los rıos Orinoco y Ventuari,

Estado Amazonas (Venezuela). Boletın RAP de Evaluacion

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

240 pp.

Lasso, C. A., J. Hernandez-Acevedo, E. Alexander, J. C. Senaris, L.

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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

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

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

francaise). Resultats preliminaires.—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.

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

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Montambault, eds., A biological assessment of the aquatic

Ecosystems of the Caura River Basin, Bolıvar State,

Venezuela. RAP Bulletin of Biological Assessment 28.

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

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Institut national de la Recherche Agronomique, Conseil

Superior de la Peche, Paris, 429 pp.

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

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

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

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uniquely colored species of catfishes from Venezuela (Silur-

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

NUMBER 17 59

APPENDIX: PLATES

Plate 1

Potamotrygonidae

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

Amazonas, Rıo Negro (Amazonas drainage), left bank sandy beach and small adjacent

backwater 7.2 km NW of San Carlos de Rio Negro, 01u589110N, 067u069100W, 19 Mar

2005, M. Sabaj, D. Werneke et al.

B. Potamotrygon orbignyi (live). AUM 43201 (171 mm maximum disk width). Venezuela,Amazonas, Rıo Orinoco ca. 60 km E of San Fernando de Atabapo, 03u589260N,

067u099460W, 3 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.

C. Potamotrygon marinae (live). ANSP 187098 (400 mm maximum disk width). Suriname,

Sipaliwini, Lawa River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan

(airstrip), 03u199310N, 054u039480W, 18 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P.

Willink, & K. Wan.

D. Potamotrygon schroederi (live). AUM 44507 (423 mm maximum disk width). Venezuela,

Amazonas, Rıo Orinoco, island W of Puerto Venado, 4.5 km S of Samariapo, 56.5 km

SW of Puerto Ayacucho, 05u129250N, 067u489320W, 28 Feb 2005, M. Sabaj, N. Lujan, D.

Werneke et al.


NUMBER 17 63

Plate 2

Arapaimidae

A. Arapaima sp. (live). UG/CSBD uncataloged (174 cm SL, male, skeleton). Guyana, Grass

Pond, Rewa River basin (Rupununi drainage), near Rewa village, Aug 2006, D. Stewart et

al. Photo by D. J. Stewart.

Osteoglossidae

B. Osteoglossum bicirrhosum (live). No voucher (ca. 500 mm TL). Guyana, Crane Pond,

southwestern part of Rupununi basin near Karanambu Ranch, 2007, D. Stewart et al.

Photo by D. J. Stewart.


NUMBER 17 65

Plate 3

Clupeidae

A. Rhinosardinia amazonica (alcohol). INHS 49009 (33.7 mm SL). Guyana, East Demerara,

Demerara River (Atlantic drainage), Land of Canaan, 11.8 mi S-SW of Georgetown at

bearing 213u, 06u389390N, 058u119460W, 14 Oct 1998, M. Sabaj, J. Armbruster, M.

Hardman et al.

Pristigasteridae

B. Pellona castelnaeana (live). MCNG 51957. Venezuela, Amazonas, Rıo Casiquiare at

mouth of Cano Caripo, 37 km W-SW of La Esmeralda, 03u069500N, 065u529380W, 5 Mar

2005, M. Sabaj, N. Lujan, D. Werneke et al.

Engraulidae

C. Amazonsprattus scintilla (live). ANSP 181134. Guyana, Pirara River (Ireng-Takutu-

Branco drainage), 3.5 km N-NW of Pirara, 03u389550N, 059u419200W, 2 Nov 2002, M.

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

D. Anchovia surinamensis (alcohol). ANSP 189252 (69.7 mm SL). Guyana, Rupununi River

(Essequibo drainage), at Massara’s Landing, 1.1 km NE village of Massara, 03u539410N,

059u179370W, 26 Oct 2002. M. Sabaj, J. Armbruster, M. Thomas et al.

E. Lycengraulis batesii (alcohol). ANSP 189251 (52.4 mm SL). Guyana, Cuyuni-Mazaruni,

Mazaruni River (Essequibo drainage) long partially exposed sandy shoal between two

islands in main channel, 6.9 km SW of Bartica, 06u229470N, 058u409320W, 12 Nov 2002,

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

F. Anchoviella sp. (alcohol). ANSP 189234 (29.5 mm SL). Suriname, Sipalawini, Lawa River

(Marowini drainage), small sand beach below cataract upstream from base camp, ca. 9 kmS-SW of Anapaike. 03u199120N, 054u039410W, 19–23 Apr 2007, J. Lundberg, J. Mol, M.

Sabaj, P. Willink, & K. Wan.

Parodontidae

G. Apareiodon agmatos (alcohol). ROM 83755 (52.6 mm SL). Guyana, Mazaruni River

(Essequibo drainage), sandy beach and embayment on right bank, upstream from villageof Jawalla, 05u41935.40N, 060u28911.80 W, 18 Apr 2008, H. Lopez-Fernandez, D.

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

Lopez-Fernandez.

H. Apareiodon orinocensis (live). ANSP 185045. Venezuela, Amazonas, Rıo Orinoco at

Puerto Venado, 4.3 km S of Samariapo, 56.4 km SW of Puerto Ayacucho, 05u129380N,

067u489180W, 26 Feb 2005, M. Sabaj, N. Lujan, D. Werneke et al.

I. Parodon guyanensis (live). ANSP 189204 (ca. 70 mm SL). Suriname, Sipaliwini, Lawa River

(Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip), 03u199310N,

054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.

J. Parodon guyanensis (alcohol). ANSP 189204 (73 mm SL). Same locality as I.


NUMBER 17 67

Plate 4

Curimatidae

A. Curimatopsis crypticus (live). ANSP 189091 (32.4 mm SL, female). Suriname, Para,

Coropinae Creek (Suriname drainage), vicinity of Republiek, 05u299570N, 055u129520W,

28 Apr 2007, M. Sabaj & P. Willink.

B. Curimatopsis crypticus (live). ANSP 189091 (28.8 mm SL, male). Same locality as A.

Prochilodontidae

C. Prochilodus rubrotaeniatus (alcohol). ANSP 175495 (160 mm SL). Guyana, Essequibo

River, sandbar ca. 800 m downstream of Essequibo (Maipuri) campsite, 04u459430N,

058u459520W, 29 Jan 1997, W. Saul et al.

D. Semaprochilodus varii (live). ANSP 187435 (210 mm SL). Suriname, Sipaliwini, LawaRiver (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),

03u199310N, 054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &

K. Wan.

Crenuchidae

E. Leptocharacidium omospilus (live). ANSP 189272 (60.5 mm SL). Venezuela, Amazonas, 1km upstream from mouth of left bank tributary of Rıo Siapa, mouth below Salto Oso and

above Salto Sardinas on Rıo Siapa, 01u269240N, 065u409010W, 14 Mar 2005, M. Sabaj, N.

Lujan, M. Arce, & T. Wesley.

F. Poecilocharax bovalii (alcohol). INHS 49600 (28.6 mm SL). Guyana, first N bank creek

tributary of Potaro River (Essequibo drainage) downstream from Waratuk Cataract, 27

Oct 1998, L. Page, M. Sabaj, J. Armbruster et al.

Hemiodontidae

G. Argonectes longiceps (alcohol). ANSP 189151 (184 mm SL). Same locality data as D.

H. Bivibranchia bimaculata (live). ANSP 189149 (111 mm SL). Same locality data as D.

Gasteropelecidae

I. Carnegiella strigata (alcohol). INHS 49173 (26 mm SL). Guyana, Mazaruni–Potaro,

‘‘Himarakus’’ Creek, tributary of Essequibo River (Atlantic drainage) at Rockstone,

05u599080N, 058u339030W, 19 Oct 1998, M. Sabaj, J. Armbruster, M. Hardman.

J. Gasteropelecus sternicla (alcohol). ANSP 189193 (46.6 mm SL). Same locality as A.


NUMBER 17 69

Plate 5

Anostomidae

A. Anostomus brevior (live). FMNH ex ANSP 189141 (51.6 mm SL). Suriname, Sipaliwini,

Lawa River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),


K. Wan.

B. Anostomus anostomus (live). ANSP 180172 (72.5 mm SL). Guyana, Essequibo River(Atlantic drainage) at Yukanopito Falls, 44.5 km SW of mouth of Kuyuwini River,

01u549530N, 058u319140W, 9 Nov 2003, M. H. Sabaj, J. Armbruster, N. Lujan et al.

C. Synaptolaemus cingulatus (live). AUM 43269 (78.8 mm SL). Venezuela, Amazonas, Rıo

Orinoco, 147 km SE of San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar

2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.

D. Leporellus vittatus (formalin). ANSP 189270 (photo voucher). Guyana, Rupununi River(Essequibo drainage), road crossing 5.9 km W-SW of village of Sand Creek, 02u579000N,

059u349100W, 4 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.

E. Leporinus fasciatus (live). ANSP 189158 (81.7 mm SL). Same locality as A.

F. Leporinus maculatus (live). FMNH ex ANSP 189041 (123.4 mm SL). Same locality as A.

G. Hypomasticus megalepis (live). AUM 37999 (68.3 mm SL). Guyana, Essequibo River atKassi-Attae Rapids, 5.5 km SE of mouth of Kuyuwini River, 02u139360N, 058u179380W, 8

Nov 2003, M. Sabaj, J. Armbruster, M. Hardman et al.

H. Leporinus ortomaculatus (live). AUM 43262 (72.9 mm SL). Same locality as C.

I. Leporinus lebaili (live). FMNH ex ANSP 189043 (56.7 mm SL). Same locality as A.

J. Caenotropus maculosus (live). ANSP 189147 (70 mm SL). Suriname, Sipalawini, LitanieRiver, side channel behind Theo’s bakery, just upstream of confluence with Marowini

River and village of Konya Kondre, 03u179240N, 054u049380W, 23 Apr 2007, J. Lundberg,

J. Mol, M. Sabaj, P. Willink, & K. Wan.


NUMBER 17 71

Plate 6

Characidae

A. Brittanichthys myersi (formalin). INHS 49053 (22.4 mm SL). Guyana, East Demerara,

Maduni River and Conservancy Canal (Mahaica-Atlantic drainage), Maduni Stop-off,

22.3 mi S of Georgetown bearing 176u, 06u309010N, 58u029140W, 15 Oct 1998, M. Sabaj, J.

Armbruster, M. Hardman et al.

B. Bryconops melanurus (live). ANSP 189268 (75.5 mm SL). Suriname, Sipalawini, LitanieRiver at mouth and confluence with Marowini River, just upstream from settlement of

Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P.

Willink, J. Mol et al.

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

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

Monduapo, 1 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.

D. Exodon paradoxus (live). AUM 36843 (61 mm SL). Guyana, Rupununi River (Essequibo

drainage), sand beach and inlet at Karanambo Ranch, 03u459000N, 059u189300W, 30 Oct

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

E. Myleus rubripinnis (live). ANSP 189267 (34 mm SL). Guyana, Suriname, Sipaliwini, Lawa

River (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),


K. Wan.

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

Orinoco, bedrock outcrop, 52.9 km SE of San Antonio, 102 km W of La Esmerelda,

03u069010N, 066u279460W, 4 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.

G. Myloplus cf. planquettei (live). ANSP 179808 (425 mm SL). Guyana, Essequibo River

(Atlantic drainage) at Yukanopito Falls, 44.5 km SW of mouth of Kuyuwini River,

01u549530N, 058u319140W, 9 Nov 2003, M. H. Sabaj, J. Armbruster, N. Lujan et al.

H. Serrasalmus rhombeus (live). ANSP 180287 (ca. 260 mm SL). Venezuela, Amazonas, Rıo

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

of San Fernando de Atabapo, 7 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.


NUMBER 17 73

Plate 7

Acestrorhynchidae

A. Acestrorhynchus falcatus (live). INHS 48983 (153.9 mm SL). Guyana, East Demerara,

Madewini River (Demerara drainage), 21.5 mi S-SW of Georgetown, bearing 207u, at

Linden highway bridge, 06u 309 05.00 N, 58u 129 44.90 W, 14 Oct 1998, M. Sabaj, J.

Armbruster, M. Hardman et al.

B. Acestrorhynchus microlepis (live). AUM 36753 (123 mm SL). Guyana, Circle West Creek,tributary of Pirara River (Ireng-Takutu drainage), 26.6 km SW of Karanambo Ranch,

03u399140N, 059u319430W, 30 Oct 2002, M. Sabaj, J. Armbruster, M. Thomas et al.

Cynodontidae

C. Cynodon meionactis (alcohol). ANSP 189129 (190 mm SL). Suriname, Sipalawini, Litanie

River at mouth and confluence with Marowini River, just upstream from settlement ofKonya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P.

Willink, J. Mol et al.

D. Hydrolycus armatus (live). MCNG 51983 (ca. 400 mm SL). Venezuela, Amazonas, Rio

Orinoco (Atlantic drainage), Pasaganado, 38 km N of San Fernando de Atabapo,


Lebiasinidae

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

tributary of Rıo Orinoco, 1.5 km S-SE of Manaka, 70 km E of San Fernando de Atabapo,

4 Apr 2004, N. Lujan & D. Werneke.

F. Copella cf. arnoldi (live). ANSP 189192 (30.4 mm SL). Suriname, Para, Coropinae Creek

(Suriname drainage), vicinity of Republiek, 05u299570N, 055u129520W, 28 Apr 2007, M.Sabaj & P. Willink.

Erythrinidae

G. Hoplias aimara (alcohol). ANSP 176723 (172 mm SL). Guyana, Tumble Down Creek,

tributary of Siparuni River (Essequibo drainage), 04u489390N, 058u519110W, 8 Dec 1997,

G. Watkins et al.

H. Hoplias lacerdae group (live). AUM 44674 (152 mm SL). Guyana, Pirara River (Ireng-

Takutu drainage), at Pirara Ranch, 03u379310, 059u409360, 26 Nov 2005, N. Lujan, D.

Taphorn et al. Photo by N. K. Lujan.

I. Erythrinus erythrinus (alcohol). ANSP 175537 (121 mm SL). Guyana, Culvert creek

(Essequibo drainage) crossing Kurupukari-Surama River road, 04u199570N, 058u519130W,

5 Feb 1997, W. Saul et al.

Ctenoluciidae

J. Boulengerella cuvieri (live). AUM 40987 (560 mm SL). Venezuela, Amazonas, Rıo Ventuari

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

de Atabapo, 04u259100N, 066u179080W, 7 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.


NUMBER 17 75

Plate 8

Cetopsidae

A. Cetopsis cf. montana (live). ANSP 178782 (78.3 mm SL). Guyana, Ireng River (Takutu-

Branco-Negro drainage), 6.9 km W-SW of village of Karasabai, 04u019100N,


B. Helogenes marmoratus (live). AUM 27960 (56 mm SL). Guyana, West Demerara,

Catabuly Creek (Demerara-Atlantic drainage), at Wismar, 1.87 mi. N-NW of Lindenbearing 335u, 06u01937.20N, 58u19925.30W, 18 Oct 1998, L. Page, M. Sabaj, J. Armbruster

et al. Photo by J. W. Armbruster.

Aspredinidae

C. Ernstichthys sp. (alcohol). ANSP 180002 (26.4 mm SL). Same locality as A.

D. Bunocephalus verrucosus (alcohol). ANSP 180015 (36.6 mm SL). Guyana, Hassar Pond

and outlet (Rupununi drainage), 5.4 km S-SE of Massara, 03u509400N, 059u179090W, 27

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

Ariidae

E. Sciades parkeri (live). ANSP 178741 (445 mm SL). Guyana, purchased at Georgetown fishmarket, 8–15 Nov 2002, M. Sabaj.

Auchenipteridae

F. Ageneiosus inermis (live). FMNH ex ANSP 187115 (ca. 200 mm SL). Suriname,

Sipalawini, Litanie River at mouth and confluence with Marowini River, just upstream

from settlement of Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg,M. Sabaj, P. Willink, J. Mol et al.

G. Auchenipterichthys punctatus (live). AUM 43416 (131 mm SL). Venezuela, Amazonas, Rıo

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


H. Trachycorystes trachycorystes (live). AUM 35933 (111 mm SL). Guyana, unnamed stream

(Rupununi drainage) at crossing on road between Massara and Karanambo, 10.3 km NWof Karanambo Ranch, 03u489270N, 059u239060W, 28 Oct 2002, M. Sabaj, J. Armbruster,

M. Thomas et al.

I. Gelanoglanis sp. (live). AUM 35908 or ANSP 178738 (male). Guyana, Ireng River

(Takutu-Branco drainage), 6.9 km WSW village of Karasabai, 04u019100N, 059u369060W, 1

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

J. Glanidium leopardum (live). ANSP 189104 (48.5 mm SL). Same locality as F.


NUMBER 17 77

Plate 9

Doradidae

A. Acanthodoras cataphractus (alcohol). NZCS 1618–1619 (84.5 mm SL). Suriname, Para,

tributary of lower Suriname River at Republiek, 14 Nov 1989, P. Outboter et al.

B. Scorpiodoras heckelii (live). ANSP 182790 (82.5 mm SL, specimen with simple secondary

gas bladder). Venezuela, Amazonas, Rıo Orinoco at Puerto Samariapo, 05u159N,

067u489W, 25 Feb 2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.

C. Leptodoras copei (live). ANSP 182225 (85.8 mm SL). Venezuela, Amazonas, Rıo Ventuari

(Orinoco drainage), beach across river from Picua village, 34 km E-NE of Macuruco, 104

km E of San Fernando de Atabapo, 04u069550N, 066u459520W, 5 Apr 2004, M. Sabaj, N.

Lujan, D. Werneke et al.

D. Leptodoras linnelli (live). AUM 41038 (171 mm SL). Venezuela, Amazonas, Rıo Ventuari

(Orinoco drainage), village of Marueta at landing, 91 kmE-NE of Macuruco, 159 km E-NE of San Fernando de Atabapo, 04u189510N,

066u179320W, 6 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.

E. Centrodoras hasemani (live). ANSP 182227 (211 mm SL). Venezuela, Amazonas, Rıo

Casiquiare, bedrock outcrop 59.5 km SW of La Esmerelda, 02u499070N, 065u579190W, 8

Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.

F. Oxydoras niger (live). AUM 35508 (340 mm SL). Pirara River, tributary of Ireng River(Takutu-Branco drainage), beach at Pirara Ranch on road to Lethem, 03u379170N,


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, Rıo Ventuari

(Orinoco drainage), beach at village of Moriche, 116 km NE of Macuruco, 169 km NE of

San Fernando de Atabapo, 04u459N, 066u219130W, 7 Apr 2004, M. Sabaj, N. Lujan, D.

Werneke et al.

I. Opsodoras morei (live). ANSP 182836 (157 mm SL). Venezuela, Amazonas, Rıo Orinoco

near confluence with Rıo Atabapo, long narrow beach between channel and laguna, San

Fernando de Atabapo, 04u029480N, 067u429170W, 2 Apr 2004, M. Sabaj, N. Lujan, D.Werneke et al.

J. Rhinodoras armbrusteri (live). ANSP 179096 (66.1 mm SL). Guyana, Takutu River (Rio

Branco-Negro drainage), ca. 2.75 km W of Saint Ignatius, 03u219180N, 059u499510W, 5-6

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


NUMBER 17 79

Plate 10

Pseudopimelodidae

A. Pseudopimelodus bufonius (live). ANSP 189098 (87.4 mm SL). Suriname, Sipalawini,

Litanie River at mouth and confluence with Marowini River, just upstream from

settlement of Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M.

Sabaj, P. Willink, J. Mol et al.

Heptapteridae

B. Mastiglanis sp. (live). FMNH ex ANSP 189106 (43.2 mm SL). Same locality as A.

C. Leptorhamdia sp. (live). AUM 43261 (51 mm SL). Venezuela, Amazonas, Rıo Orinoco,

147 km SE of San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar 2005, M.

Sabaj, N. Lujan, M. Arce, & T. Wesley.

D. Pimelodella geryi (live). ANSP 189109 (114 mm SL). Suriname, Sipaliwini, Lawa River


054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.

Pimelodidae

E. Hypophthalmus marginatus (market). ANSP 187103 (330 mm SL). Suriname, Paramaribo,purchased at main fish market in Paramaribo, presumably from vicinity in Surinam River,

17 Apr 2007, J. Lundberg, M. Sabaj, & P. Willink.

F. Megalonema platycephalum (live). AUM 36018 (97.2 mm SL). Guyana, Rupununi River

(Essequibo drainage), sand beach and inlet at Karanambo Ranch, 03u459000N,

059u189300W, 30 Oct 2002, M. Sabaj, J. Armbruster, M. Thomas et al.

G. Brachyplatystoma filamentosum (live). ANSP 187070 (540 mm SL). Venezuela, Amazonas,Rıo Casiquiare, bedrock outcrop 59.5 km SW of La Esmerelda, 02u499070N,


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

Plate 11

Callichthyidae

A. Callichthys callichthys (live). ANSP 179110 (68.2 mm SL). Guyana, Orokang River

(Mazaruni drainage), 1.2 km S of Chi Chi Falls airstrip, 21.1 km S-SW of Imbaimadai,

05u319310N, 060u139560W, 13 Nov 2002, J. Armbruster, M. Sabaj, M. Thomas et al.

B. Megalechis thoracata (alcohol). ANSP 179795 (62 mm SL). Guyana, Hassar Pond and

outlet (Rupununi drainage), 5.4 km S-SE of Massara, 03u509400N, 059u179090W, 27 Oct2002, J. Armbruster, D. Werneke, C. Allison et al.

Trichomycteridae

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

inlet between two islands in channel, 84.5 km N of San Fernando de Atabapo, 9 km S of

Monduapo, 04u479540N, 067u499160W, 1 Mar 2005, M. Sabaj, N. Lujan, D. Werneke etal.

D. Trichomycterus hasemani (alcohol, top 5 lateral view, bottom 5 dorsal view). ANSP

179154 (13.5 mm SL). Guyana, Rupununi River (Essequibo drainage), 3.7 km S-SE of

village of Massara, 03u519440N, 059u179040W, 27 Oct 2002, M. Sabaj, J. Armbruster, M.

Thomas et al.

E. Henonemus taxistigmus (alcohol). ANSP 179953 (90.6 mm SL). Guyana, Rupununi River(Essequibo drainage) at Kwatamang, 4 km SE of Annai, 03u559030N, 059u069010W, 25 Oct

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

F. Trichomycterus guianensis (alcohol). ANSP 179109 (52.5 mm SL). Guyana, Orokang

River (Mazaruni drainage), at Chi Chi Falls airstrip, 20.1 km S-SW of Imbaimadai,

05u329060N, 060u139590W, 13 Nov 2002, J. Armbruster, M. Sabaj, M. Thomas et al.

G. Typhlobelus sp. (alcohol, dorsal view). AUM 35802 (22.3 mm SL). Guyana, Ireng River(Takutu-Branco-Negro drainage), 6.9 km W-SW of village of Karasabai, 04u019100N,


H. Typhlobelus sp. (alcohol, left 5 dorsal view, right 5 ventral view of head). Same data as

G.

I. Pygidianops sp. (live). ANSP 179820 (23.1 mm SL). Guyana, Takutu River (Branco-Negro

drainage), 3.77 km S-SW of Lethem, 03u219180N, 059u499510W, 16 Nov 2003, M. Sabaj, J.Armbruster, M. Hardman et al.

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


NUMBER 17 83

Plate 12

Loricariidae

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

Ventuari (Orinoco drainage), 23 km NE of Macaruco, 94 km E of San Fernando de

Atabapo, 04u049500N, 066u519540W, 5 Apr 2004, N. Lujan, D. Werneke, M. Sabaj et al.

B. Leporacanthicus cf. galaxias (live). AUM 39226 (93 mm SL). Venezuela, Amazonas, Rıo

Orinoco at Cucue Amerindian village, opposite Trapichote village, 60 km E of SanFernando de Atabapo, 03u589260N, 067u099300W, 3 Apr 2004, N. Lujan, D. Werneke, M.

Sabaj et al.

C. Hemiancistrus medians (live). ANSP 187122 (156 mm SL). Suriname, Sipaliwini, Lawa



K. Wan.

D. Peckoltia sabaji (live). ANSP 185094 (109.7 mm SL). Guyana, Takutu River (Branco-

Negro drainage), 3.77 km S-SW of Lethem, 03u219180N, 059u499510W, 1 Nov 2003, M.

Sabaj, J. Armbruster, M. Hardman et al.

E. Peckoltia braueri (live). AUM 38882 (103 mm SL). Same locality as D.

F. Lithoxus stocki (live). ANSP 189131 (66.5 mm SL). Suriname, Sipalawini, Litanie River at

mouth and confluence with Marowini River, just upstream from settlement of KonyaKondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P. Willink, J.

Mol et al.

G. Pseudolithoxus dumus (live). AUM 43267 (87 mm SL). Venezuela, Amazonas, Rıo

Orinoco, 147 km SE of San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar

2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.

H. Pseudancistrus pectegenitor (alcohol). MCNG 54797 (241.6 mm SL). Venezuela,Amazonas, Rıo Casiquiare, bedrock in stream, 73 km NE of San Carlos de Rıo Negro,

02u219090N, 066u349310W, 9 Mar 2005, N. Lujan, M. Sabaj, D. Werneke et al.

I. Metaloricaria paucidens (live). ANSP 187325 (146.5 mm SL). Suriname, Sipaliwini, Lawa



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,



NUMBER 17 85

Plate 13

Gymnotidae

A. Electrophorus electricus (live). MCNG 51982. Venezuela, Amazonas, Rıo Orinoco, 147 km

SE of San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar 2005, M. Sabaj, N.

Lujan, M. Arce, & T. Wesley.

Sternopygidae

B. Sternopygus sp. (live). ANSP 189018 (146 mm TL). Suriname, Sipaliwini, Lawa River


054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.

Rhamphichthyidae

C. Gymnorhamphichthys hypostomus (alcohol). INHS 49454 (161 mm LEA). Guyana, Potaro

River (Essequibo drainage), beach on N bank, downstream of Tumatumari Cataract,

05u21948.40N, 59u00904.40W, 22 Oct 1998, M. Sabaj, J. Armbruster, & M. Hardman.

Hypopomidae

D. Hypopomus artedi (alcohol). ANSP 189266 (152 mm TL). Suriname, Sipaliwini, LawaRiver (Marowini drainage), ca. 8 km S-SW of Anapaike/Kawemhakan (airstrip),


K. Wan.

Apteronotidae

E. Apteronotus albifrons (live). AUM 40678 (109 mm SL). Venezuela, Amazonas, RıoManapiare (Ventuari-Orinoco drainage), at San Juan de Manapiare landing, 15 Apr 2004,

J. Para.


NUMBER 17 87

Plate 14

Rivulidae

A. Rivulus waimacui (alcohol). INHS 49635 (55.5 mm SL). Guyana, tributary of Potaro River

(Essequibo drainage) near Tukeit Cataract, 05u12916.50N, 059u279090W, 27-28 Oct 1998,

L. Page, J. Armbruster, M. Hardman et al.

Poeciliidae

B. Tomeurus gracilis (alcohol). INHS 49017 (27 mm SL). Guyana, East Demerara, Demerara

River (Atlantic drainage), Land of Canaan, 11.8 mi S-SW of Georgetown bearing 213u,06u389390N, 058u119460W, 14 Oct 1998, M. Sabaj, J. Armbruster, M. Hardman et al.

Anablepidae

C. Anableps anableps (alcohol). INHS 49016 (140 mm SL). Same locality as B.

Belonidae

D. Potamorrhaphis guianensis (live). ANSP 179480 (228 mm SL). Guyana, Yuora River

(Ireng-Takutu-Branco drainage), 6.7 km NE of village of Karasabai on road to Tiger

Creek village, 04u039140N, 059u299070W, 31 Oct 2002, M. Sabaj, J. Armbruster, M.Thomas et al.

E. Potamorrhaphis petersi (alcohol). ANSP 163026 (231 mm SL). Venezuela, Amazonas, Rıo

Sipapo (Orinoco drainage), backwater channel behind sandbar 6-7 km above Pendare,

04u519N, 67u439W, 12 Nov 1985, B. Chernoff et al.

F. Pseudotylosurus microps (alcohol). ANSP 179633 (178 mm SL). Guyana, Rupununi River

(Essequibo drainage) at Kwatamang, 4 km SE of Annai, 03u559030N, 059u069010W, 25 Oct2002, M. Sabaj, J. Armbruster, M. Thomas et al.


NUMBER 17 89

Plate 15

Cichlidae

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 Riverat mouth and confluence with Marowini River, just upstream from settlement of Konya

Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P. Willink, J.

Mol et al.

C. Geophagus sp. (live). AUM 38940 (103 mm SL). Guyana, Kuyuwini River, main channel

and backwater 19.5 km W of confluence with Essequibo River (Atlantic drainage).

02u149280N, 058u309030W, 11 Nov 2003, M. Sabaj, J. Armbruster, M. Hardman et al.

D. Hoplarchus psittacus (live). AUM 41425 (147 mm SL). Venezuela, Rıo Ventuari (Orinoco

drainage), bedrock outcrops 83 km E-NE of Macuruco, 153 km E-NE of San Fernando

de Atabapo, 04u159120N, 066u209410W, 6 Apr 2004, M. Sabaj, N. Lujan, D. Werneke

et al.

E. Mesonauta insignis (live). AUM 40590 (82 mm SL). Venezuela, Rıo Orinoco near mouth

of Rıo Ventuari, Macuruco Landing, 75 km E of San Fernando de Atabapo, 03u579290N,

067u019560W, 4 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.

F. Cichla intermedia (live). ANSP 189269 . Venezuela, Amazonas, Rıo Orinoco, bedrock

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


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


NUMBER 17 91

Plate 16

Synbranchidae

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,

Rıo Iguapo approximately 1 hour above its confluence with Rıo Orinoco, 03u099N,

065u289W, 13 Mar 1987, H. Lopez et al.

D. Plagioscion squamosissimus (alcohol). ANSP 177421 (257 mm SL). Guyana, Siparuni

River (Essequibo drainage), Black Water camp, 04u449N, 058u599W, 6 Dec 1997, G.

Watkins et al.

Eleotridae

E. Microphilypnus sp. (live). ANSP 180643. Venezuela, Amazonas, Rıo Negro (Amazonasdrainage), left bank sandy beach and small adjacent backwater 7.2 km NW of San Carlos

de Rio Negro, 01u589110N, 067u069100W, 19 Mar 2005, M. Sabaj, D. Werneke et al.

Achiridae

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

cano of Rıo Orinoco separating island and beach just downstream of Quiratare, 02u599N,066u049W, 11 Mar 1987, B. Chernoff et al.

G. Soleonasus finis (alcohol). ANSP 179510 (75.5 mm TL). Guyana, Essequibo River

(Atlantic drainage), E bank at Kurukupari, 04u399410N, 058u409310W, 24 Oct 2002, M.H.

Sabaj et al.

Polycentridae

H. Polycentrus schomburgkii (live). FMNH ex ANSP 189014 (22.3 mm SL). Suriname, Para,

Coropinae Creek (Suriname drainage), vicinity of Republiek, 05u299570N, 055u129520W,

28 Apr 2007, M. Sabaj & P. Willink.

Tetraodontidae

I. Colomesus asellus (live). AUM 37945 (35 mm SL). Guyana, Essequibo River (E bank) at

Kurukupari, 04u399410N, 058u409310W, 17 Nov 2003, M. Sabaj, M. Hardman, N. Lujan

et al.


NUMBER 17 93

INDEX TO ORDERS, FAMILIES, AND SUBFAMILIES

Acestrorhynchidae

Achiridae

Agoniatinae

Anguilliformes

Anostomidae

Aphyocharacinae

Apteronotidae

Arapaimidae

Aspredinidae

Auchenipteridae

Belonidae

Beloniformes

Bryconinae

Callichthyidae

Cetopsidae

Characidae

Characiformes

Characinae

Cheirodontinae

Chilodontidae

Cichlidae

Clupeidae

Clupeiformes

Crenuchidae

Ctenoluciidae

Curimatidae

Cynodontidae

Cyprinodontiformes

Doradidae

Engraulidae

Erythrinidae

Gasteropelecidae

Genera Incerta Sedis, family

Characidae

Glandulocaudinae

Gobiidae

Gymnotidae

Gymnotiformes

Hemiodontidae

Hemiramphidae

Heptapteridae

Hypopomidae

Hypoptopomatinae

Hypostominae

Iguanodectinae

Lebiasinidae

Lepidosirenidae

Lepidosireniformes

Lithogeninae

Loricariidae

Loricariinae

Myliobatiformes

Ophichthidae

Osteoglossidae

Osteoglossiformes

Parodontidae

PerciformesPimelodidae

Pleuronectiformes

Poeciliidae

Polycentridae

Potamotrygonidae

Pristidae

Pristiformes

PristigasteridaeProchilodontidae

Pseudopimelodidae

Rhamphichthyidae

Rivulidae

Sciaenidae

SerrasalminaeSiluriformes

Sternopygidae

Stethaprioninae

Synbranchidae

Synbranchiformes

Tetragonopterinae

TetraodontidaeTetraodontiformes

Trichomycteridae

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