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BULLETIN <strong>of</strong> the<br />
<strong>Biological</strong> <strong>Society</strong><br />
<strong>of</strong> <strong>Washington</strong><br />
ISSN 0097-0298<br />
checklist <strong>of</strong> freshwater fishes <strong>of</strong> the guiana shield<br />
10 September 2009<br />
NUMBER 17
Bulletin Editor: Stephen L. Gardiner<br />
Review Editor: Bruce B. Collette<br />
Copies available as the supply lasts from:<br />
The Custodian <strong>of</strong> Publications<br />
<strong>Biological</strong> <strong>Society</strong> <strong>of</strong> <strong>Washington</strong><br />
MRC 116<br />
National Museum <strong>of</strong> Natural History<br />
P.O. Box 37012<br />
<strong>Washington</strong>, D.C. 20013-7012<br />
(Cost $20.00, including postage and handling)<br />
© <strong>Biological</strong> <strong>Society</strong> <strong>of</strong> <strong>Washington</strong>, 2009
CHECKLIST OF THE FRESHWATER FISHES OF THE<br />
GUIANA SHIELD<br />
Richard P. Vari, Carl J. Ferraris, Jr., Aleksandar Radosavljevic, and Vicki A. Funk<br />
i
Front cover illustration: Pseudolithoxus dumus, family Loricariidae (see Plate 12, Figure G).<br />
Illustrations facing each section:<br />
For the Introduction, montage <strong>of</strong> radiographs <strong>of</strong> fishes from the rivers <strong>of</strong> the Guiana Shield, upper left –<br />
Rhaphiodon vulpinis, Cynodontidae; upper right – Prochilodus mariae, Prochilodontidae; center – Serrasalmus<br />
irritans, Characidae, Serrasalminae; lower left – Corydoras filamentosus, Callichthyidae; and lower right,<br />
Sternarchorhynchus roseni, Apteronotidae, mature male with enlarged dentary dentition. Images and plate<br />
prepared by S. Raredon.<br />
For the Fishes <strong>of</strong> the Guiana Shield, Acnodon oligocanthus (Serrasalminae, juvenile) from Steindachner, F. 1915.<br />
Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen<br />
Classe, Wien 93:15–105 (date based on release <strong>of</strong> separates <strong>of</strong> main work published in 1917).<br />
For the Guide to the Checklist, Acroronia nassa (Cichlidae) from Steindachner, F. 1875. Sitzungsberichte der<br />
Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen Classe, Wien 71:61–137.<br />
For the Photographic Atlas <strong>of</strong> Fishes <strong>of</strong> the Guiana Shield, Leptodoras hasemani (Doradidae) from Steindachner,<br />
F. 1915. Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen<br />
Classe, Wien 93:15–105 (date based on release <strong>of</strong> separates <strong>of</strong> main work published in 1917).<br />
Preferred citations:<br />
Vari, R. P., C. J. Ferraris, Jr., A. Radosavljevic, & V. A. Funk, eds., 2009. Checklist <strong>of</strong> the freshwater fishes <strong>of</strong> the<br />
Guiana Shield.—Bulletin <strong>of</strong> the <strong>Biological</strong> <strong>Society</strong> <strong>of</strong> <strong>Washington</strong>, no. 17.<br />
or, e.g.,<br />
Vari, R. P., & C. J. Ferraris, Jr. 2009. Fishes <strong>of</strong> the Guiana Shield. In Vari, R. P., C. J. Ferraris, Jr., A.<br />
Radosavljevic, & V. A. Funk, eds., 2009. Checklist <strong>of</strong> the freshwater fishes <strong>of</strong> the Guiana Shield.—Bulletin <strong>of</strong><br />
the <strong>Biological</strong> <strong>Society</strong> <strong>of</strong> <strong>Washington</strong>, no. 17.<br />
ii
CONTENTS<br />
CONTRIBUTORS. . . ......................................................... iv<br />
ABSTRACT . . . ............................................................. vii<br />
INTRODUCTION. . . .........................................................<br />
Vicki A. Funk and Carol L. Kell<strong>of</strong>f<br />
1<br />
FISHES OF THE GUIANA SHIELD. .............................................<br />
Richard P. Vari and Carl J. Ferraris, Jr.<br />
9<br />
GUIDE TO THE CHECKLIST . .................................................<br />
Aleksandar Radosavljevic<br />
21<br />
CHECKLIST. . . ............................................................. 25<br />
Pristiformes . ............................................................. 25<br />
Mylobatiformes . . ......................................................... 25<br />
Osteoglossiformes. ......................................................... 25<br />
Anguilliformes . . . ......................................................... 25<br />
Clupeiformes ............................................................. 25<br />
Characiformes . . . ......................................................... 25<br />
Siluriformes . ............................................................. 36<br />
Gymnotiformes . . ......................................................... 45<br />
Cyprinodontiformes . . . ..................................................... 47<br />
Beloniformes ............................................................. 48<br />
Synbranchiformes. ......................................................... 48<br />
Perciformes . ............................................................. 48<br />
Pleuronectiformes. ......................................................... 51<br />
Tetraodontiformes ......................................................... 51<br />
Lepidosireniformes ......................................................... 51<br />
PHOTOGRAPHIC ATLAS OF FISHES OF THE GUIANA SHIELD. .....................<br />
Mark Sabaj Pérez<br />
INTRODUCTION<br />
53<br />
APPENDIX: PLATES ..................................................... 61<br />
INDEX TO ORDERS, FAMILIES, AND SUBFAMILIES . ............................. 95<br />
iii
CONTRIBUTORS<br />
James S. Albert, <strong>Department</strong> <strong>of</strong> Biology, University <strong>of</strong> Louisiana at Lafayette, Lafayette, Louisiana 70504-2451,<br />
U.S.A., e-mail: jxa4003@louisiana.edu<br />
Jonathan W. Armbruster, <strong>Department</strong> <strong>of</strong> <strong>Biological</strong> Sciences, 331 Funchess, Auburn University, Alabama 36849,<br />
U.S.A., e-mail: armbrjw@auburn.edu<br />
Paulo A. Buckup, Departamento de Vertebrados, Museu Nacional, Quinta da Boa Vista, 20940-040 Rio de<br />
Janeiro, RJ, Brazil, e-mail: buckup@acd.ufrj.br<br />
Ricardo Campos-da-Paz, Escola de Ciências Biológicas, Universidade Federal do Estado do Rio de Janeiro, Av.<br />
Pasteur, 458/sala 408, Urca - Rio de Janeiro, RJ, 22290-240, Brazil, e-mail: rcpaz@acd.ufrj.br<br />
Marcelo R. de Carvalho, Instituto de Biociências, Universidade de Sâo Paulo, Rua do Matão, Trav. 14, no. 101,<br />
São Paulo, SP, 05508-900, Brazil, e-mail: mrcarvalho@ib.usp.br<br />
Lilian Cassati, Laboratório de Ictiologia, Departamento de Zoologia e Botânica, IBILCE-UNESP, Rua Cristóvão<br />
Colombo, 2265, São José do Rio Preto, SP, 15054-000, Brazil, e-mail: licasatti@hotmail.com<br />
Ning Labbish Chao, Universidade Federal do Amazonas, Caixa Postal 3695, Manaus, AM 69051-970, Brazil,<br />
e-mail: piabas@gmail.com<br />
Bruce B. Collette, National Marine Fisheries Service, Systematics Laboratory MRC 153, National Museum <strong>of</strong><br />
Natural History, <strong>Washington</strong> D.C. 20560-0153, U.S.A., e-mail: collettb@si.edu<br />
Wilson J. E. M. Costa, Laboratório de Ictiologia Geral e Aplicada, Departamento de Zoologia -- UFRJ, Caixa<br />
Postal 68049, 21944-970 Rio de Janeiro, RJ, Brazil, e-mail: wcosta@acd.ufrj.br<br />
Fabio di Dario, Universidade Federal do Rio de Janeiro, Av. São José do Barreto, São José do Barreto, Caixa<br />
Postal 119331, Macae, RJ, 27921-550, Brazil, e-mail: didario@gmail.com<br />
Carl J. Ferraris, Jr., 2944 NE Couch St., Portland, OR 97232, USA; and Division <strong>of</strong> Fishes, National Museum <strong>of</strong><br />
Natural History, Smithsonian Institution, <strong>Washington</strong>, D.C. 20560-0159, U.S.A., e-mail: carlferraris@comcast.net<br />
John P. Friel, Museum <strong>of</strong> Vertebrates, Cornell University, E151 Corson Hall, Ithaca, New York 14853-2701,<br />
U.S.A., e-mail: jpf19@cornell.edu<br />
Vicki A. Funk, <strong>Department</strong> <strong>of</strong> <strong>Botany</strong>, National Museum <strong>of</strong> Natural History, Smithsonian Institution,<br />
<strong>Washington</strong>, D.C. 20560-0166, U.S.A., e-mail: funkv@si.edu<br />
Michel Jégu, Antenne IRD, UR 131, Laboratoire d’Ichtyologie, M.N.H.N., 43 rue Cuvier, 75231 Paris Cedex 05,<br />
France, e-mail: jegu@mnhn.fr<br />
Carol L. Kell<strong>of</strong>f, <strong>Department</strong> <strong>of</strong> <strong>Botany</strong>, National Museum <strong>of</strong> Natural History, Smithsonian Institution,<br />
<strong>Washington</strong>, D.C. 20560-0166, U.S.A., e-mail: kell<strong>of</strong>fc@si.edu<br />
Sven O. Kullander, <strong>Department</strong> <strong>of</strong> Vertebrate Zoology, Swedish Museum <strong>of</strong> Natural History, P. O. Box 50007, SE-<br />
104 05 Stockholm, Sweden, e-mail: sven.kullander@nrm.se<br />
Francisco Langeani Neto, Laboratório de Ictiologia, Departamento de Zoologia e Botânica, IBILCE-UNESP, Rua<br />
Cristóvão Colombo, 2265, São José do Rio Preto, SP, 15054-000, Brazil, e-mail: langeani@zoo.ibilce.unesp.br<br />
Flávio C. T. Lima, Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42594, São Paulo, SP, 04299-970,<br />
Brazil, e-mail: fctlima@usp.br<br />
Rosana S. Lima, Universidade de Estado de Rio de Janeiro, Faculdade de Formação de Pr<strong>of</strong>essores, Rua Dr.<br />
Francisco Portela, 794, São Gonçalo, RJ, 24435-000, Brazil, e-mail: rosanasl@yahoo.com.br<br />
Carlos A. S. Lucena, Laboratory <strong>of</strong> Ichthyology, Museu de Ciências e Tecnologia PUCRS, Caixa Postal 1429,<br />
Porto Alegre, RS, 90619-900, Brazil, e-mail: lucena@pucrs.br<br />
iv
Paulo H. F. Lucinda, Laboratório de Ictiologia, Universidade do Tocantins - Campus de Porto Nacional Caixa<br />
Postal 25, Porto Nacional, TO, 77500-000, Brazil, e-mail: plucinda@unitins.br<br />
John G. Lundberg, The Academy <strong>of</strong> Natural Sciences, <strong>Department</strong> <strong>of</strong> Ichthyology, 1900 Benjamin Franklin<br />
Parkway, Philadelphia, Pennsylvania 19103, U.S.A., e-mail: lundberg@acnatsci.org<br />
Luiz R. Malabarba, Departamento de Zoologia -- IB, Universidade Federal do Rio Grande do Sul, Av. Bento<br />
Gonçalves, 9500 - bloco IV - Prédio 43435, Porto Alegre, RS, 91501-970, Brazil, e-mail: malabarb@ufrgs.br<br />
John D. McEachran, <strong>Department</strong> <strong>of</strong> Wildlife & Fishery Science, Texas A&M University, 22587 AMU, College<br />
Station, Texas 77843-2258, U.S.A., e-mail: jmceachran@tamu.edu<br />
Naércio A. Menezes, Museu de Zoologia, Universidade de São Paulo, Caixa Postal 42594, São Paulo, SP, 04299-<br />
970, Brazil, e-mail: naercio@usp.br<br />
Cristiano Moreira, Departamento de Ciências Biológicas, Universidade Federal de São Paulo, Rua Pr<strong>of</strong>. Artur<br />
Riedel, 275, Diadema, SP, 09972-270, Brazil, e-mail: Moreira.c.r@gmail.com<br />
Carla S. Pavanelli, Fundação Universidade Estadual de Maringá, Nupelia, Av. Colombo, 3690, Maringá, PR,<br />
87020-900, Brazil, e-mail: carlasp@nupelia.uem.br<br />
Aleksandar Radosavljevic, The City College <strong>of</strong> New York, 526 Marshal Science Building, 160 Convent Avenue,<br />
New York, New York 10031, U.S.A., e-mail: alex.radosavljevic@gmail.com<br />
Robson T. C. Ramos, Depto de Sistemática e Ecologia, Universidade Federal da Paraíba, PB, João Pessoa, 58059-<br />
900, Brazil, e-mail: robtamar@dse.ufpb.br<br />
Roberto E. Reis, Laboratory <strong>of</strong> Ichthyology, Museu de Ciências e Tecnologia PUCRS, Caixa Postal 1429, Porto<br />
Alegre, RS, 90619-900, Brazil, e-mail: reis@pucrs.br<br />
Ricardo S. Rosa, Depto de Sistemática e Ecologia, Universidade Federal da Paraíba, João Pessoa, PB, 58059-900,<br />
Brazil, e-mail: rsrosa@dse.ufpb.br<br />
Mark Sabaj Pérez, The Academy <strong>of</strong> Natural Sciences, <strong>Department</strong> <strong>of</strong> Ichthyology, 1900 Benjamin Franklin<br />
Parkway, Philadelphia, Pennsylvania 19103, U.S.A., e-mail: sabaj@acnatsci.org<br />
Scott A. Schaefer, Division <strong>of</strong> Vertebrate Zoology, American Museum <strong>of</strong> Natural History, Central Park West at<br />
79th Street, New York, New York 10024-5192, U.S.A., e-mail: schaefer@amnh.org<br />
Oscar A. Shibatta, Departamento de Biologia Animal e Vegetal, Centro de Ciências Biológicas, Universidade<br />
Estadual de Londrina, 86051-990 Londrina, PR, Brazil, e-mail: shibatta@uel.br<br />
Mônica Toledo-Piza, Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Caixa<br />
Postal 11461, 05422-970 São Paulo, SP, Brazil, e-mail: mtpiza@usp.br<br />
Richard P. Vari, Division <strong>of</strong> Fishes, National Museum <strong>of</strong> Natural History, Smithsonian Institution, <strong>Washington</strong>,<br />
D.C. 20560-0159, U.S.A., e-mail: varir@si.edu<br />
Claude Weber, <strong>Department</strong> d’Herpetologie et d’Ichtyologie, Museum d’Histoire Naturelle, P. O. Box 434, CH-<br />
1211 Genève, Switzerland, e-mail: claude.weber@mhn.ville-ge.ch<br />
Marilyn Weitzman, Division <strong>of</strong> Fishes, National Museum <strong>of</strong> Natural History, Smithsonian Institution,<br />
<strong>Washington</strong>, D.C. 20560-0159, U.S.A., e-mail: weitzmam@si.edu<br />
Stanley H. Weitzman, Division <strong>of</strong> Fishes, National Museum <strong>of</strong> Natural History, Smithsonian Institution,<br />
<strong>Washington</strong>, D.C. 20560-0159, U.S.A., e-mail: weitzmas@si.edu<br />
Wolmar Wosiacki, Museu Paraense Emílio Goeldi, Laboratório de Ictiologia, Av. Magalhães Barata 376 Caixa<br />
Postal Box 399, Belém, PA, 66040-170, Brazil, e-mail: wolmar@museu-goeldi.br<br />
Angela M. Zanata, Departamento de Zoologia, Universidad Federal da Bahia, Rua Barão de Geremoabo, Campus<br />
de Ondina, Salvador, BA, 40170-290, Brazil, e-mail: a_zanata@yahoo.com.br<br />
v
Abstract.—Distributions are provided for 1168 species <strong>of</strong> fishes that live in the<br />
freshwater drainage systems overlying the Guiana Shield in Brazil, Colombia,<br />
French Guiana, Guyana, and Suriname. This total includes representatives <strong>of</strong><br />
376 genera, 49 families, and 15 orders, with five orders (Characiformes,<br />
Siluriformes, Perciformes, Gymnotiformes, and Cyprinodontiformes) accounting<br />
for 96.7% <strong>of</strong> the diversity. Drainage systems on the Guiana Shield are home<br />
to approximately 23% <strong>of</strong> the freshwater species known from Central and South<br />
America. A summary is provided <strong>of</strong> ichthyological collecting on the Shield along<br />
with summaries <strong>of</strong> major publications dealing with fishes <strong>of</strong> each region on the<br />
Shield. Factors that may account for the high species level diversity in that<br />
region are discussed. Methods for photographing fishes are detailed, and a<br />
photographic album <strong>of</strong> 127 species <strong>of</strong> 46 families that occur on the Shield is<br />
included.<br />
Key Words.—Guiana Shield Fishes, Freshwaters, Brazil, Colombia, French<br />
Guiana, Guyana, Suriname, Venezuela.<br />
vii
In the face <strong>of</strong> the growing biodiversity crisis, we must<br />
move to document and evaluate the biota <strong>of</strong> our<br />
natural areas. As taxonomists, it is part <strong>of</strong> our job to<br />
name, place into groups, and keep track <strong>of</strong> the species<br />
that we study. Often we use the distribution <strong>of</strong> the taxa<br />
to help with our work and to provide a broader context<br />
for the results. In doing this job, taxonomists produce<br />
checklists, floras and faunas, and monographs. In fact,<br />
collecting specimens and producing these documents<br />
are essential elements in our quest to understand the<br />
natural world and how it evolved. Checklists are the<br />
‘‘first pass’’ in our attempts to understand the diversity<br />
<strong>of</strong> an area. They give us the first approximation <strong>of</strong> the<br />
known diversity <strong>of</strong> any group or groups <strong>of</strong> organisms<br />
and they <strong>of</strong>ten provide the first pr<strong>of</strong>ile <strong>of</strong> the biodiversity<br />
in relatively poorly known parts <strong>of</strong> the globe. For<br />
many groups, they may represent the most complete<br />
information available and since basic taxonomic data<br />
provide the essential information for studies in systematic<br />
and evolutionary biology, we continually seek<br />
to improve and update it. Checklists have many uses:<br />
they are aids in the identification and correct naming <strong>of</strong><br />
species, they serve as essential resources for biodiversity<br />
estimates and biogeographic studies, and they are<br />
starting points for more detailed studies <strong>of</strong> an area’s<br />
biota. When reviewed by many specialists, they <strong>of</strong>ten<br />
represent the most advanced state <strong>of</strong> knowledge available<br />
in the field. But they do more than that, beyond<br />
these basics they increase our knowledge <strong>of</strong> this fascinating<br />
region and act as a starting point for additional<br />
biodiversity research because they give insight into the<br />
species richness, endemicity, and floral and faunal<br />
affinities for the region and provide the baseline<br />
information for analyzing species turnover rates and<br />
migration or invasion events and many other biological<br />
phenomena (e.g., Funk & Richardson 2002, Funk et al.<br />
2002, 2005; Ferrier et al. 2004, Kell<strong>of</strong>f & Funk 2004).<br />
Checklists also increase national and regional pride by<br />
demonstrating the diversity <strong>of</strong> the area and provide<br />
important public outreach and fundamental information<br />
to be used by governments and conservation<br />
organizations in addressing the biodiversity crisis. This<br />
Checklist <strong>of</strong> the Fishes <strong>of</strong> the Guiana Shield, when<br />
combined with the recently published Checklist <strong>of</strong> the<br />
Terrestrial Vertebrates <strong>of</strong> the Guiana Shield (Hollowell<br />
& Reynolds 2005), and Checklist <strong>of</strong> the Plants <strong>of</strong> the<br />
Guiana Shield (Funk et al. 2007), provides a sound<br />
basis for future research and conservation efforts.<br />
This Checklist <strong>of</strong> the Fishes <strong>of</strong> the Guiana Shield is an<br />
excellent example <strong>of</strong> effective interaction, involving<br />
ichthyologists from around the world who contributed<br />
in diverse ways to its completion. It began as an<br />
INTRODUCTION<br />
V. A. FUNK and CAROL L. KELLOFF<br />
extraction from Reis et al. (2003) but quickly grew into<br />
a larger and more involved project. As with any<br />
endeavor <strong>of</strong> this type, insights that specialists have<br />
gained through their experience are irreplaceable in<br />
correcting errors and updating classifications. In order<br />
to make future editions <strong>of</strong> this checklist as current and<br />
accurate as possible, specialists are encouraged to<br />
contact the Smithsonian’s <strong>Biological</strong> Diversity <strong>of</strong> the<br />
Guiana Shield Program with additions or corrections.<br />
The Guiana Shield<br />
The Guiana Shield region is a biologically rich area<br />
that includes much <strong>of</strong> northeastern South America<br />
(Fig. 1). It is strictly defined by the underlying<br />
geological formation known as the Guiana Shield,<br />
and in the context <strong>of</strong> this volume the term Guiana<br />
Shield also refers to the corresponding geographic<br />
region. That region includes the Venezuelan states <strong>of</strong><br />
Bolívar and Amazonas, and a portion <strong>of</strong> Delta<br />
Amaçuro; all <strong>of</strong> Guyana, Suriname, and French<br />
Guiana; and parts <strong>of</strong> northern Brazil. Several geological<br />
outliers <strong>of</strong> the Guiana Shield occur west <strong>of</strong> the<br />
Orinoco River in Colombia. In Spanish and Portuguese<br />
speaking countries, the region is <strong>of</strong>ten referred to<br />
as the ‘‘Guayana’’; thus the terms Colombian<br />
Guayana, Brazilian Guayana, and Venezuelan<br />
Guayana are <strong>of</strong>ten used. The total area <strong>of</strong> the Guiana<br />
Shield is approximately 1,520,000 km 2 . Table 1 lists the<br />
square kilometers <strong>of</strong> political divisions that occur<br />
within the region. See Berry et al. (1995) for a review<br />
<strong>of</strong> definitions <strong>of</strong> and terminology related to the Guiana<br />
Shield region.<br />
Geology<br />
The Guiana Shield (Fig. 1) is a distinct geologic<br />
region that underlies Guyana, Suriname, French<br />
Guiana, parts <strong>of</strong> Venezuela, and Brazil, and a small<br />
area in Colombia. It is roughly bounded by the<br />
Atlantic Ocean to the north and east, the Orinoco<br />
River to the north and west, the Río Negro (a major<br />
tributary <strong>of</strong> the Amazon River) to the southwest, and<br />
the Amazon River to the south (Gibbs & Barron 1993).<br />
The Guiana Shield is a distinct ancient geological<br />
formation that includes the mountain systems that<br />
form the watershed boundary between the Amazon<br />
and Orinoco rivers. On the Shield’s western side, the<br />
Orinoco River and Río Negro are connected by the<br />
Río Casiquiare, making most <strong>of</strong> the region somewhat<br />
like an island; however, there are some areas <strong>of</strong> nearby<br />
Colombia that have remnants <strong>of</strong> the Shield formation.<br />
The southern boundary <strong>of</strong> the Guiana Shield is
2 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Figure 1. The Guiana Shield; adapted from Gibbs & Barron (1993), with the region <strong>of</strong> western outliers indicated.<br />
difficult to define precisely, as a broad band <strong>of</strong> outwash<br />
materials resulting from erosion occurs between<br />
mountains on the southern boundary <strong>of</strong> the Shield<br />
and the Amazon and Negro rivers stretching into parts<br />
<strong>of</strong> northern Brazil. Also, much <strong>of</strong> the Venezuelan state<br />
<strong>of</strong> Delta Amacuro occurs over thick sediments<br />
deposited primarily by the Orinoco River and is really<br />
not strictly part <strong>of</strong> the Shield; however, some mountains<br />
<strong>of</strong> the Guiana Shield occur in this state’s southern<br />
section, and a large proportion <strong>of</strong> the sediments <strong>of</strong> the<br />
delta are derived from outwash from the highlands <strong>of</strong><br />
the Shield. For more detailed discussions <strong>of</strong> the<br />
Table 1.—Divisions <strong>of</strong> the Guiana Shield in approximately west to<br />
east arrangement, with abbreviations used and estimated areas.<br />
Division Abbr. Area (km 2 )<br />
*Colombian Guayana CG 120,325<br />
Amazonas, Venezuela VA 175,750<br />
Bolívar, Venenzuela BO 238,000<br />
*Delta Amacuro, Venezuela DA 40,200<br />
*Amazonas, Brazil BA 125,550<br />
*Roraima, Brazil RO 173,750<br />
*Pará, Brazil PA 243,280<br />
*Amapá, Brazil AP 98,750<br />
Guyana GU 214,970<br />
Suriname SU 163,270<br />
French Guiana FG 91,000<br />
Total (km 2 ) 1,896,845<br />
*5 not all parts <strong>of</strong> these politically defined areas are included in<br />
the Guiana Shield region.<br />
geology <strong>of</strong> the area, readers should refer to Gibbs &<br />
Barron (1993) and Huber (1995a).<br />
The base <strong>of</strong> the Guiana Shield is composed <strong>of</strong><br />
crystalline rocks <strong>of</strong> Proterozoic origin; these are mainly<br />
granites and gneisses formed between 3.6 and 0.8<br />
billion years ago (Mendoza 1977, Schubert & Huber<br />
1990). Large areas <strong>of</strong> the Shield were overlain with<br />
sediments from 1.6 to 1 billion years ago and cemented<br />
during thermal events (Huber 1995a). These quartzite<br />
and sandstone rocks comprise the Roraima formation<br />
and today remnants are scattered across the central<br />
portion <strong>of</strong> the Shield extending west from the Potaro<br />
Plateau <strong>of</strong> the Pakaraima Mountains in central<br />
Guyana through parts <strong>of</strong> Venezuela and Colombia<br />
(Arbeláez & Callejas 1999) and south into northern<br />
Brazil (Leechman 1913, Gibbs & Barron 1993). Within<br />
this area, erosion has resulted in numerous verticalwalled,<br />
frequently flat-topped mountains called ‘‘tepuis,’’<br />
among them are Chimantí-tepui (2550 m), Cerro<br />
Duida (2358 m), and Auyán-tepui (2450 m). Pico de<br />
Neblina (3014 m) is the Shield’s western-most tepui<br />
and highest point, located on the southern-most<br />
segment <strong>of</strong> the border between Venezuela and Brazil.<br />
Mount Roraima (2810 m) is located at the juncture <strong>of</strong><br />
Guyana, Venezuela, and Brazil and includes the<br />
highest point within Guyana. The eastern-most peaks<br />
in Guyana reach approximately 2000 m elevation,<br />
including Mt. Ayanganna (2041 m). Several <strong>of</strong> these<br />
mesa-like formations are virtually inaccessible by foot
NUMBER 17 3<br />
and are so unusual that they inspired a fictional<br />
scientific expedition referred to one as ‘‘The Lost<br />
World’’ (Doyle 1912), a term sometimes applied to all<br />
tepuis. Notable waterfalls <strong>of</strong> the region include Angel<br />
Falls (979 m) on Auyán-tepui in Venezuela and<br />
Kaieteur Falls (226 m), which flows year around, on<br />
the eastern-most edge <strong>of</strong> the Roraima formation<br />
known as the Potaro Plateau in Guyana.<br />
Granitic dome mountains occur on the Shield in the<br />
southern part <strong>of</strong> the three Guianas (Guyana, Suriname,<br />
French Guiana), where they are known as<br />
‘‘inselbergs,’’ as well as in the western extreme <strong>of</strong> the<br />
Shield in the Puerto Ayacucho region in Venezuela<br />
where they are called ‘‘lajas.’’ Deposits <strong>of</strong> low-nutrient<br />
white sands occur inland <strong>of</strong> the coastal plain, in belts<br />
across the Shield, and in isolated pockets. Large areas<br />
<strong>of</strong> savanna are found in the region, particularly the<br />
complex <strong>of</strong> savannas that includes the Rupununi<br />
Savanna in southwestern Guyana, the Gran Sabana<br />
in eastern Venezuela, and the savannas <strong>of</strong> northern<br />
Roraima, Brazil. In some <strong>of</strong> these areas the sands<br />
overlay a clay hardpan that is resistant to penetration<br />
by tree roots and that floods during the heavy rainy<br />
season, resulting in limited forest growth. Tertiary and<br />
Quaternary sediments separate the southern edge <strong>of</strong><br />
the Guiana Shield from the Amazon River and the<br />
eastern edge from the Atlantic Ocean.<br />
Climate<br />
As a whole, the Guiana Shield region has a tropical<br />
climate characterized by a relatively high mean annual<br />
temperature exceeding 25uC at sea level, an annual<br />
monthly maximum temperature range <strong>of</strong> less than 5uC,<br />
and an average daily temperature range <strong>of</strong> approximately<br />
6uC (Snow 1976). Because <strong>of</strong> the Guiana<br />
Shield’s location just north <strong>of</strong> the equator, its climate<br />
varies primarily according to elevation and effects <strong>of</strong><br />
the trade winds that combine to affect rainfall patterns.<br />
The trade winds blow consistently from the east and<br />
northeast, <strong>of</strong>f <strong>of</strong> the Atlantic Ocean onto northeastern<br />
South America, with wind speeds averaging from 3–4<br />
m per second. Due to orographic effects, the easternmost<br />
escarpments <strong>of</strong> the mountains <strong>of</strong> the Guiana<br />
Shield are generally localities <strong>of</strong> increased precipitation<br />
where these moisture-laden winds meet the slopes<br />
(Clarke et al. 2001). Seasonal oscillations <strong>of</strong> the<br />
Intertropical Convergence Zone (ITCZ) also bring<br />
variations in rainfall as the locations <strong>of</strong> low pressure<br />
zones near the equator change (Snow 1976). Varying<br />
primarily by latitude, one or two rainy seasons result<br />
from shifts in the ITCZ. The heaviest rains usually<br />
occur between May and August, whereas the rainy<br />
season running from December to January is shorter<br />
and less intense, with rains that do not penetrate as far<br />
inland. Even during most dry seasons, frequent storms<br />
provide adequate moisture to allow evergreen tropical<br />
moist forests to persist in most low elevation parts <strong>of</strong><br />
the region.<br />
<strong>Biological</strong> Diversity<br />
The variety <strong>of</strong> landscapes <strong>of</strong> the Guiana Shield<br />
includes sandstone tepuis, granite inselbergs, white<br />
sands, seasonally flooded tropical savannas, lowlands<br />
with numerous rivers, isolated mountain ranges, and<br />
coastal swamps, each supporting a characteristic<br />
vegetation (Huber 1995b, Huber et al. 1995). This<br />
variety accounts for a great deal <strong>of</strong> the high diversity<br />
and endemicity <strong>of</strong> the Shield’s biota. The highlands <strong>of</strong><br />
the Shield have a flora and fauna with numerous<br />
endemic species. Some tepui endemic species occur as<br />
low as 300 m in elevation, with increasing numbers by<br />
1500 to 1800 m, and fully developed communities<br />
occurring by 2000 m. Few if any plant or animal<br />
specimens have been collected from most medium to<br />
high elevation areas <strong>of</strong> the Guiana Shield. Many parts<br />
<strong>of</strong> the Shield are poorly explored, including parts <strong>of</strong><br />
Brazil north <strong>of</strong> the Amazon River, much <strong>of</strong> eastern<br />
Colombia, and the southern parts <strong>of</strong> Venezuela,<br />
Guyana, Suriname, and French Guiana.<br />
Conservation<br />
With the exceptions <strong>of</strong> the populated localities such<br />
as Puerto Ayacucho, Ciudad Guayana, Ciudad Bolívar,<br />
Boa Vista, Georgetown, Paramaribo, Cayenne, the<br />
agricultural coastal areas, and open areas like the<br />
Rupununi Savanna, the environment <strong>of</strong> the Guiana<br />
Shield has benefited from limited access and low<br />
population densities, although this same isolation has<br />
hindered biodiversity research. Estimates vary, but<br />
much <strong>of</strong> the vegetation is still relatively undisturbed by<br />
human activities. Recently, however, the pace <strong>of</strong><br />
disturbance has greatly increased. Current threats to<br />
the environment include large-scale logging by Asian<br />
and local companies, large- and small-scale gold and<br />
diamond mining, oil prospecting, bauxite mining,<br />
hydroelectric dams, wildlife trade, and populationrelated<br />
pressures such as burning, grazing, agriculture,<br />
and the expansion <strong>of</strong> towns and villages. Taken<br />
together, these impacts have begun to take their toll,<br />
with vast areas vulnerable to increasing disturbance a<br />
fact easily observed by using Google Earth and<br />
‘‘flying’’ over the area.<br />
The status <strong>of</strong> conservation efforts varies by country.<br />
Throughout the Guiana Shield, many areas that are<br />
designated as protected or otherwise restricted are<br />
<strong>of</strong>ten only ‘‘paper’’ parks because <strong>of</strong> a lack <strong>of</strong><br />
infrastructure, funds, and will to actually protect the<br />
areas. Over the last four decades, Venezuela has<br />
established seven national parks, 29 natural monuments,<br />
and two biosphere reserves covering about<br />
142,280 km 2 , more than 30% <strong>of</strong> its share <strong>of</strong> the
4 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Guiana Shield (Funk & Berry 2005). In Guyana, the<br />
progress <strong>of</strong> conservation efforts has been slower, with<br />
the only substantial protected area being Kaieteur<br />
National Park, its 627 km 2 comprising about 3% <strong>of</strong><br />
the country’s area (Kell<strong>of</strong>f 2003, Kell<strong>of</strong>f & Funk<br />
2004), with additional reserves under consideration.<br />
Guyana’s 3710 km 2 Iwokrama forest (Clarke et al.<br />
2001) has parts listed as reserves, but overall it is<br />
dedicated to sustainable use; unfortunately, logging<br />
has begun, and the section <strong>of</strong> the road from Boa Vista,<br />
Brazil, to Georgetown, Guyana, that runs through<br />
Iwokrama is about to be paved. Suriname’s protected<br />
areas system includes one national park and a network<br />
<strong>of</strong> 11 reserves, totaling almost 20,000 km 2 , over 12%<br />
<strong>of</strong> its total area. This includes the recently created<br />
16,000 km 2 Central Suriname Nature Reserve, a<br />
UNESCO World Heritage Site that joined and<br />
expanded three existing reserves (see http://www.<br />
stinasu.com). French Guiana has no <strong>of</strong>ficially designated<br />
protected areas, but 18 proposed sites total 6710<br />
km 2 , about 7.5% <strong>of</strong> its area (Lindeman & Mori 1989).<br />
The natural areas <strong>of</strong> Venezuela and Guyana are<br />
currently under the most anthropogenic pressure,<br />
while those <strong>of</strong> French Guiana are probably less<br />
threatened.<br />
The Shield encompasses part or all <strong>of</strong> six countries<br />
with six different governments, five <strong>of</strong>ficial languages<br />
and many more indigenous languages. Cooperation is<br />
sometimes hampered by border disputes, illegal crossborder<br />
activities involving gold and wildlife, and a lack<br />
<strong>of</strong> interest by governments that are located far away.<br />
The implementation <strong>of</strong> conservation practices is<br />
further complicated by many issues concerning the<br />
indigenous peoples <strong>of</strong> the region. All <strong>of</strong> these<br />
challenges will have to be overcome on the way to<br />
designing and maintaining a viable reserve system for<br />
the Guiana Shield. However, it is critical that we gain<br />
an understanding <strong>of</strong> the flora and fauna <strong>of</strong> the Shield<br />
area so that decisions can be made on critical areas that<br />
have high priority for conservation and so data can be<br />
collected from areas that might ultimately be destroyed.<br />
Because it is an ancient, fairly isolated<br />
geological area, it is rich in endemic plant and animal<br />
taxa, with many more likely to be discovered with<br />
additional exploration. In addition, because this area<br />
has been long neglected by biologists, it is <strong>of</strong>ten an area<br />
<strong>of</strong> ‘‘inadequate information’’ for many biodiversity<br />
analyses.<br />
This volume contains the fishes from the Guiana<br />
Shield, when paired with the previously published<br />
Checklist <strong>of</strong> the Terrestrial Vertebrates <strong>of</strong> the Guiana<br />
Shield (Hollowell & Reynolds 2005), we can examine<br />
the size and scope <strong>of</strong> Vertebrates, an important<br />
monophyletic group, known to inhabit the Guiana<br />
Shield. Table 2 lists the vertebrate groups and their<br />
sizes. The two checklists include a total <strong>of</strong> 53 orders,<br />
189 families, 1190 genera, and 301 species. A large<br />
Table 2.—Number <strong>of</strong> vertebrate taxa at different ranks.<br />
Orders Families Genera Species<br />
Amphibians 2 13 59 269<br />
Reptiles 3 22 119 295<br />
Mammals 11 35 143 282<br />
Birds 22 70 493 1004<br />
Fishes 15 49 376 1168<br />
Total 53 189 1190 3018<br />
percentage <strong>of</strong> the species (38%) are contributed by the<br />
fishes listed in this volume.<br />
Figure 2 compares the vertebrate diversity across the<br />
major political areas <strong>of</strong> the Guiana Shield and shows<br />
the species turnover between different areas. The<br />
mammals and reptiles have the most similar fauna<br />
across the Shield with a 58% and 53% overlap,<br />
respectively, between French Guiana (the extreme east)<br />
and Venezuela-Amazonas (the extreme west). Fish and<br />
birds have the least (24% and 10%, respectively). With<br />
fishes this can probably be explained by the fact that<br />
the headwaters <strong>of</strong> the rivers in the east are widely<br />
separated from those <strong>of</strong> the west. The rivers <strong>of</strong> the<br />
Shield in the west (Venezuela) have their source in the<br />
Venezuelan Guayana and the Andes, in the central<br />
portion (Guyana) the Essequibo drains mainly from<br />
the Acari Mountains which lie on the border with<br />
Brazil as does the Corantijn River (border between<br />
Guyana and Suriname). To the east, rivers such as the<br />
Maroni and Oyapock drain from the Tumuk-Humak<br />
Mountains. The bird diversity percentage <strong>of</strong> ‘turn over’<br />
was surprisingly small until one realizes that there are<br />
very different flyways that go across the eastern and<br />
western parts <strong>of</strong> the Shield. When the three major<br />
avenues <strong>of</strong> vertebrate mobility are examined (land, air,<br />
water), it seems that the land provides the most stable<br />
species make up and the air and water provide the<br />
least. Could this have anything to do with the resulting<br />
high species diversity <strong>of</strong> the birds and fishes?<br />
In the wider scope <strong>of</strong> biological understanding, the<br />
goal <strong>of</strong> checklists <strong>of</strong> this type is to understand diversity<br />
in terms <strong>of</strong> the spatial, evolutionary, and ecological<br />
settings <strong>of</strong> physical environments, rather than simply<br />
by political boundaries. The assembly <strong>of</strong> these lists is a<br />
step toward considering the fauna in terms <strong>of</strong> the<br />
geological entity <strong>of</strong> the Guiana Shield. Future studies<br />
will include the analyses <strong>of</strong> animal community composition<br />
on finer landscape scales, using developing<br />
abilities to produce customized checklists for research<br />
and conservation with Geographic Information System<br />
(GIS) technologies drawing upon comprehensive databases<br />
that include georeferenced museum specimen<br />
records.<br />
<strong>Biological</strong> Diversity <strong>of</strong> the Guiana Shield (BDG)<br />
The ‘‘<strong>Biological</strong> Diversity <strong>of</strong> the Guiana Shield<br />
Program’’ (BDG) is a field-oriented program <strong>of</strong> the
NUMBER 17 5<br />
Figure 2. A comparison <strong>of</strong> the species lists <strong>of</strong> the political areas <strong>of</strong> the Guiana Shield Region.<br />
National Museum <strong>of</strong> Natural History that began in<br />
1983 (federally funded since 1987). The goal <strong>of</strong> the<br />
BDG is to ‘‘study, document and preserve the<br />
biological diversity <strong>of</strong> the Guiana Shield.’’ Most <strong>of</strong><br />
the program’s field work has taken place in Guyana,<br />
but data analyses cover the majority <strong>of</strong> the Shield. In<br />
Guyana, the BDG operates under the auspices <strong>of</strong> the<br />
University <strong>of</strong> Guyana (UG). The BDG program is<br />
designed to provide specimens and data to address<br />
questions about many groups <strong>of</strong> organisms from<br />
locations across the Shield. Information from BDG<br />
collections and from other herbarium collections is<br />
used to produce checklists, vegetation maps, floristic<br />
and faunistic studies, revisions, and monographs. The<br />
data generated from these studies are used to ask<br />
questions about the make up <strong>of</strong> Guiana Shield<br />
biological diversity, such as species turnover rates,<br />
surrogate taxa, and areas <strong>of</strong> high diversity. Finally, the<br />
BDG is exploring practical applications <strong>of</strong> the data<br />
that have been collected through regular collaborations<br />
with conservation and government agencies.<br />
In addition to collecting and research, the BDG<br />
Program trains students and scientists in both the<br />
U.S.A. and Guyana, assists in their research, and has<br />
established and helped to maintain collections. Over<br />
the years several events have been hosted in Guyana,<br />
including two Amerindian training courses, two bird<br />
preparation courses, two plant identification courses, a<br />
variety <strong>of</strong> lectures at the University and public venues,<br />
and a public scientific symposium on the biological<br />
diversity <strong>of</strong> Guyana. We also <strong>of</strong>fer training opportunities;<br />
nearly every year since 1987 the Program has<br />
hosted at least one Guyanese student or UG staff<br />
member at the Smithsonian. Many have participated in<br />
the Natural History Museum’s Research Training<br />
Program or the SI/MAB training courses. BDG<br />
worked with the University <strong>of</strong> Guyana to raise funds<br />
from the Royal Bank <strong>of</strong> Canada to construct a new<br />
building, the ‘‘Centre for the Study <strong>of</strong> <strong>Biological</strong><br />
Diversity,’’ located on the campus <strong>of</strong> UG. More<br />
recently, we worked with UG to raise funds from<br />
USAID to build an extension on the original building.
6 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
The Centre houses collections and research space,<br />
provides a library, and houses a Geographic Information<br />
System (GIS) facility. The goal <strong>of</strong> the Centre is to<br />
combine research, education and conservation in the<br />
study <strong>of</strong> biological diversity. The Centre is funded from<br />
outside grants, but the staff is part <strong>of</strong> the University.<br />
Currently, the plant database maintained by BDG has<br />
161,108 specimen records and all sheets have been<br />
barcoded. The BDG Program is working to make its<br />
data available to the scientific community. The<br />
collections are being mapped using ArcMap and then<br />
displayed on Google Earth as place marks. The<br />
project, Georeferencing Plants <strong>of</strong> the Guiana Shield is<br />
available on the <strong>Department</strong> <strong>of</strong> <strong>Botany</strong> public website<br />
(http://botany.si.edu/bdg/georeferencing.cfm).<br />
Acknowledgments<br />
Special thanks go to the University <strong>of</strong> Guyana and<br />
the Guyana EPA who have consistently supported our<br />
efforts, including Mike Tamessar, Indarjit Ramdass,<br />
and Philip da Silva, as well as past and present staff<br />
members <strong>of</strong> the Centre for the Study <strong>of</strong> <strong>Biological</strong><br />
Diversity, in particular Calvin Bernard. This is number<br />
153 in the Smithsonian’s <strong>Biological</strong> Diversity <strong>of</strong> the<br />
Guiana Shield Program publication series.<br />
References<br />
Arbeláez, M. V., & R. Callejas. 1999. Flórula de la Meseta de<br />
Arensica de la comunidad de Monochoa (Región de<br />
Araracuara, Medio Caquetá). Tropenbos, Bogotá, Colombia.<br />
Berry, P. E., B. K. Holst, & K. Yatskievych (eds.). 1995. Flora <strong>of</strong> the<br />
Venezuelan Guayana. Vol. 1: Introduction. J. A. Steyermark,<br />
P. E. Berry, & B. K. Holst, general eds. Missouri Botanical<br />
Garden, St. Louis, 306 pp.<br />
Clarke, H. D., V. Funk, & T. Hollowell. 2001. Using checklists and<br />
collections data to investigate plant diversity. I: a comparative<br />
checklist <strong>of</strong> the plant diversity <strong>of</strong> the Iwokrama Forest,<br />
Guyana.—Sida Botanical Miscellany 21:1–86.<br />
Doyle, A. C. 1912. The Lost World. Puffin Books, London.<br />
Ferrier, S., G. V. N. Powell, K. S. Richardson, G. Manion, J. M.<br />
Overton, T. F. Allnutt, S. E. Cameron, K. Mantle, N. D.<br />
Burgess, D. P. Faith, J. F. Lamoreux, G. Kier, R. J. Hijmans,<br />
V. A. Funk, G. A. Cassis, B. L. Fisher, P. Flemons, D. Lees, J.<br />
C. Lovett, & R. S. A. R. Van Rompaey. 2004. Mapping more<br />
<strong>of</strong> terrestrial biodiversity for Global Conservation assessment.—BioScience<br />
54(12):1101–1109.<br />
Funk, V. A., & P. E. Berry. 2005. The Guiana Shield. Pp. 76–79 in G.<br />
A. Krupnick & W. J. Kress, eds., Plant conservation: a natural<br />
history approach. University <strong>of</strong> Chicago Press, Chicago, 235<br />
pp.<br />
———, & K. S. Richardson. 2002. Systematic data in biodiversity<br />
studies: use it or lose it.—Systematic Biology 51:303–316.<br />
———, ———, & S. Ferrier. 2005. Survey-gap analysis in<br />
expeditionary research: where do we go from here?—<br />
<strong>Biological</strong> Journal <strong>of</strong> the Linnean <strong>Society</strong> 85:549–567.<br />
———, A. K. Sakai, & K. Richardson. 2002. Biodiversity: the<br />
interface between systematics and conservation.—Systematic<br />
Biology 51:235–237.<br />
———, T. Hollowell, P. Berry, C. Kell<strong>of</strong>f, & S. N. Alexander. 2007.<br />
Checklist <strong>of</strong> the plants <strong>of</strong> the Guiana Shield (Venezuela:<br />
Amazonas, Bolívar, Delta Amaçuro; Guyana, Surinam,<br />
French Guiana).—Contributions from the United States<br />
National Herbarium 55:1–584.<br />
Gibbs, A. K., & C. N. Barron. 1993. The geology <strong>of</strong> the Guiana<br />
Shield. Oxford University Press, New York, 246 pp.<br />
Hollowell, T., & R. P. Reynolds (eds.). 2005. Checklist <strong>of</strong> the<br />
terrestrial vertebrates <strong>of</strong> the Guiana Shield.—Bulletin <strong>of</strong> the<br />
<strong>Biological</strong> <strong>Society</strong> <strong>of</strong> <strong>Washington</strong> 13:i–ix + 1–98.<br />
Huber, O. 1995a. Geography and physical features. Pp. 1–61 in P. E.<br />
Berry, B. K. Holst, & K. Yatskievych, eds., Flora <strong>of</strong> the<br />
Venezuelan Guayana. Vol. 1: Introduction. J. A. Steyermark,<br />
P. E. Berry, & B. K. Holst, general eds. Missouri Botanical<br />
Garden, St. Louis.<br />
——— 1995b. Vegetation. Pp. 97–160 in P. E. Berry, B. K. Holst, &<br />
K. Yatskievych, eds., Flora <strong>of</strong> the Venezuelan Guayana. Vol.<br />
1: Introduction. J. A. Steyermark, P. E. Berry, & B. K. Holst,<br />
general eds. Missouri Botanical Garden, St. Louis.<br />
———, G. Gharbarran, & V. A. Funk. 1995. Preliminary vegetation<br />
map <strong>of</strong> Guyana. <strong>Biological</strong> Diversity <strong>of</strong> the Guianas Program,<br />
Smithsonian Institution,<strong>Washington</strong>, D.C.<br />
Kell<strong>of</strong>f, C. L. 2003. The use <strong>of</strong> biodiversity data in developing<br />
Kaieteur National Park, Guyana, for ecotourism and<br />
conservation.—Contributions to the Study <strong>of</strong> <strong>Biological</strong><br />
Diversity, University <strong>of</strong> Guyana, Georgetown 1:1–44.<br />
———, & V. A. Funk. 2004. Phytogeography <strong>of</strong> the Kaieteur Falls,<br />
Potaro Plateau, Guyana: floral distributions and affinities.—<br />
Journal <strong>of</strong> Biogeography 31:501–513.<br />
Leechman, A. 1913. The British Guiana handbook. ‘‘The Argosy’’<br />
Co., Ltd., Georgetown, British Guiana, and Dulau & Co.,<br />
London, 283 pp.<br />
Lindeman, J. C., & S. A. Mori. 1989. The Guianas. Pp. 375–391 in D.<br />
G. Campbell & H. D. Hammond, eds., Floristic inventory <strong>of</strong><br />
tropical countries. New York Botanical Garden, New York.<br />
Mendoza, V. 1977. Evolución tectónica del Escudo de Guayana.—<br />
Boletín de Geología. Publicación Especial 7(3):2237–2270.<br />
Reis, R. E., S. O. Kullander, & C. J. Ferraris, Jr. (eds.). 2003. Check<br />
list <strong>of</strong> the freshwater fishes <strong>of</strong> South and Central America.<br />
Edipucrs, Porto Alegre, Brazil, 729 pp.<br />
Schubert, C., & O. Huber. 1990. The Gran Sabana: panorama <strong>of</strong> a<br />
region. Lagoven Booklets, Caracas, 107 pp.<br />
Snow, J. W. 1976. Climates <strong>of</strong> northern South America. Pp. 295–403<br />
in W. Schwerdtfeger, ed., Climates <strong>of</strong> Central and South<br />
America. Elsevier Scientific Publishing Company, Amsterdam.
History<br />
A vast complex <strong>of</strong> wetlands, lakes, streams, and<br />
rivers drains the broad savannas, dense rainforests,<br />
extensive uplands, and tepuis <strong>of</strong> the Guiana Shield.<br />
Early European explorers and colonists were impressed<br />
not only by the many unusual fish species dwelling in<br />
these water systems but also by the diversity <strong>of</strong> the<br />
ichthy<strong>of</strong>auna. Accounts <strong>of</strong> fishes from those drainage<br />
systems commenced with descriptions by pre-Linnaean<br />
naturalists (e.g., Gronovius 1754, 1756) based on<br />
specimens returned to Europe, with Linnaeus (1758)<br />
formally describing a number <strong>of</strong> these species.<br />
Much <strong>of</strong> the early descriptive activity involving<br />
fishes <strong>of</strong> the Guiana Shield centered on the ichthy<strong>of</strong>auna<br />
<strong>of</strong> British Guiana (5 Guyana). Commentaries<br />
on fishes from that colony by Bancr<strong>of</strong>t (1769) and<br />
Hilhouse (1825) preceded the formal description <strong>of</strong><br />
catfish species from the Demerara by Hancock (1828).<br />
Cuvier and Valenciennes summarized the available<br />
information on the ichthy<strong>of</strong>auna <strong>of</strong> all <strong>of</strong> the Guianas<br />
in their series entitled Histoire Naturelle des Poissons<br />
that documented the fish species known worldwide to<br />
science to that time; with the catfishes being the first <strong>of</strong><br />
the groups inhabiting the shield discussed by those<br />
authors (Cuvier & Valenciennes 1840a, b). These and<br />
the other treatments <strong>of</strong> that era were, however, largely<br />
opportunistic accounts based on scattered samples<br />
returned to Europe rather than derivative <strong>of</strong> focused<br />
studies on the fish fauna <strong>of</strong> any region on the shield.<br />
Consequently, the scale <strong>of</strong> the species-level diversity <strong>of</strong><br />
that ichthy<strong>of</strong>auna remained unknown and underappreciated.<br />
Indications <strong>of</strong> the scale <strong>of</strong> the richness <strong>of</strong> the fish<br />
fauna inhabiting the rivers <strong>of</strong> the Guiana Shield<br />
commenced with the expeditions <strong>of</strong> the Schomburgk<br />
brothers, Richard and Robert. In a remarkable<br />
endeavor for that era, Robert collected fishes from<br />
1835 to 1839 both in the more accessible shorter<br />
northerly flowing rivers <strong>of</strong> the Guianas and through<br />
portions <strong>of</strong> the Río Orinoco and Río Negro and the<br />
Rio Branco. Drawings <strong>of</strong> fishes prepared during<br />
Schomburgk’s travels across the shield served as the<br />
basis for 83 species accounts in Jardine’s ‘‘Naturalist’s<br />
Library’’ (1841, 1843), including the formal descriptions<br />
<strong>of</strong> a series <strong>of</strong> species. Unfortunately, the specimens<br />
that were the basis for the drawings were not<br />
preserved, and some illustrations combined details <strong>of</strong><br />
more than one species. Subsequent expeditions by the<br />
Schomburgks traversed portions <strong>of</strong> what are now<br />
Guyana, Venezuela, Suriname, and Brazil and yielded,<br />
what was for that time, large numbers <strong>of</strong> fish specimens.<br />
In a series <strong>of</strong> publications Müller & Troschel (1845,<br />
FISHES OF THE GUIANA SHIELD<br />
RICHARD P. VARI and CARL J. FERRARIS, JR.<br />
1848, 1849) recognized 141 species in the Schomburgk<br />
collections and provided the first detailed illustrations<br />
<strong>of</strong> fishes from South American freshwaters.<br />
Diverse factors resulted in a lag in the state <strong>of</strong><br />
knowledge <strong>of</strong> the fishes inhabiting many portions <strong>of</strong><br />
the shield, with the comparative difficulty in accessibility<br />
to inland regions clearly a paramount issue for<br />
many areas. Supplementing that impediment were a<br />
series <strong>of</strong> misadventures that bedeviled collectors who<br />
sampled the fish fauna <strong>of</strong> the western portions <strong>of</strong> the<br />
shield. Alexandre Rodriques Ferreira headed an<br />
expedition that explored a significant portion <strong>of</strong> the<br />
Rio Negro basin, commencing with a major collecting<br />
effort through the Rio Branco system in 1786 (Ferreira<br />
et al. 2007:12). Confounding Ferreira’s attempts to<br />
publish his results were a string <strong>of</strong> unfortunate events<br />
that culminated with the 1807 invasion <strong>of</strong> Portugal by<br />
Napoleonic forces and the seizure and shipment <strong>of</strong><br />
Ferreira’s collections to Paris. Ferreira’s report on<br />
animals from the Rio Branco region remained unpublished<br />
until long after his death, and even then, only<br />
parts appeared in print (see references in Ferreira<br />
1983). In two expeditions between 1850 and 1852,<br />
Alfred Russel Wallace (<strong>of</strong> Natural Selection fame)<br />
collected over 200 species <strong>of</strong> fishes throughout the Rio<br />
Negro basin including rivers draining the shield.<br />
Wallace’s collections were lost with the sinking <strong>of</strong> the<br />
ship returning him to England. Nonetheless, his field<br />
sketches (Wallace 2002) document that the lost<br />
collection included a number <strong>of</strong> species <strong>of</strong> fishes then<br />
unknown to science (Regan 1905a, Toledo-Piza et al.<br />
1999, Vari & Ferraris 2006).<br />
An accelerating pace <strong>of</strong> ichthyological collecting<br />
across many portions <strong>of</strong> the Guiana Shield during the<br />
latter part <strong>of</strong> the nineteenth century resulted in the<br />
discovery and description <strong>of</strong> numerous species. These<br />
collections also documented the presence on the Shield<br />
<strong>of</strong> many species originally described from elsewhere in<br />
cis-Andean South America. Notwithstanding those<br />
advances, the information was dispersed through<br />
revisionary (e.g., Regan 1905b, c) and monographic<br />
studies (e.g., Eigenmann & Eigenmann 1890), general<br />
ichthy<strong>of</strong>aunal summaries <strong>of</strong> regions on the shield (e.g.,<br />
Pellegrin 1908), and species descriptions in multiple<br />
languages.<br />
Exceptions to this pattern <strong>of</strong> scattered publication<br />
were limited to a handful <strong>of</strong> papers focused on subsets<br />
<strong>of</strong> the ichthy<strong>of</strong>auna from comparatively small regions.<br />
Among the more notable <strong>of</strong> these were the analysis <strong>of</strong><br />
the catfishes <strong>of</strong> Suriname (Bleeker 1862), discussions <strong>of</strong><br />
the fishes present in portions <strong>of</strong> French Guiana<br />
(Vaillant 1899, 1900), and a semi-popular overview <strong>of</strong>
10 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
the fishes <strong>of</strong> French Guiana (Pellegrin 1908). Compendia<br />
<strong>of</strong> the freshwater fish species known from<br />
individual colonies, countries, or regions were not<br />
developed, let alone summaries <strong>of</strong> the fishes inhabiting<br />
in the numerous streams, rivers, and lakes across the<br />
shield. The dispersed literature prevented an appreciation<br />
<strong>of</strong> the scale <strong>of</strong> the diversity <strong>of</strong> the shield<br />
ichthy<strong>of</strong>auna.<br />
The first overview <strong>of</strong> the freshwater fishes <strong>of</strong><br />
northeastern South America, including the Guiana<br />
Shield, was Eigenmann’s (1912) treatise on the<br />
freshwater fishes <strong>of</strong> British Guiana. Although Eigenmann<br />
sampled the fish fauna <strong>of</strong> only a comparatively<br />
small section <strong>of</strong> British Guiana, his collections were<br />
extensive for that era. In a series <strong>of</strong> papers, he and his<br />
students described 128 new species from those collections<br />
(Eigenmann 1912:133). Eigenmann’s monograph<br />
included data from his own collections, information<br />
from the literature, and records <strong>of</strong> fishes that<br />
originated on the Shield in various museums. Summary<br />
tables (Eigenmann 1912:64) detailed the fish species<br />
known from ten subunits that fall, at least in part,<br />
within the boundaries <strong>of</strong> the Guiana Shield (Río<br />
Orinoco basin, ‘‘West Coast’’ <strong>of</strong> British Guiana<br />
[5 Barima River basin], Rio Branco basin, Rupununi<br />
River, Lower Essequibo River, Lower Potaro River,<br />
Demerara River, Dutch Guiana [5 Suriname], and<br />
French Guiana [5 Guyane Française]).<br />
Eigenmann (1912) reported 493 species from those<br />
ten geographic units; a total that was in excess <strong>of</strong> the<br />
species reported to that time from the rivers <strong>of</strong> the<br />
Shield. These additional species were a function <strong>of</strong> two<br />
factors. His total included all <strong>of</strong> the fish species then<br />
known to inhabit the Río Orinoco; however, that vast<br />
river system extends far beyond the Shield boundaries<br />
with approximately only 40% <strong>of</strong> that watershed<br />
overlying the Shield. Many <strong>of</strong> the fish species known<br />
at the end <strong>of</strong> the first decade <strong>of</strong> the twentieth century<br />
from the Río Orinoco basin originated in the llanos<br />
(savannas) <strong>of</strong> the north central and western portions <strong>of</strong><br />
the basin. Aquatic habitats and the fish faunas in these<br />
floodplain savanna settings differ dramatically from<br />
the ecosystems and fish communities <strong>of</strong> the more<br />
rapidly flowing rivers that drain the forested northern<br />
slope <strong>of</strong> the Guiana Shield. Further inflating Eigenmann’s<br />
species total was his inclusion <strong>of</strong> some<br />
primarily marine forms. Such species penetrate the<br />
lower reaches <strong>of</strong> the rivers draining the Guianas during<br />
periods <strong>of</strong> low river flow and consequent increased<br />
estuarine salinity. Few, if any, <strong>of</strong> these species are likely<br />
to range upriver onto the Shield even during the height<br />
<strong>of</strong> the dry season.<br />
The decades since Eigenmann’s monograph have<br />
seen numerous ichthyological collecting expeditions in<br />
many systems on the Shield. Two wide-ranging and<br />
productive collecting endeavors through that region<br />
during the first half <strong>of</strong> the twentieth century remain<br />
relatively poorly known. John Haseman, who collected<br />
throughout the Rio Branco basin and the southernmost<br />
portion <strong>of</strong> the Rupununi River system in 1912<br />
and 1913, made the first <strong>of</strong> these. Haseman deposited<br />
these extensive collections in the Naturhistorisches<br />
Museum in Vienna where he studied them for a year in<br />
collaboration with Franz Steindachner. Nonetheless,<br />
only one major publication based on those collections<br />
was published (Steindachner 1915), most likely because<br />
<strong>of</strong> the onset <strong>of</strong> World War I, disruptions during and<br />
immediately after the conflict, and the death <strong>of</strong><br />
Steindachner soon after the cessation <strong>of</strong> hostilities.<br />
Various revisionary studies in recent decades incorporated<br />
subsets <strong>of</strong> Haseman’s collection; nonetheless,<br />
much <strong>of</strong> the material is yet-to-be analyzed critically.<br />
The second collector, Carl Ternetz, sampled fishes<br />
through the Rio Negro, Río Casiquiare, and Río<br />
Orinoco basins during 1924 and 1925. Myers<br />
(1927:107) remarked that the collection was ‘‘a<br />
magnificent series <strong>of</strong> fishes, most <strong>of</strong> them hitherto<br />
unexplored systematically by an ichthyologist.’’ Notwithstanding<br />
the description <strong>of</strong> some new species<br />
collected by Ternetz in rivers <strong>of</strong> the shield by Myers<br />
(1927) and other authors and the use <strong>of</strong> portions <strong>of</strong><br />
that collection in some studies (e.g., Myers & Weitzman<br />
1960), most <strong>of</strong> the material remains unstudied,<br />
even after its transfer from Indiana University to the<br />
California Academy <strong>of</strong> Sciences.<br />
The 1960s brought a resurgence <strong>of</strong> major ichthyological<br />
collecting efforts in many <strong>of</strong> the river systems<br />
on the shield (e.g., the Brokopondo Project; Boeseman<br />
1968:4), with the pace <strong>of</strong> these endeavors accelerating<br />
during recent decades. A compendium <strong>of</strong> these<br />
collecting efforts lies beyond the purpose and scope<br />
<strong>of</strong> this paper; however, as summarized in the next<br />
section many <strong>of</strong> these expeditions were integral to<br />
checklists, regional revisionary studies, and summaries<br />
<strong>of</strong> the ichthy<strong>of</strong>auna in river basins or regions <strong>of</strong> the<br />
Shield.<br />
State <strong>of</strong> Knowledge <strong>of</strong> the Shield Fish Fauna<br />
The nearly ten decades since the preparation <strong>of</strong><br />
Eigenmann’s 1912 magnum opus saw numerous<br />
publications on fishes <strong>of</strong> the Guiana Shield. Many<br />
were revisionary studies <strong>of</strong> genera or families whose<br />
ranges extend far beyond the limits <strong>of</strong> the Shield, <strong>of</strong>ten<br />
across major portions <strong>of</strong> cis-Andean South America<br />
and in some instances into trans-Andean regions or<br />
occasionally Central America. Other publications were<br />
restricted to the members <strong>of</strong> a genus, subfamily, or<br />
family from a country within the Shield region (e.g.,<br />
Suriname: Boeseman 1968, Nijssen 1970, Kullander &<br />
Nijssen 1989) or across a major portion <strong>of</strong> that area<br />
(e.g., Boeseman 1982). Relatively few <strong>of</strong> these papers<br />
involved broad surveys <strong>of</strong> an entire ichthy<strong>of</strong>auna in a<br />
river system or country on the Shield with those that
NUMBER 17 11<br />
did so summarized below arranged by the geographic<br />
subdivisions in the checklist. Many include associated<br />
ecological and life history information for fish communities<br />
and individual species.<br />
Brazil, Pará (PA). The single publication <strong>of</strong> note<br />
from this region is Ferreira (1993) that summarized the<br />
results <strong>of</strong> intensive collecting efforts at sites within the<br />
Rio Trombetas, one <strong>of</strong> the major northern tributaries<br />
<strong>of</strong> the Amazon River east <strong>of</strong> the Rio Negro.<br />
Brazil, Roraima (RO). Ferreira et al. (1988) summarized<br />
the ichthy<strong>of</strong>auna at several closely situated<br />
localities in the Rio Mucajai, a tributary <strong>of</strong> the Rio<br />
Branco. More recently, Ferreira et al. (2007) provided<br />
detailed information on the ichthy<strong>of</strong>auna across the<br />
expanse <strong>of</strong> the Rio Branco basin, supplemented by<br />
numerous color photographs, discussions <strong>of</strong> habitats,<br />
and comments on the anthropogenic impact on the<br />
aquatic systems within the basin.<br />
French Guiana (FG). French Guiana has the most<br />
intensely studied ichthy<strong>of</strong>auna <strong>of</strong> any portion <strong>of</strong> the<br />
Guyana Shield. The first attempt to summarize<br />
information on the fishes <strong>of</strong> the entire department<br />
was that <strong>of</strong> Puyo (1949). Géry (1972) followed up with<br />
studies <strong>of</strong> the characiforms (his characoids) from the<br />
Guianas with a particular focus on French Guiana.<br />
Planquette et al. (1996), Keith et al. (2000), and Le Bail<br />
et al. (2000), in a groundbreaking series <strong>of</strong> publications,<br />
brought together information on the spectrum <strong>of</strong><br />
the freshwater fish fauna in that department. Each<br />
species account includes a description, illustration, and<br />
comments on its biology. Distributions within and<br />
beyond French Guiana are discussed, and the sites <strong>of</strong><br />
occurrences <strong>of</strong> the species in the department are<br />
plotted.<br />
Guyana (GU). Notwithstanding the title <strong>of</strong> the<br />
publication, Eigenmann’s (1912) monographic study<br />
was based primarily on collections from the northern<br />
portions <strong>of</strong> Guyana, in particular the Potaro River and<br />
lower courses <strong>of</strong> the Essequibo and Demerara rivers,<br />
albeit with that data supplemented with information<br />
from the literature. Hardman et al. (2002) reported on<br />
the fish fauna captured at Eigenmann’s collecting<br />
localities nine decades after his expedition and provided<br />
a checklist <strong>of</strong> the 272 species collected in that survey.<br />
Lowe (McConnell) (1964) included lists <strong>of</strong> fishes from<br />
the southern most reaches <strong>of</strong> the Essequibo River<br />
system along with observations on their ecology and on<br />
the movements <strong>of</strong> various species during the yearly<br />
flood and drought cycles. Watkins et al. (2004)<br />
provided a summary <strong>of</strong> the fishes <strong>of</strong> the Iwokrama<br />
Forest Reserve.<br />
Suriname (SU). The ichthy<strong>of</strong>auna <strong>of</strong> Suriname<br />
remains relatively poorly documented, with the listing<br />
<strong>of</strong> the freshwater fishes in the country by Eigenmann<br />
(1912:64–73) largely derived from literature information.<br />
Boeseman (1952, 1953, 1954) supplemented<br />
Eigenmann’s listing. Ouboter & Mol (1993) presented<br />
the most comprehensive published list <strong>of</strong> the freshwater<br />
fishes <strong>of</strong> Suriname. Kullander & Nijssen (1989)<br />
published a detailed analysis <strong>of</strong> the cichlids <strong>of</strong><br />
Suriname.<br />
Venezuela, Amazonas (VA). The state <strong>of</strong> Amazonas,<br />
Venezuela, includes portions <strong>of</strong> the south-flowing Río<br />
Negro <strong>of</strong> the Amazon basin, the north-draining Río<br />
Orinoco and the entirety <strong>of</strong> the intervening Río<br />
Casiquiare. Mago-Leccia (1971) produced the first<br />
summary <strong>of</strong> the fishes <strong>of</strong> the Río Casiquiare. Lasso<br />
(1992), Royero et al. (1992), and Lasso et al. (2004a,<br />
2004b) provided information on the fish faunas <strong>of</strong><br />
various river systems within the state.<br />
Venezuela, Bolivar (BO). A series <strong>of</strong> studies treated<br />
the fish fauna <strong>of</strong> several right bank tributaries <strong>of</strong> the<br />
Río Orinoco that drain the northern slopes <strong>of</strong> the<br />
Guyana Shield. These included summaries <strong>of</strong> species in<br />
various basins, with those listings supplemented in<br />
some instances by information on fish biology and<br />
distribution. Significant publications on the ichthy<strong>of</strong>auna<br />
<strong>of</strong> the Río Caroni were published by Lasso<br />
(1991) and Lasso et al. (1991a, b) and for the Río<br />
Caura by Lasso et al. (2003a, b), Rodríquez-Olarte et<br />
al. (2003), and Vispo et al. (2003). The Río Cuyuni, a<br />
western tributary <strong>of</strong> the Essequibo River, drains the<br />
eastern portions <strong>of</strong> the Shield in the state <strong>of</strong> Bolivar.<br />
Machado-Allison et al. (2000) summarized the fish<br />
fauna <strong>of</strong> the Venezuelan portions <strong>of</strong> that river system.<br />
Lasso et al. (2004a) and Girardo et al. (2007) provide<br />
supplemental information on the ichthy<strong>of</strong>auna <strong>of</strong> that<br />
drainage basin.<br />
Ichthy<strong>of</strong>aunal Richness<br />
This checklist <strong>of</strong> fishes known from the water<br />
systems <strong>of</strong> the Guiana Shield includes 1168 species.<br />
Included in that total are representatives <strong>of</strong> 376 genera,<br />
49 families, and 15 orders. Five orders are dominant in<br />
terms <strong>of</strong> number <strong>of</strong> species living on the shield and<br />
account for 96.7% <strong>of</strong> the species (Characiformes, 478<br />
species and 41.0%; Siluriformes, 425 species and 36.4%;<br />
Perciformes, 126 species and 10.8%; Gymnotiformes,<br />
52 species and 4.5%; Cyprinodontiformes, 47 species<br />
and 4.0%). This sum <strong>of</strong> 1168 species attests to the<br />
dramatic improvement <strong>of</strong> our knowledge <strong>of</strong> the<br />
freshwater fish fauna on the Shield in slightly less than<br />
a century since Eigenmann (1912) documented fewer<br />
than 500 species from that region. The 1168 species are<br />
approximately 4.1% <strong>of</strong> the 28,400 fish species recently<br />
estimated to be present in all marine and freshwater<br />
systems worldwide (Nelson 2006), a percentage that<br />
amply testifies to the striking diversity <strong>of</strong> the ichthy<strong>of</strong>auna<br />
within that region. All the more noteworthy is<br />
the species-level richness <strong>of</strong> the ichthy<strong>of</strong>auna within the<br />
context <strong>of</strong> the overall Neotropical freshwater fish<br />
fauna. According to a recent summary, approximately<br />
5000 species <strong>of</strong> freshwater fishes occur across the
12 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
entirety <strong>of</strong> Central and South America (Reis et al.<br />
2003). Thus, the drainage systems <strong>of</strong> the Guiana Shield<br />
are home to approximately 23% <strong>of</strong> the freshwater fish<br />
species that occur across the vast expanse between<br />
southern South America and the southern border <strong>of</strong><br />
Mexico. Many factors contributed to the Shield region<br />
being a repository <strong>of</strong> freshwater ichthyological diversity,<br />
with a few particularly worthy <strong>of</strong> comment.<br />
Physiography<br />
The Guiana Shield is the ancient Precambrian<br />
Guianan formation resulting from the uplift <strong>of</strong> the<br />
underlying craton (Gibbs & Barron 1993) and demonstrates<br />
attributes that generally lead to high levels <strong>of</strong><br />
biodiversity: geological diversity, a topographically<br />
variable landscape, and transitions between ecosystems<br />
(Killeen et al. 2002). Overall the region has a primarily<br />
low to somewhat hilly physiography, albeit with some<br />
abrupt changes in topography in the regions proximate<br />
to the tepui formations that extend across much <strong>of</strong> the<br />
region in an approximately east to west alignment.<br />
Some river valleys have marked shifts in topography<br />
with resultant waterfalls, rapids, and riffles that<br />
increase the complexity <strong>of</strong> drainage system structure.<br />
These topographic factors result in multiple aquatic<br />
habitats with differing levels <strong>of</strong> physical complexity. At<br />
one extreme are the lentic waters <strong>of</strong> swamps, wetlands,<br />
channels, and lowland rivers. With increasing gradient,<br />
the drainage systems progress through variably flowing<br />
waters interrupted by higher energy settings such as<br />
riffles and isolated lower scale rapids. Finally, there are<br />
regions <strong>of</strong> greater gradients with rapidly flowing waters<br />
and major repetitive rapids and waterfalls. To the<br />
degree that differences in stream structure directly<br />
correlate with elevational gradients, there also occur<br />
differences in water temperatures.<br />
Water Chemistry<br />
Physical river system attributes are the most obvious<br />
manifestation <strong>of</strong> variation in aquatic systems across the<br />
Shield but represent a distinct subset <strong>of</strong> the spectrum <strong>of</strong><br />
factors that contribute to shifts in the composition and<br />
relative biomass <strong>of</strong> fish communities across that region.<br />
Complementing diversity in river structure are variations<br />
in water chemistry that occur not only between<br />
but also within drainage basins across the Shield. Three<br />
major water types occur in the tropics. The first <strong>of</strong><br />
these are white waters carrying nutrient-rich sediment<br />
loads for at least part <strong>of</strong> the year and which, despite<br />
their name, are actually brown (e.g., Rio Branco;<br />
Goulding et al. 2003:42). A second major group are<br />
clear water streams and rivers, including those draining<br />
many regions <strong>of</strong> the eastern portion <strong>of</strong> the Amazonian<br />
portion <strong>of</strong> the Shield (Pará and Amapá; Goulding et al.<br />
2003:43). Finally there are acidic black water rivers<br />
that drain heavily leached soils where decomposing<br />
plant matter produces high levels <strong>of</strong> fluvic and humic<br />
acids but with the water poor in dissolved solids and<br />
nutrients (e.g., Rio Negro; Goulding et al. 2003:44;<br />
Savanna Belt rivers in Suriname, Ouboter & Mol<br />
1993:134). Such water type differences occur both at<br />
the level <strong>of</strong> major river systems and at a much smaller<br />
scale within some river basins (Arbeláez et al. 2008).<br />
Admixtures <strong>of</strong> water types resulting from within basin<br />
water type differences yield conjoined drainages with<br />
variably intermediate chemistries (e.g., downriver <strong>of</strong><br />
where the black water Rio Negro empties into the<br />
white water Rio Solimões at Manaus, Goulding et al.<br />
2003:44; or where the white water Rupununi River<br />
empties into the black water upper Essequibo River,<br />
Watkins et al. 2005:40).<br />
Species <strong>of</strong> freshwater fishes demonstrate physiological,<br />
morphological, and behavioral adaptations that<br />
<strong>of</strong>ten allow them to specialize for life in particular<br />
water types but <strong>of</strong>ten simultaneously exclude them<br />
from other water types. Some species or genera are,<br />
therefore, more common in, or effectively limited to<br />
waters with particular chemical characteristics (Lowe-<br />
McConnell 1995). Exemplifying this situation are<br />
acidic black waters such as those in the Rio Negro<br />
that appear to be the primary, if not exclusive, habitat<br />
for some fish species (Goulding et al. 1988, table 2).<br />
These acidic waters are at the same time apparently<br />
inimical to other species and some genera notwithstanding<br />
the presence <strong>of</strong> these taxa in adjoining rivers<br />
<strong>of</strong> different water types (Goulding et al. 1988:98). The<br />
different water types also differ in degrees <strong>of</strong> primary<br />
productivity and dependence on detritus-based energy<br />
systems (De Jesús & Kohler 2004), factors that further<br />
impact fish diversity and community composition.<br />
Allochthonous Influences<br />
Terrestrial habitats further influence freshwater<br />
systems via the shift <strong>of</strong> nutrients and organic matter,<br />
with the <strong>of</strong>ten-substantial input into water bodies<br />
mediated by both water and wind. Terrestrial to<br />
freshwater inputs and their impacts on aquatic systems<br />
span a broad spectrum <strong>of</strong> scale. At one extreme, both<br />
continuing upland erosion and periodic floods transport<br />
dissolved and particulate matter into water bodies<br />
(Sioli 1975, Polis et al. 1997). Alternative forms <strong>of</strong><br />
riparian vegetation also affect the amount and types <strong>of</strong><br />
allochthonous materials, including detritus, seeds and<br />
fruits, and animals (primarily terrestrial insects) input<br />
into the aquatic food web via run<strong>of</strong>f and wind. The<br />
input <strong>of</strong> allochthonous detritus into Neotropical water<br />
systems supports particularly large populations <strong>of</strong><br />
detritivorous fishes (Flecker 1996). Most notably,<br />
many species and groups <strong>of</strong> fishes specialize on<br />
exploiting allochthonous seeds, fruits, and insects, with<br />
variation in the input <strong>of</strong> these items impacting the
NUMBER 17 13<br />
composition <strong>of</strong> ichthy<strong>of</strong>aunal communities (see Goulding<br />
et al. 1988, Boujard et al. 1990).<br />
At the other end <strong>of</strong> the scale spectrum, alternate soil<br />
types in conjunction with factors such as rainfall<br />
regimes and temperature also support dramatically<br />
different plant communities ranging from dense rainforests<br />
through open savannas. Physical attributes <strong>of</strong><br />
differing marginal plant communities directly influence<br />
fish community composition and structure in the water<br />
bodies that they border. Most obvious <strong>of</strong> these effects<br />
is differential shading by riparian forests, an influence<br />
that is particularly significant in terms <strong>of</strong> the species<br />
dwelling in streams and smaller rivers. Marginal<br />
vegetation and submerged macrophytes affect the<br />
physical complexity <strong>of</strong> water bodies and thus the<br />
composition <strong>of</strong> the resident ichthy<strong>of</strong>aunas in various<br />
fashions. Most noteworthy among these are differential<br />
inputs to the aquatic systems in the amount and type <strong>of</strong><br />
woody and leafy debris, variation in the submerged<br />
portions <strong>of</strong> overhanging terrestrial plants along water<br />
margins, and differing amounts and types <strong>of</strong> submerged<br />
emergent vegetation. Synergy <strong>of</strong> drainage<br />
structure and energy, differences in water types,<br />
variation in associated riparian animals and plants,<br />
and differences in input <strong>of</strong> nutrients yield dramatically<br />
different fish communities. That variation at the local<br />
level is a major contributor to the overall species-level<br />
richness <strong>of</strong> the ichthy<strong>of</strong>auna across the totality <strong>of</strong> the<br />
Shield.<br />
Drainage System Interconnections<br />
Physiological and physical factors closely tie freshwater<br />
fishes to drainage patterns. Historical separations<br />
<strong>of</strong>, and associations between, drainage systems<br />
thereby contributed to the present day distributions <strong>of</strong><br />
many species on the Shield and the richness <strong>of</strong> that<br />
fauna. Notwithstanding the tectonic quiescence <strong>of</strong> the<br />
Guiana Shield for over 550 million years, the highlands<br />
resulting from the uplift <strong>of</strong> the craton underwent<br />
progressive erosion <strong>of</strong> the sedimentary layers overlying<br />
that base (Gibbs & Barron 1993). The pronounced<br />
degree <strong>of</strong> endemicity in many <strong>of</strong> the basins across the<br />
Shield is indicative <strong>of</strong> their long isolation, with factors<br />
including differences in water types being influential in<br />
this regard. Nonetheless, there have been changes in<br />
the water flow patterns on and along the margins <strong>of</strong> the<br />
Shield that influenced the present composition <strong>of</strong> the<br />
ichthy<strong>of</strong>auna in those systems.<br />
At the large scale, tectonic events resulted in the<br />
broader details <strong>of</strong> the present Orinoco and Amazon<br />
basins, both <strong>of</strong> which contribute to the Shield<br />
ichthy<strong>of</strong>auna. A large paleo-drainage encompassing<br />
much <strong>of</strong> the present Orinoco and Amazon basin<br />
drained north into the Caribbean Sea approximately<br />
at the present location <strong>of</strong> Lago Maracaibo. Tectonic<br />
events at the end <strong>of</strong> the Miocene resulted in separation<br />
<strong>of</strong> the Amazon from Orinoco basin (Hoorn 1994).<br />
Another consequence <strong>of</strong> these changes was the shift<br />
eastward <strong>of</strong> the mainstream Orinoco along the<br />
northern boundary <strong>of</strong> the Shield to its present mouth<br />
slightly north <strong>of</strong> the northeastern margin <strong>of</strong> the Shield.<br />
This dramatic realignment led to its capture <strong>of</strong><br />
drainages flowing from the northern slopes <strong>of</strong> the<br />
Shield. The shift <strong>of</strong> the mainstream Amazon to it<br />
present mouth similarly resulted in the capture <strong>of</strong> the<br />
southern draining rivers <strong>of</strong> the Shield. Disrupting the<br />
continuity between many <strong>of</strong> those freshwater systems<br />
for varying periods were subsequent marine transgressions<br />
into the eastern portions <strong>of</strong> the Amazon valley.<br />
Superimposed on these large-scale drainage pattern<br />
changes were long-term, <strong>of</strong>ten pronounced, climate<br />
changes through the region (Baker et al. 2001) with<br />
resultant sequential contraction and expansion <strong>of</strong><br />
suitable aquatic habitats. This combination <strong>of</strong> hydrographic<br />
and climatic changes resulted in disruptions, in<br />
some instances repeated disruptions, <strong>of</strong> previously<br />
continuous species ranges, thereby setting the stage<br />
for subsequent speciation.<br />
The complex hydrological history <strong>of</strong> the drainages<br />
on the Shield involved not only division <strong>of</strong> previously<br />
continuous drainages but also in new connections<br />
between various river systems. Connectivity resulted<br />
from landscape tilting during uplift events and via<br />
headwater stream capture. Those events permitted, and<br />
continue to permit, movements <strong>of</strong> fishes between what<br />
have been isolated basins on the Shield. Subsequent<br />
disruptions <strong>of</strong> connectivity provided an opportunity<br />
for the evolution <strong>of</strong> species. Such past connections may<br />
account for unusual present day distribution patterns<br />
previously highlighted by some authors (e.g., Armbruster<br />
2005). Prominent among these possible connections<br />
was the river hypothesized to have drained directly<br />
from the south slope <strong>of</strong> the Guiana Shield into the<br />
Atlantic Ocean through what is now northeastern<br />
Guyana (Hammond 2005:137). Faunal similarities and<br />
close phylogenetic relationships among included fish<br />
species also point to possible past associations between<br />
the upper Río Caroni <strong>of</strong> the Río Orinoco basin and the<br />
Río Cuyuni, a western tributary <strong>of</strong> the Essequibo River<br />
(Lasso et al. 1991a, Sabaj Pérez & Birindelli 2008). An<br />
interconnection between the Amazon and the upper<br />
portions <strong>of</strong> the Maroni River was proposed by<br />
Cardoso & Montoya-Burgos (2009) who also proposed<br />
that temporary connections between adjoining river<br />
systems during periods <strong>of</strong> lower sea levels permitted<br />
dispersal <strong>of</strong> freshwater fishes along the coasts <strong>of</strong> the<br />
Guianas.<br />
The paramount example <strong>of</strong> an extant interconnection<br />
between major river systems on the Shield, or<br />
indeed across the continent, is the Rio Casiquiare. This<br />
over 300 km long natural canal connects the Río Negro<br />
<strong>of</strong> the Amazon River basin with the upper portions <strong>of</strong><br />
the Río Orinoco. This unusual drainage begins as a
14 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
bifurcation <strong>of</strong> the upper portion <strong>of</strong> the Río Orinoco<br />
and represents an ongoing capture <strong>of</strong> the upper portion<br />
<strong>of</strong> the latter river system by the Rio Negro (Sternberg<br />
1975, Winemiller at al. 2008). Despite its substantial<br />
size at the divergence, where it is approximately 100 m<br />
wide, and the fact that it carries a significant portion <strong>of</strong><br />
the total flow <strong>of</strong> that portion <strong>of</strong> the Río Orinoco, the<br />
Río Casiquiare does not provide unimpeded transit for<br />
all species <strong>of</strong> fishes between those basins. Lack <strong>of</strong><br />
interbasin species panmixis is, in part, a function <strong>of</strong> the<br />
fact that the Río Casiquiare conjoins headwaters<br />
habitats inhabited by a subset <strong>of</strong> the species resident<br />
across the entirety <strong>of</strong> each basin. Equally, or perhaps<br />
more significantly, the gradient in water types along<br />
the course <strong>of</strong> the Río Casiquiare acts as a partial filter<br />
that impedes movement <strong>of</strong> many fish species (Winemiller<br />
et al. 2008), thereby maintaining differences in<br />
ichthy<strong>of</strong>aunal composition between the upper Río<br />
Negro and upper Río Orinoco.<br />
Although the Río Casiquiare is the most notable<br />
connection between major rivers on the Guiana Shield<br />
and the only year-round continuity, some degree <strong>of</strong><br />
seasonal connectivity occurs between the upper reaches<br />
<strong>of</strong> the Rio Branco and the southern most portions <strong>of</strong><br />
the Essequibo River in the Rupununi Savannas <strong>of</strong><br />
southwestern Guyana. That region is an expansive<br />
floodplain where the headwaters <strong>of</strong> the Rupununi<br />
River (Essequibo basin) and the Takutu and Ireng<br />
rivers (both components <strong>of</strong> the Rio Branco <strong>of</strong> the<br />
Amazon River basin) come into close proximity. This<br />
proximity and varying degree <strong>of</strong> continuity <strong>of</strong> the<br />
headwaters <strong>of</strong> these rivers across a vast flooded plan<br />
during high water periods [Lowe (McConnell) 1964,<br />
Watkins et al. 2005] facilitate movement <strong>of</strong> at least<br />
some fish species between the headwaters <strong>of</strong> the Rio<br />
Branco and Essequibo River basins. Although such<br />
movements potentially enrich the ichthy<strong>of</strong>aunas <strong>of</strong><br />
each <strong>of</strong> those river systems, they do not add to the<br />
overall richness <strong>of</strong> the fish fauna <strong>of</strong> the shield.<br />
Future Directions<br />
Notwithstanding the series <strong>of</strong> papers listed under<br />
‘‘State <strong>of</strong> Knowledge <strong>of</strong> the Shield Fish Fauna’’ and<br />
many other publications, the continuing discovery and<br />
description <strong>of</strong> freshwater species from water systems on<br />
the Shield testifies to the incomplete state <strong>of</strong> our<br />
knowledge <strong>of</strong> that fish fauna. The primary impediment<br />
is the lack <strong>of</strong> exhaustive ichthyological collecting across<br />
that vast region, with many river systems still<br />
effectively unsampled (e.g., the upper Mazaruni River<br />
in Guyana; Taphorn et al. 2008). Headwater tributary<br />
streams, deep mainstream channels, and difficult-tosample<br />
habitats such as swamps and rapids remain<br />
unsampled or poorly sampled even in those drainage<br />
systems that have been the subject <strong>of</strong> ichthyological<br />
collecting efforts. Those habitats and some largely<br />
ichthyologically unexplored drainage systems hold the<br />
greatest promise as sources <strong>of</strong> undescribed species;<br />
however, we must not lose sight <strong>of</strong> the fact that areas<br />
that have been long the foci <strong>of</strong> ichthyological sampling<br />
continue to yield new species and are deserving <strong>of</strong><br />
continued attention. Recent monographic studies <strong>of</strong><br />
speciose genera and families <strong>of</strong> Neotropical freshwater<br />
fishes demonstrate that collections in museums, universities,<br />
and research institutions <strong>of</strong> North and South<br />
America and Europe house numerous species as <strong>of</strong> yet<br />
unknown to science. Indeed in many groups, the vast<br />
majority, if not all, <strong>of</strong> recently described species were<br />
already represented in collections and awaited discovery,<br />
<strong>of</strong>ten for many decades.<br />
Thorough sampling <strong>of</strong> the freshwater fish fauna is<br />
vital as is the thorough examination <strong>of</strong> materials in<br />
collections, but equally or perhaps more important for<br />
furthering our knowledge <strong>of</strong> the fishes on the Shield are<br />
comprehensive revisionary studies <strong>of</strong> all groups <strong>of</strong><br />
fishes represented in that ichthy<strong>of</strong>auna. Such in-depth<br />
studies are critical given our inadequate understanding<br />
<strong>of</strong> the species-level diversity <strong>of</strong> many Neotropical<br />
freshwater groups (Vari & Malabarba 1998). Inclusive<br />
revisions <strong>of</strong> complex groups <strong>of</strong> Neotropical freshwater<br />
fishes have repeatedly demonstrated that the sum <strong>of</strong><br />
long recognized species within a genus typically<br />
underestimates the actual number <strong>of</strong> species in a taxon,<br />
sometimes to a pronounced degree. An example is the<br />
67 species now recognized in Creagrutus; a total three<br />
and one-half times the number <strong>of</strong> species (19)<br />
recognized in the genus prior to 1994 (Harold & Vari<br />
1994, Vari & Harold 2001, Vari & Lima 2003, Ribeiro<br />
et al. 2004, Torres-Mejia & Vari 2005). Comprehensive<br />
revisions similarly further the subsequent identification<br />
<strong>of</strong> additional previously unrecognized species by other<br />
researchers (Reis 2004).<br />
It is impossible to estimate the degree to which the<br />
total number <strong>of</strong> freshwater fish species summarized in<br />
this checklist falls short <strong>of</strong> the actual count <strong>of</strong> species<br />
dwelling on the Shield. Nonetheless, it is clear that the<br />
rate <strong>of</strong> continued additions to this speciose fish fauna,<br />
the many regions and habitats that have not yet been<br />
thoroughly explored ichthyologically, and the large<br />
numbers <strong>of</strong> groups <strong>of</strong> fishes in the region that have not<br />
yet been exhaustively studied portend a significant<br />
increase in the species total.<br />
Conservation Challenges<br />
Deleterious anthropogenic activities impact freshwater<br />
ichthy<strong>of</strong>aunas across the Neotropics (Killeen 2007),<br />
with many affecting various portions <strong>of</strong> the Guiana<br />
Shield and adversely influencing fish communities in<br />
the region. Adverse impacts are pervasive across<br />
freshwater ichthy<strong>of</strong>aunas worldwide (Millennium Ecosystem<br />
Assessment 2005, Revenga et al. 2005), with<br />
freshwater fishes consequently the most threatened
NUMBER 17 15<br />
groups <strong>of</strong> vertebrates across the world in terms <strong>of</strong><br />
affected species (Dudgeon et al. 2006, Chapman et al.<br />
2008). Major impacts on freshwater ichthy<strong>of</strong>aunas <strong>of</strong><br />
the Guiana Shield cover the spectrum <strong>of</strong> human<br />
activities. These include overfishing for human consumption<br />
and the aquarium trade, pollution from<br />
agricultural, domestic, industrial and mining sources,<br />
diversion <strong>of</strong> water for agricultural, domestic, and<br />
industrial purposes, mining within river channels,<br />
introductions <strong>of</strong> exotic species, transplanting <strong>of</strong> native<br />
species between separate drainage systems, deforestation<br />
within drainage basins with consequent changes in<br />
water flow patterns and quality, increased erosion and<br />
siltation as a consequence <strong>of</strong> development, agriculture,<br />
and mining operations, and impoundments for hydroelectric<br />
and irrigation systems with disruption <strong>of</strong><br />
migration routes for fishes.<br />
Major advances are necessary before we approach a<br />
definitive understanding <strong>of</strong> the species-level diversity<br />
for the fishes inhabiting the rivers, streams, lakes, and<br />
other water bodies on the Guiana Shield. Nonetheless,<br />
the following Checklist can serve as a foundation for<br />
future studies, leading to a better appreciation <strong>of</strong> the<br />
diversity <strong>of</strong> that ichthy<strong>of</strong>auna. Such information is<br />
vital to inform decisions by resource managers,<br />
government agencies, and members <strong>of</strong> the public<br />
interested in protecting both the fish fauna and the<br />
broader aquatic communities, both <strong>of</strong> which provide<br />
essential and important ecosystem services across the<br />
Shield.<br />
Species <strong>of</strong> the Guiana Shield<br />
The Checklist includes species recognized at the time<br />
that contributors completed their accounts (mid-2008)<br />
with these supplemented whenever possible by information<br />
on new species described from the Shield<br />
through early 2009. Readers interested in further<br />
information on the families and species included in<br />
the listing can refer to CLOFFSCA (Reis et al. 2003).<br />
That listing includes bibliographic information for all<br />
fish species in Central and South America, including<br />
those known to occur on the Guiana Shield, through<br />
the end <strong>of</strong> 2002. References to the original descriptions<br />
<strong>of</strong> species published post that date are listed under the<br />
following Guide to the Checklist. The regions utilized<br />
in the checklist correspond to those used for terrestrial<br />
vertebrates in Hollowell & Reynolds (2005). Abell et al.<br />
(2008) recently proposed a hydrographically delimited<br />
series <strong>of</strong> zones for South America. The more finescaled<br />
resolution <strong>of</strong> that system is potentially more<br />
informative in terms <strong>of</strong> areas <strong>of</strong> regional endemism for<br />
aquatic organisms. We defer, however, from applying<br />
it to the freshwater fishes <strong>of</strong> the Guiana Shield given<br />
the large degree <strong>of</strong> uncertainty as to distributional<br />
limits for most species in that fish fauna.<br />
Acknowledgments<br />
Completion <strong>of</strong> this Bulletin was facilitated by S.<br />
Raredon who prepared the various images that face<br />
each section. Numerous individuals assisted our field<br />
efforts and studies <strong>of</strong> the fishes on the Shield over the<br />
years, with particular thanks to L. Aguana, A.<br />
Machado-Allison, O. Castillo, J. Fernandez, S. Jewett,<br />
C. Lasso, H. Madarie, F. Mago-Leccia, L. Parenti, F.<br />
Provenzano, and D. Taphorn.<br />
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The cornerstone <strong>of</strong> this project was the exhaustive<br />
Checklist <strong>of</strong> the freshwater fishes <strong>of</strong> South and Central<br />
America (abbreviated hereafter as CLOFFSCA; Reis et<br />
al. 2003). Only those species with confirmed collections<br />
within the geographic bounds <strong>of</strong> the Guiana Shield<br />
(after Gibbs & Barron 1993; excluding outlier formations<br />
in Colombia) were extracted from the<br />
CLOFFSCA accounts. Records from the main channel<br />
<strong>of</strong> the Amazon River that lies at a distance from Shield<br />
boundaries were excluded, but those from the main<br />
channel <strong>of</strong> the Río Orinoco that runs closer to the<br />
northern limit <strong>of</strong> the Shield were included. This<br />
preliminary assemblage was updated from the literature<br />
published since CLOFFSCA’s release and the<br />
resultant summaries sent to relevant authorities for<br />
review. Whenever possible, the families and/or subfamilies<br />
were reviewed by the original CLOFFSCA<br />
authors. References include papers describing new<br />
species from the area <strong>of</strong> interest that appeared after<br />
the completion <strong>of</strong> CLOFFSCA.<br />
Classification in the checklist follows CLOFFSCA.<br />
Orders and families are listed in systematic order with<br />
genera and species listed alphabetically within families/<br />
subfamilies and genera, respectively. Subfamilies are<br />
limited to the Characidae (Characiformes) and Loricariidae<br />
(Siluriformes). Synonyms and common names<br />
were not included, but that information is available in<br />
CLOFFSCA.<br />
The Guiana Shield was divided into 11 regions (see<br />
Table 3 for abbreviations and Fig. 1 for map) to<br />
illustrate distributions. The four regions in Brazil<br />
include only those parts <strong>of</strong> each state (Amapá,<br />
Amazonas, Pará, Roraima) falling within the Shield’s<br />
boundaries. Species from rivers forming borders<br />
between two regions were listed as occurring in both<br />
regions. Records from river systems not totally<br />
contained within the Guiana Shield were only included<br />
if it could be confirmed that a particular species was<br />
collected in the portion <strong>of</strong> the drainage basin overlying<br />
Table 3.—Regions <strong>of</strong> the Guiana Shield used in checklist in<br />
approximately west to east order.<br />
CG Colombian Guayana<br />
VA Venezuela—Amazonas<br />
BO Venezuela—Bolívar<br />
DA Venezuela—Delta Amacuro<br />
BA Brazil—Amazonas<br />
RO Brazil—Roraima<br />
PA Brazil—Pará<br />
AP Brazil—Amapá<br />
GU Guyana<br />
SU Suriname<br />
FG French Guiana<br />
GUIDE TO THE CHECKLIST<br />
ALEKSANDAR RADOSAVLJEVIC<br />
Shield. Regional abbreviations in the checklist are<br />
followed by a ‘‘?’’ in instances when distributions were<br />
uncertain or questionable.<br />
Literature Cited<br />
(Including papers describing new species published after completion<br />
<strong>of</strong> CLOFFSCA.)<br />
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———. 2004. Pseudancistrus sidereus, a new species from southern<br />
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Pseudancistrus.—Zootaxa 628:1–15.<br />
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from the Río Caroní, Venezuela (Siluriformes: Loricariidae).—Zootaxa<br />
1731:33–41.<br />
———, & D. C. Werneke. 2005. Peckoltia cavatica, a new loricariid<br />
catfish from Guyana and a redescription <strong>of</strong> P. braueri<br />
(Eigenmann 1912) (Siluriformes).—Zootaxa 882:1–14.<br />
———, N. K. Lujan, & D. C. Taphorn. 2007. Four new Hypancistrus<br />
(Siluriformes: Loricariidae) from Amazonas, Venezuela.—<br />
Copeia 2007(1):62–79.<br />
———, L. A. Tansey, & N. K. Lujan. 2007. Hypostomus rhantos<br />
(Siluriformes: Loricariidae), a new species from southern<br />
Venezuela.—Zootaxa 1553:59–68.<br />
Benine, R. C., & G. A. M. Lopes. 2007. A new species <strong>of</strong><br />
Hemigrammus Gill, 1858 (Characiformes: Characidae) from<br />
Río Caura, Venezuela.—Zootaxa 1610:53–59.<br />
———, G. Z. Pelição, & R. P. Vari. 2004. Tetragonopterus<br />
lemniscatus (Characiformes: Characidae), a new species from<br />
the Corantijn River basin in Suriname.—Proceedings <strong>of</strong> the<br />
<strong>Biological</strong> <strong>Society</strong> <strong>of</strong> <strong>Washington</strong> 117:339–345.<br />
Birindelli, J. L. O., M. H. Sabaj, & D. C. Taphorn. 2007. New species<br />
<strong>of</strong> Rhynchodoras from the Río Orinoco, Venezuela, with<br />
comments on the genus (Siluriformes: Doradidae).—Copeia<br />
2007(3):672–684.<br />
Bührnheim, C. M., & L. R. Malabarba. 2007. Redescription <strong>of</strong><br />
Odontostilbe pulchra (Gill, 1858) (Teleostei: Characidae:<br />
Cheirodontinae), and description <strong>of</strong> two new species from<br />
the río Orinoco basin.—Neotropical Ichthyology 5(1):1–20.<br />
Buitrago-Suárez, U. A., & B. M. Burr. 2007. Taxonomy <strong>of</strong> the catfish<br />
genus Pseudoplatystoma Bleeker (Siluriformes: Pimelodidae)<br />
with recognition <strong>of</strong> eight species.—Zootaxa 1512:1–38.<br />
Chern<strong>of</strong>f, B., & A. Machado-Allison. 2005. Bryconops magoi and<br />
Bryconops collettei (Characiformes: Characidae), two new<br />
freshwater fish species from Venezuela, with comments on B.<br />
caudomaculatus (Günther).—Zootaxa 1094:1–23.<br />
de Chambrier, S., & J. I. Montoya-Burgos. 2008. Pseudancistrus<br />
corantijniensis, a new species from the Guyana Shield
22 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
(Siluriformes: Loricariidae) with a molecular and morphological<br />
description <strong>of</strong> the Pseudancistrus barbatus group.—<br />
Zootaxa 1918:45–58.<br />
Deynat, P. 2006. Potamotrygon marinae n. sp., a new species <strong>of</strong><br />
freshwater stingrays from French Guiana (Myliobatiformes:<br />
Potamotrygonidae).—Comptes Rendus Biologies 329(7):483–<br />
493.<br />
DoNascimiento, C., & J. G. Lundberg. 2005. Myoglanis aspredinoides<br />
(Siluriformes: Heptapteridae), a new catfish from the<br />
Río Ventuari, Venezuela.—Zootaxa 1009:37–49.<br />
Friel, J. P. 2008. Pseudobunocephalus, a new genus <strong>of</strong> banjo catfish<br />
with the description <strong>of</strong> a new species from the Orinoco River<br />
system <strong>of</strong> Colombia and Venezuela (Siluriformes: Aspredinidae).—Neotropical<br />
Ichthyology 6(3):293–300.<br />
Garutti, V. 2003. Revalidação de Astyanax rupununi Fowler, 1914<br />
(Teleostei, Characidae) e descrição de duas espécies novas para<br />
ogênero.—Papéis Avulsos de Zoologia (São Paulo) 43(1):1–9.<br />
Gibbs, A. K., & C. N. Barron. 1993. The geology <strong>of</strong> the Guiana<br />
Shield. Oxford University Press, New York, 246 pp.<br />
Hrbek, T., D. C. Taphorn, & J. E. Thomerson. 2005. Molecular<br />
phylogeny <strong>of</strong> Austr<strong>of</strong>undulus Myers (Cyprinodontiformes:<br />
Rivulidae), with revision <strong>of</strong> the genus and the description <strong>of</strong><br />
four new species.—Zootaxa 825:1–39.<br />
Jégu, M., P. Keith, & P.-Y. Le Bail. 2003. Myloplus planquettei sp. n.<br />
(Teleostei: Characidae) une nouvelle espèce de grand Serrasalminae<br />
phytophage du bouclier guyanais.—Revue Suisse de<br />
Zoologie 1024:833–853.<br />
Keith, P., L. Nandrin, & P.-Y. Le Bail. 2006. Rivulus gaucheri, a new<br />
species <strong>of</strong> rivuline (Cyprinodontiformes: Rivulidae) from<br />
French Guiana.—Cybium 30(2):133–137.<br />
Kullander, S. O., & E. J. G. Ferreira. 2005. Two new species <strong>of</strong><br />
Apistogramma Regan (Teleostei: Cichlidae) from the rio<br />
Trombetas, Pará State, Brazil.—Neotropical Ichthyology<br />
3(3):361–371.<br />
———, & ———. 2006. A review <strong>of</strong> the South American cichlid<br />
genus Cichla, with descriptions <strong>of</strong> nine new species (Teleostei:<br />
Cichlidae).—Ichthyological Exploration <strong>of</strong> Freshwaters 17(4):<br />
289–398.<br />
Lasso, C. A., & F. Provenzano. 2002. Dos nuevas especies de bagres<br />
del género Trichomycterus (Siluriformes: Trichomycteridae)<br />
de la Gran Sabana, Escudo de las Guayanas, Venezuela.—<br />
Revista de Biología Tropical 50(3/4):1139–1149.<br />
———, J. I. Mojica, J. S. Usma, J. A. Maldonaldo O., C.<br />
DoNascimiento, D. C. Taphorn, F. Provenzano, O. M. Lasso<br />
Alcalá, G. Galvis, L. Vásquez, M. Lugo, A. Machado Allison,<br />
R. Royero, C. Suárez, & A. Ortega Lara. 2004. Peces de las<br />
cuenca del río Orinoco. Parte I: lista de especies y distribución<br />
por subcuencas.—Biota Colombiana 5:95–157.<br />
Lasso-Alcalá, O. M., D. C. Taphorn B., C. A. Lasso, & O. León-<br />
Mata. 2006. Rivulus sape, a new species <strong>of</strong> killifish (Cyprinodontiformes:<br />
Rivulidae) from the Paragua River system,<br />
Caroní River drainage, Guyana Shield, Venezuela.—Zootaxa<br />
1275:21–29.<br />
Lehman, A. P., & R. E. Reis. 2004. Callichthys serralabium: a new<br />
species <strong>of</strong> neotropical catfish from the upper Orinoco and<br />
Negro Rivers (Siluriformes: Callichthyidae).—Copeia 2004(2):<br />
336–343.<br />
López-Fernández, H., & D. C. Taphorn. 2004. Geophagus abalios, G.<br />
dicrozoster and G. winemilleri (Perciformes: Cichlidae), three<br />
new species from Venezuela.—Zootaxa 439:1–27.<br />
———, D. C. Taphorn Baechle, & S. O. Kullander. 2006. Two new<br />
species <strong>of</strong> Guianacara from the Guiana Shield <strong>of</strong> eastern<br />
Venezuela (Perciformes: Cichlidae).—Copeia 2006(3):384–<br />
395.<br />
Lujan, N. K. 2008. Description <strong>of</strong> a new Lithoxus (Siluriformes:<br />
Loricariidae) from the Guayana Highlands with a discussion<br />
<strong>of</strong> Guiana Shield biogeography.—Neotropical Ichthyology<br />
6(3):413–418.<br />
———, M. Arce, & J. W. Armbruster. 2009. A new black<br />
Baryancistrus with blue sheen from the upper Orinoco<br />
(Siluriformes: Loricariidae).—Copeia 2009(1):50–56.<br />
———, J. W. Armbruster, & M. H. Sabaj. 2007. Two new species <strong>of</strong><br />
Pseudancistrus from southern Venezuela (Siluriformes: Loricariidae).—Ichthyological<br />
Exploration <strong>of</strong> Freshwaters, 18(2):<br />
163–174.<br />
Lundberg, J. G., & W. M. Dahdul. 2008. Two new cis-Andean<br />
species <strong>of</strong> the South American catfish genus Megalonema<br />
allied to trans-Andean Megalonema xanthum, with description<br />
<strong>of</strong> a new subgenus (Siluriformes: Pimelodidae).—Neotropical<br />
Ichthyology 6(3):439–454.<br />
Malabarba, M. C. S. L. 2004. Revision <strong>of</strong> the Neotropical genus<br />
Triportheus Cope, 1872 (Characiformes: Characidae).—Neotropical<br />
Ichthyology 2(4):167–204.<br />
Montaña, C. G., H. López-Fernández, & D. C. Taphorn. 2008. A<br />
new species <strong>of</strong> Crenicichla (Perciformes: Cichlidae) from the<br />
Ventuari River, Upper Orinoco River Basin, Amazonas State,<br />
Venezuela.—Zootaxa 1856:33–40.<br />
de Pinna, M., & P. Keith. 2003. A new species <strong>of</strong> the catfish genus<br />
Ituglanis from French Guyana (Osteichthyes: Siluriformes:<br />
Trichomycteridae).—Proceedings <strong>of</strong> the <strong>Biological</strong> <strong>Society</strong> <strong>of</strong><br />
<strong>Washington</strong> 116:873–882.<br />
Provenzano, R. F., A. Machado-Allison, B. Chern<strong>of</strong>f, P. Willink, &<br />
P. Petry. 2005. Harttia merevari, a new species <strong>of</strong> catfish<br />
(Siluriformes: Loricariidae) from Venezuela.—Neotropical<br />
Ichthyology 3(4):519–524.<br />
Radda, A. C. 2004. Description <strong>of</strong> a new species <strong>of</strong> the rivuline genus<br />
Rivulus Poey, 1869 (Rivulidae, Osteichthyes) from Rio Caura,<br />
Bolivar State, Venezuela.—Annalen des Naturhistorischen<br />
Museums in Wien, Serie B, 105B:21–25.<br />
Ramos, R. T. C. 2003. Systematic review <strong>of</strong> Apionichthys (Pleuronectiformes:<br />
Achiridae), with description <strong>of</strong> four new species.—<br />
Ichthyological Exploration <strong>of</strong> Freshwaters 14(2):97–126.<br />
Reis, R. E., S. O. Kullander, & C. J. Ferraris, Jr. (eds.). 2003. Check<br />
list <strong>of</strong> the freshwater fishes <strong>of</strong> South and Central America.<br />
Edipucrs, Porto Alegre, Brazil, 729 pp.<br />
Román-Valencia, C., D. C. Taphorn B., & R. I. Ruiz-C. 2008. Two<br />
new Bryconamericus: B. cinarucoense n. sp. and B. singularis n.<br />
sp. (Characiformes, Characidae) from the Cinaruco River,<br />
Orinoco Basin, with keys to all Venezuelan species.—Animal<br />
Biodiversity and Conservation 31(1):15–27.<br />
Römer, U., I. Hahn, & A. Conrad. 2006. Apistogramma wapisana<br />
sp. n. Description <strong>of</strong> a dwarf cichlid from northern Brazil.<br />
Cichlid Atlas 2. Mergus Verlag, Melle, Germany, pp. 748–<br />
763.<br />
Rosa, R. S., M. R. de Carvalho, & C. de Almeida Wanderley. 2008.<br />
Potamotrygon boesemani (Chondrichthyes: Myliobatiformes:<br />
Potamotrygonidae), a new species <strong>of</strong> Neotropical freshwater<br />
stingray from Surinam.—Neotropical Ichthyology 6(1):1–8.<br />
Sabaj, M. H. 2005. Taxonomic assessment <strong>of</strong> Leptodoras (Siluriformes:<br />
Doradidae) with descriptions <strong>of</strong> three new species.—<br />
Neotropical Ichthyology 3(4):637–678.<br />
———, D. C. Taphorn, & O. E. Castillo G. 2008. Two new species <strong>of</strong><br />
thicklip thornycats, genus Rhinodoras (Teleostei: Siluriformes:<br />
Doradidae).—Copeia 2008(1):209–226.<br />
Sabaj Pérez, M. H., & J. L. O. Birindelli. 2008. Taxonomic revision <strong>of</strong><br />
extant Doras Lacepède, 1803 (Siluriformes: Doradidae) with<br />
descriptions <strong>of</strong> three new species.—Proceedings <strong>of</strong> the<br />
Academy <strong>of</strong> Natural Sciences <strong>of</strong> Philadelphia 157:189–233.<br />
de Santana, C. D., & D. C. Taphorn. 2006. Sternarchorhynchus<br />
gnomus, a new species <strong>of</strong> electric knifefish from the Lower Río<br />
Caroni, Venezuela (Gymnotiformes: Apteronotidae).—Ichthyological<br />
Exploration <strong>of</strong> Freshwaters 17(1):1–8.<br />
———, & R. P. Vari. 2009. The South American electric fish genus<br />
Platyurosternarchus (Gymnotiformes: Apteronotidae).—Copeia<br />
2009(2):233–244.
NUMBER 17 23<br />
Sarmento-Soares, L. M., & R. F. Martins-Pinheiro. 2008. A<br />
systematic revision <strong>of</strong> Tatia (Siluriformes: Auchenipteridae:<br />
Centromochlinae).—Neotropical Ichthyology 6(3):495–542.<br />
Schaefer, S. A., & F. Provenzano, 2008. The Lithogeninae<br />
(Siluriformes, Loricariidae): anatomy, interrelationships, and<br />
description <strong>of</strong> a new species.—American Museum Novitates<br />
3637:1–49.<br />
———, ———, M. de Pinna, & J. N. Baskin. 2005. New and<br />
noteworthy Venezuelan glanapterygine catfishes (Siluriformes,<br />
Trichomycteridae), with discussion <strong>of</strong> their biogeography and<br />
psammophily.—American Museum Novitates 3496:1–27.<br />
Schindler, I., & W. Staeck. 2006. Geophagus gottwaldi sp. n. – a new<br />
species <strong>of</strong> cichlid fish (Teleostei: Perciformes: Cichlidae) from<br />
the drainage <strong>of</strong> the upper río Orinoco in Venezuela.—<br />
Zoologische Abhandlungen (Dresden) 56:91–97.<br />
———, & ———. 2008. Dicrossus gladicauda sp. n. – a new species <strong>of</strong><br />
crenicarine dwarf cichlids (Teleostei: Perciformes: Cichlidae)<br />
from Colombia, South-America.—Vertebrate Zoology 58(1):<br />
67–73.<br />
Sidlauskas, B. L., & G. M. dos Santos. 2005. Pseudanos winterbottomi:<br />
a new anostomine species (Teleostei: Characiformes:<br />
Anostomidae) from Venezuela and Brazil, and comments on<br />
its phylogenetic relationships.—Copeia 2005(1):109–123.<br />
———, J. C. Garavello, & J. Jellen. 2007. A new Schizodon<br />
(Characiformes: Anostomidae) from the Río Orinoco system,<br />
with a redescription <strong>of</strong> S. Isognathus from the Río Paraguay<br />
system.—Copeia 2007(3):711–725.<br />
Staeck, W. 2003. Cichliden-Lexikon, Teil 3: Südamerikanische<br />
Zwergbuntbarsche. Dähne Verlag, Ettlingen, Germany, 219 pp.<br />
———, & I. Schindler. 2007. Description <strong>of</strong> Laetacara fulvipinnis sp.<br />
n. (Teleostei: Perciformes: Cichlidae) from the upper drainages<br />
<strong>of</strong> the rio Orinoco and rio Negro in Venezuela.—<br />
Vertebrate Zoology 57(1):63–71.<br />
Suijker, W. H., & G. E. Collier. 2006. Rivulus mahdiaensis, a new<br />
killifish from central Guyana (Cyprinodontiformes: Rivulidae).—Zootaxa<br />
1246:1–13.<br />
Taphorn, B. D. C., H. López-Fernández, & C. R. Bernard. 2008.<br />
Apareiodon agmatos, a new species from the upper Mazaruni<br />
river, Guyana (Teleostei: Characiformes: Parodontidae).—<br />
Zootaxa 1925:31–38.<br />
Taphorn, B., C. G. Montaña, & P. Buckup. 2006. Characidium<br />
longum (Characiformes: Crenuchidae), a new fish from<br />
Venezuela.—Zootaxa 1247:1–12.<br />
Thomas, M. R., & L. H. Rapp Py-Daniel. 2008. Three new species <strong>of</strong><br />
the armored catfish genus Loricaria (Siluriformes: Loricariidae)<br />
from river channels <strong>of</strong> the Amazon basin.—Neotropical<br />
Ichthyology 6(3):379–394.<br />
Vari, R. P., & C. J. Ferraris, Jr. 2006. The catfish genus Tetranematichthys<br />
(auchenipteridae).—Copeia 2006(2):168–180.<br />
———, ———, & M. C. C. de Pinna. 2005. The Neotropical whale<br />
catfishes (Siluriformes: Cetopsidae: Cetopsinae), a revisionary<br />
study.—Neotropical Ichthyology 3(2):127–238.<br />
Vermeulen, F. B. M., & T. Hrbek. 2005 Kryptolebias sepia n. sp.<br />
(Actinopterygii: Cyprinodontiformes: Rivulidae), a new killifish<br />
from the Tapanahony River drainage in southeast<br />
Surinam.—Zootaxa 928:1–20.<br />
Werneke, D. C., J. W. Armbruster, N. K. Lujan, & D. C. Taphorn.<br />
2005. Hemiancistrus guahiborum, a new suckermouth armored<br />
catfish from Southern Venezuela (Siluriformes: Loricariidae).—Neotropical<br />
Ichthyology 3(4):543–548.<br />
———, M. H. Sabaj, N. K. Lujan, & J. W. Armbruster. 2005.<br />
Baryancistrus demantoides and Hemiancistrus subviridis, two<br />
new uniquely colored species <strong>of</strong> catfishes from Venezuela<br />
(Siluriformes: Loricariidae).—Neotropical Ichthyology 3(4):<br />
533–542.<br />
Willink, P. W., B. Chern<strong>of</strong>f, A. Machado-Allison, F. Provenzano, &<br />
P. Petry. 2003. Aphyocharax yekwanae, a new species <strong>of</strong><br />
bloodfin tetra (Teleostei: Characiformes: Characidae) from<br />
the Guyana Shield <strong>of</strong> Venezuela.—Ichthyological Exploration<br />
<strong>of</strong> Freshwaters 14(1):1–8.<br />
Zanata, A. M., & M. Toledo-Piza. 2004. Taxonomic revision <strong>of</strong> the<br />
South American fish genus Chalceus Cuvier (Teleostei:<br />
Ostariophysi: Characiformes) with the description <strong>of</strong> three<br />
new species.—Zoological Journal <strong>of</strong> the Linnean <strong>Society</strong><br />
140(1):103–135.<br />
Zarske, A., P.-Y. Le Bail, & J. Géry. 2006. New and poorly known<br />
Characiform fishes from French Guiana. 1. Two new Tetras<br />
<strong>of</strong> the genera Hemigrammus and Hyphessobrycon (Teleostei:<br />
Characiformes: Characidae).—Zoologische Abhandlungen<br />
(Dresden) 55:17–30.<br />
———, J. Géry, & I. Isbrücker. 2004. Moenkhausia rara sp. n. – eine<br />
neue, bereits bestandsgefährdete Salmler-Art (Teleostei: Characiformes:<br />
Characidae) aus Surinam und Französisch Guayana<br />
mit einer ergänzenden Beschreibung von M. simulata (Eigenmann<br />
in Pearson, 1924).—Zoologische Abhandlungen (Dresden)<br />
54:19–30.
CHECKLIST OF THE FISHES OF THE GUIANA SHIELD<br />
Order: Pristiformes<br />
Family: Pristidae—John D. McEachran & M. R. de Carvalho<br />
Pristis pectinata Latham, 1794 DA GU?<br />
Pristis pristis (Linnaeus, 1758) DA<br />
Order: Myliobatiformes<br />
Family: Potamotrygonidae—Marcelo R. de Carvalho & Ricardo<br />
S. Rosa<br />
Paratrygon aiereba (Müller & Henle, 1841) BO DA RO<br />
Potamotrygon boesemani Rosa, Carvalho & Almeida, 2008 GU SU<br />
Potamotrygon marinae Deynat, 2006 FG<br />
Potamotrygon motoro (Müller & Henle, 1841) CG VA BO RO<br />
Potamotrygon orbignyi (Castelnau, 1855) VA BO DA GU SU FG<br />
Potamotrygon schroederi Fernández-Yépez, 1957 VA BO DA<br />
Potamotrygon scobina Garman, 1913 RO PA<br />
Order: Osteoglossiformes<br />
Family: Osteoglossidae—Carl J. Ferraris, Jr.<br />
Osteoglossum bicirrhosum (Cuvier, 1829)<br />
Family: Arapaimidae—Carl J. Ferraris, Jr.<br />
CG RO AP GU FG<br />
Arapaima gigas (Schinz, 1822) GU<br />
Order: Anguilliformes<br />
Family: Ophichthidae—Sven O. Kullander<br />
Stictorhinus potamius (Böhlke & McCosker, 1975) BO DA<br />
Order: Clupeiformes<br />
Family: Clupeidae—Carl J. Ferraris, Jr.<br />
Rhinosardinia bahiensis (Steindachner, 1879)<br />
Family: Engraulidae—Carl J. Ferraris, Jr. & Sven O. Kullander<br />
BO DA AP GU SU FG<br />
Anchoa spinifer (Valenciennes, 1848) DA GU SU FG<br />
Anchovia surinamensis (Bleeker, 1866) BO DA GU SU FG<br />
Anchoviella brevirostris (Günther, 1868) BO DA GU SU FG<br />
Anchoviella cayennensis (Puyo, 1946) SU FG<br />
Anchoviella guianensis (Eigenmann, 1912) BO DA GU SU FG<br />
Anchoviella jamesi (Jordan & Seale, 1926) VA BO BA<br />
Anchoviella lepidentostole (Fowler 1911) GU SU FG<br />
Lycengraulis batesii (Günther, 1868) BO DA GU SU FG<br />
Lycengraulis grossidens (Spix & Agassiz, 1829) DA SU FG<br />
Pterengraulis atherinoides (Linnaeus, 1766)<br />
Family: Pristigasteridae—Fabio di Dario<br />
BO DA SU FG<br />
Pellona castelnaeana (Valenciennes, 1847) CG VA BO DA<br />
Pellona flavipinnis (Valenciennes, 1836) CG VA BO DA GU SU FG<br />
Family: Parodontidae—Carla S. Pavanelli<br />
Apareiodon agmatos Taphorn, López-Fernández & Bernard,<br />
2008<br />
Order: Characiformes<br />
Apareiodon gransabana Starnes & Schindler, 1993 BO GU SU FG<br />
Apareiodon orinocensis Bonilla et al., 1999 VA BO<br />
GU
26 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Parodon apolinari Myers, 1930 CG VA BO<br />
Parodon bifasciatus Eigenmann, 1912 RO GU<br />
Parodon guyanensis Géry, 1959 VA BO GU SU FG<br />
Parodon suborbitalis Valenciennes, 1850<br />
Family: Curimatidae—Richard P. Vari<br />
BO<br />
Curimata cerasina Vari, 1984 VA BO DA<br />
Curimata cisandina (Allen, 1942) BA<br />
Curimata cyprinoides (Linnaeus, 1766) DA GU SU FG<br />
Curimata incompta Vari, 1984 VA BO DA<br />
Curimata ocellata (Eigenmann & Eigenmann, 1889) CG VA BO BA<br />
Curimata roseni Vari, 1989 VA BO DA GU<br />
Curimata vittata (Kner, 1858) CG VA BO GU<br />
Curimatella alburna (Müller & Troschel, 1844) RO GU<br />
Curimatella dorsalis (Eigenmann & Eigenmann, 1889) CG VA BO DA<br />
Curimatella immaculata (Fernández-Yépez, 1948) CG VA BO DA RO GU<br />
Curimatopsis crypticus Vari, 1982 AP GU SU FG<br />
Curimatopsis evelynae Géry, 1964 CG VA BA?<br />
Curimatopsis macrolepis (Steindachner, 1876) CG VA BO DA BA<br />
Cyphocharax abramoides (Kner, 1859) CG VA BO PA<br />
Cyphocharax festivus Vari, 1992 VA BO GU<br />
Cyphocharax helleri (Steindachner, 1910) BO AP GU SU FG<br />
Cyphocharax leucostictus (Eigenmann & Eigenmann, 1889) VA RO<br />
Cyphocharax meniscaprorus Vari, 1992 VA BO<br />
Cyphocharax mestomyllon Vari, 1992 BA<br />
Cyphocharax microcephalus (Eigenmann & Eigenmann, 1889) GU SU FG?<br />
Cyphocharax multilineatus (Myers, 1927) VA BO BA<br />
Cyphocharax notatus (Steindachner, 1908) VA<br />
Cyphocharax oenas Vari, 1992 VA BO DA<br />
Cyphocharax punctatus (Vari & Nijssen, 1986) SU FG<br />
Cyphocharax spilurus (Günther, 1864) CG VA BO BA RO GU SU FG<br />
Potamorhina altamazonica (Cope, 1878) CG VA BO DA<br />
Psectrogaster ciliata (Müller & Troschel, 1844) CG VA BO DA RO GU<br />
Psectrogaster essequibensis (Günther, 1864) GU<br />
Steindachnerina argentea (Gill, 1858) CG VA BO DA<br />
Steindachnerina bimaculata (Steindachner, 1876) BO DA<br />
Steindachnerina guentheri (Eigenmann & Eigenmann, 1889) BO DA GU<br />
Steindachnerina planiventris Vari & Vari, 1989 RO<br />
Steindachnerina pupula Vari, 1991 BO<br />
Steindachnerina varii Géry, Planquette & Le Bail, 1991<br />
Family: Prochilodontidae—Richard P. Vari<br />
AP SU FG<br />
Prochilodus mariae Eigenmann, 1922 CG VA BO DA<br />
Prochilodus rubrotaeniatus Jardine, 1841 VA BO RO GU SU FG<br />
Semaprochilodus insignis (Jardine, 1841) CG GU<br />
Semaprochilodus kneri (Pellegrin, 1909) VA BO DA<br />
Semaprochilodus laticeps (Steindachner, 1879) CG VA BO DA<br />
Semaprochilodus varii Castro, 1988<br />
Family: Anostomidae—Richard P. Vari<br />
SU FG<br />
Abramites hypselonotus (Günther, 1868) VA BO DA GU
NUMBER 17 27<br />
Anostomoides atrianalis Pellegrin, 1908 VA BO<br />
Anostomoides laticeps (Eigenmann, 1912) VA BO GU<br />
Anostomus anostomus (Linnaeus, 1758) GU<br />
Anostomus brevior Géry, 1961 FG<br />
Anostomus plicatus Eigenmann, 1912 GU SU<br />
Anostomus spiloclistron Winterbottom, 1974 SU<br />
Anostomus ternetzi Fernández-Yépez, 1949 VA BO DA GU SU FG<br />
Gnathodolus bidens Myers, 1927 VA BO<br />
Laemolyta fernandezi Myers, 1950 VA BO DA<br />
Laemolyta orinocensis (Steindachner, 1879) VA BO DA<br />
Laemolyta proximate (Garman, 1890) BA GU<br />
Laemolyta taeniata (Kner, 1859) CG VA BO<br />
Leporinus acutidens (Valenciennes, 1836) FG<br />
Leporinus affinis Günther, 1864 BO<br />
Leporinus agassizi Steindachner, 1876 VA BO<br />
Leporinus alternus Eigenmann, 1912 DA GU<br />
Leporinus arcus Eigenmann, 1912 VA BO GU SU FG<br />
Leporinus badueli Puyo, 1948 FG<br />
Leporinus brunneus Myers, 1950 VA BO BA<br />
Leporinus desmotes Fowler, 1914 GU SU<br />
Leporinus despaxi Puyo, 1943 RO FG<br />
Leporinus fasciatus (Bloch, 1794) SU FG<br />
Leporinus friderici (Bloch, 1794) GU SU FG<br />
Leporinus gossei Géry, Planquette & Le Bail, 1991 SU FG<br />
Leporinus granti Eigenmann, 1912 GU FG<br />
Leporinus lat<strong>of</strong>asciatus Steindachner, 1910 VA BO DA<br />
Leporinus lebaili Géry & Planquette, 1983 SU FG<br />
Leporinus leschenaulti Valenciennes, 1850 FG<br />
Leporinus maculatus Müller & Troschel, 1844 GU SU FG<br />
Leporinus megalepis Günther, 1863 GU<br />
Leporinus melanostictus Norman, 1926 AP FG<br />
Leporinus nigrotaeniatus (Jardine, 1841) GU<br />
Leporinus nijsseni Garavello, 1990 SU FG<br />
Leporinus ortomaculatus Garavello, 2000 VA BO<br />
Leporinus paralternus Fowler, 1914 GU<br />
Leporinus pellegrinii Steindachner, 1910 GU SU FG<br />
Leporinus pitingai Santos & Jegu, 1996 BA<br />
Leporinus punctatus Garavello, 2000 VA BO<br />
Leporinus spilopleura Norman, 1926 AP FG<br />
Leporinus steyermarki Inger, 1956 VA BO DA<br />
Leporinus uatumaensis Santos & Jegu, 1996 BA<br />
Leporinus yophorus Eigenmann, 1922 VA BO<br />
Pseudanos gracilis (Kner, 1858) VA BO BA<br />
Pseudanos irinae Winterbottom, 1980 VA BO<br />
Pseudanos trimaculatus (Kner, 1858) GU<br />
Pseudanos winterbottomi Sidlauskas & Santos, 2005 VA BO<br />
Sartor elongatus Santos & Jégu, 1987 PA<br />
Schizodon fasciatus Spix & Agassiz, 1829 BO FG
28 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Schizodon scotorhabdotus Sidlauskas, Garavello & Jellen, 2007 VA BO DA<br />
Synaptolaemus cingulatus Myers & Fernández-Yépez, 1950<br />
Family: Chilodontidae—Richard P. Vari<br />
VA BO<br />
Caenotropus labyrinthicus (Kner, 1858) CG VA BO DA GU SU<br />
Caenotropus maculosus (Eigenmann, 1912) BO GU SU FG<br />
Caenotropus mestomorgmatos Vari, Castro & Raredon, 1995 VA BO BA<br />
Chilodus punctatus Müller & Troschel, 1844 CG VA BO DA GU SU<br />
Chilodus zunevei Puyo, 1946<br />
Family: Crenuchidae—Paulo A. Buckup<br />
FG<br />
Ammocryptocharax elegans Weitzman & Kanazawa, 1976 CG VA BO<br />
Ammocryptocharax lateralis (Eigenmann, 1909) GU<br />
Ammocryptocharax minutus Buckup, 1993 CG VA BA<br />
Ammocryptocharax vintonae (Eigenmann, 1909) BO GU<br />
Characidium boaevistae Steindachner, 1915 VA BO RO<br />
Characidium chupa Schultz, 1944 BO<br />
Characidium declivirostre Steindachner, 1915 VA BO BA<br />
Characidium hasemani Steindachner, 1915 BO RO<br />
Characidium longum Taphorn, Montaña & Buckup, 2006 VA BO BA<br />
Characidium pellucidum Eigenmann, 1909 GU SU FG<br />
Characidium pteroides Eigenmann, 1909 GU<br />
Characidium steindachneri Cope, 1878 CG VA BO BA RO PA GU<br />
Characidium zebra Eigenmann, 1909 CG VA BO DA BA RO PA AP GU SU FG<br />
Crenuchus spilurus Günther, 1863 CG VA BO GU SU FG<br />
Elachocharax geryi Weitzman & Kanazawa, 1978 CG VA BO BA<br />
Elachocharax mitopterus Weitzman, 1986 VA BA<br />
Elachocharax pulcher Myers, 1927 CG VA BO BA PA<br />
Leptocharacidium omospilus Buckup, 1993 VA BO BA RO<br />
Melanocharacidium blennioides (Eigenmann, 1909) BO GU<br />
Melanocharacidium compressus Buckup, 1993 VA BO<br />
Melanocharacidium depressum Buckup, 1993 VA BA<br />
Melanocharacidium dispilomma Buckup, 1993 CG VA BO BA RO PA FG<br />
Melanocharacidium melanopteron Buckup, 1993 VA BO<br />
Melanocharacidium nigrum Buckup, 1993 BA RO<br />
Melanocharacidium pectorale Buckup, 1993 VA BO BA RO PA<br />
Microcharacidium eleotrioides (Géry, 1960) SU FG<br />
Microcharacidium gnomus Buckup, 1993 CG VA BO BA<br />
Microcharacidium weitzmani Buckup, 1993 CG VA BO BA PA<br />
Odontocharacidium aphanes (Weitzman & Kanazawa, 1977) CG VA BA<br />
Poecilocharax bovalii Eigenmann, 1909 GU<br />
Poecilocharax weitzmani Géry, 1965 CG VA BO<br />
Skiotocharax meizon Presswell, Weitzman & Bergquist, 2000<br />
Family: Hemiodontidae—Francisco Langeani Neto<br />
GU<br />
Anodus elongatus Agassiz, 1829 PA<br />
Anodus orinocensis (Steindachner, 1887) CG VA BO DA<br />
Argonectes longiceps (Kner, 1858) PA SU FG<br />
Bivibranchia bimaculata Vari, 1985 GU SU FG<br />
Bivibranchia fowleri (Steindachner, 1908) CG VA BO DA BA RO GU<br />
Bivibranchia simulata Géry, Planquette & Le Bail, 1991 AP FG
NUMBER 17 29<br />
Bivibranchia velox (Eigenmann & Myers, 1927) AP<br />
Hemiodus amazonum (Humboldt, 1821) GU<br />
Hemiodus argenteus Pellegrin, 1908 CG VA BO DA GU SU<br />
Hemiodus atranalis (Fowler, 1940) RO PA GU<br />
Hemiodus goeldii Steindachner, 1908 AP<br />
Hemiodus gracilis Günther, 1864 VA BO<br />
Hemiodus huraulti (Géry, 1964) SU FG<br />
Hemiodus immaculatus Kner, 1858 CG VA BO DA PA<br />
Hemiodus microlepis Kner, 1858 VA BO<br />
Hemiodus quadrimaculatus Pellegrin, 1908 RO AP GU SU FG<br />
Hemiodus semitaeniatus Kner, 1858 CG VA BO GU<br />
Hemiodus thayeria Böhlke, 1955 CG VA BO BA<br />
Hemiodus unimaculatus (Bloch, 1794) AP GU SU FG<br />
Hemiodus vorderwinkleri (Géry, 1964)<br />
Family: Gasteropelecidae—Stanley H. Weitzman<br />
BO GU<br />
Carnegiella marthae Myers, 1927 CG VA BO DA BA<br />
Carnegiella strigata (Günther, 1864) CG VA BO DA GU SU<br />
Gasteropelecus sternicla (Linnaeus, 1758) BO DA GU FG<br />
Thoracocharax stellatus (Kner, 1858)<br />
Family: Characidae<br />
Genera Incerta Sedis—Flávio C. T. Lima<br />
CG VA BO DA<br />
Aphyocharacidium melandetum (Eigenmann, 1912) GU<br />
Aphyodite grammica Eigenmann, 1912 GU<br />
Astyanax bimaculatus (Linnaeus, 1758) CG VA BO DA GU SU FG<br />
Astyanax clavitaeniatus Garutti, 2003 RO<br />
Astyanax fasciatus (Cuvier, 1819) BO<br />
Astyanax guianensis Eigenmann, 1909 VA BO GU<br />
Astyanax leopoldi Géry, Planquette & Le Bail, 1988 AP FG<br />
Astyanax metae Eigenmann, 1914 VA BO<br />
Astyanax mutator Eigenmann, 1909 GU<br />
Astyanax myersi (Fernández-Yépez, 1950) VA BO<br />
Astyanax rupununi Garutti, 2003 GU<br />
Astyanax scintillans Myers, 1928 VA BO<br />
Astyanax siapae Garutti, 2003 VA<br />
Astyanax validus Géry, Planquette & Le Bail, 1991 FG<br />
Astyanax venezuelae Schultz, 1944 VA BO<br />
Aulixidens eugeniae Böhlke, 1952 VA BO<br />
Bryconamericus alpha Eigenmann, 1914 CG VA BO<br />
Bryconamericus beta Eigenmann, 1914 VA BO<br />
Bryconamericus cinarucoense Román-Valencia, Taphorn & Ruiz-<br />
C., 2008<br />
GU<br />
Bryconamericus cismontanus Eigenmann, 1914 VA BO<br />
Bryconamericus cristiani Román-Valencia, 1998 CG<br />
Bryconamericus deuterodonoides Eigenmann, 1914 VA BO<br />
Bryconamericus hyphesson Eigenmann, 1909 GU<br />
Bryconamericus macropthalmus Román-Valencia, 2003 VA<br />
Bryconamericus orinocoense Román-Valencia, 2003 VA<br />
Bryconamericus subtilisform Román-Valencia, 2003 BO<br />
Bryconexodon trombetasi Jégu, Santos & Ferreira, 1991 PA
30 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Bryconops affinis (Günther, 1864) GU SU FG<br />
Bryconops alburnoides Kner, 1858 CG VA BO<br />
Bryconops caudomaculatus (Günther, 1864) VA BO DA GU SU FG<br />
Bryconops colanegra Chern<strong>of</strong>f & Machado-Allison, 1999 VA BO<br />
Bryconops colaroja Chern<strong>of</strong>f & Machado-Allison, 1999 BO<br />
Bryconops collettei Chern<strong>of</strong>f & Machado-Allison, 2005 VA BO<br />
Bryconops cyrtogaster (Norman, 1926) AP FG<br />
Bryconops disruptus Machado-Allison & Chern<strong>of</strong>f, 1997 VA BA<br />
Bryconops giacopinii (Fernández-Yépez, 1950) VA BO<br />
Bryconops humeralis Machado-Allison, Chern<strong>of</strong>f & Buckup,<br />
1996<br />
VA BO<br />
Bryconops imitator Chern<strong>of</strong>f & Machado-Allison, 2002 VA BO<br />
Bryconops inpai Knöppel, Junk & Géry, 1968 VA BA<br />
Bryconops melanurus (Bloch, 1794) GU SU FG<br />
Bryconops vibex Machado-Allison, Chern<strong>of</strong>f & Buckup, 1996 VA BO<br />
Ceratobranchia joanae Chern<strong>of</strong>f & Machado-Allison, 1990 VA BO<br />
Chalceus epakros Zanata & Toleda-Piza, 2004 VA BO RO PA GU<br />
Chalceus macrolepidotus Cuvier, 1816 CG VA BO DA GU SU FG<br />
Creagrutus bolivari Schultz, 1944 CG VA BO DA<br />
Creagrutus ephippiatus Vari & Harold, 2001 VA BO DA<br />
Creagrutus gyrospilus Vari & Harold, 2001 VA BO<br />
Creagrutus machadoi Vari & Harold, 2001 BO<br />
Creagrutus magoi Vari & Harold, 2001 VA BO<br />
Creagrutus maxillaris (Myers, 1927) CG VA BO BA<br />
Creagrutus melanzonus Eigenmann, 1909 BO GU FG<br />
Creagrutus melasma Vari, Harold & Taphorn, 1994 VA BO<br />
Creagrutus menezesi Vari & Harold, 2001 RO<br />
Creagrutus phasma Myers, 1927 CG VA BO BA<br />
Creagrutus planquettei Géry & Renno, 1989 FG<br />
Creagrutus provenzanoi Vari & Harold, 2001 VA BO<br />
Creagrutus runa Vari & Harold, 2001 CG VA BO BA<br />
Creagrutus veruina Vari & Harold, 2001 VA<br />
Creagrutus vexillapinnus Vari & Harold, 2001 CG VA BA?<br />
Creagrutus xiphos Vari & Harold, 2001 BO<br />
Creagrutus zephyrus Vari & Harold, 2001 CG VA BA?<br />
Ctenobrycon spilurus (Valenciennes, 1850) VA BO DA GU SU FG<br />
Deuterodon potaroensis Eigenmann, 1909 GU<br />
Exodon paradoxus Müller & Troschel, 1844 GU<br />
Gymnocorymbus thayeri Eigenmann, 1908 CG VA BO DA GU<br />
Gymnotichthys hildae Fernández-Yépez, 1950 VA BO<br />
Hemibrycon surinamensis Géry, 1962 SU FG<br />
Hemigrammus aereus Géry, 1959 FG<br />
Hemigrammus analis Durbin, 1909 CG VA BO BA GU<br />
Hemigrammus barrigonae Eigenmann & Henn, 1914 CG VA BO<br />
Hemigrammus belottii (Steindachner, 1882) FG<br />
Hemigrammus boesemani Géry, 1959 SU FG<br />
Hemigrammus cylindricus Durbin, 1909 GU<br />
Hemigrammus erythrozonus Durbin, 1909 GU<br />
Hemigrammus guyanensis Géry, 1959 FG
NUMBER 17 31<br />
Hemigrammus iota Durbin, 1909 GU<br />
Hemigrammus lunatus Durbin, 1918 SU<br />
Hemigrammus marginatus Ellis, 1911 VA BO DA<br />
Hemigrammus micropterus Meek, 1907 VA BO DA<br />
Hemigrammus microstomus Durbin, 1918 VA BO<br />
Hemigrammus mimus Böhlke, 1955 VA BO<br />
Hemigrammus newboldi (Fernández-Yépez, 1949) VA BO DA<br />
Hemigrammus ocellifer (Steindachner, 1882) GU SU FG<br />
Hemigrammus ora Zarske, Le Bail & Géry, 2006 FG<br />
Hemigrammus orthus Durbin, 1909 GU<br />
Hemigrammus rhodostomus Ahl, 1924 CG VA BO<br />
Hemigrammus rodwayi Durbin, 1909 GU SU FG<br />
Hemigrammus schmardae (Steindachner, 1882) CG VA BO BA<br />
Hemigrammus stictus (Durbin, 1909) CG VA BO GU<br />
Hemigrammus taphorni Benine & Lopes, 2007 BO<br />
Hemigrammus unilineatus (Gill, 1858) GU SU FG<br />
Hyphessobrycon albolineatum Fernández-Yépez, 1950 VA BO<br />
Hyphessobrycon borealis Zarske, Le Bail & Géry, 2006 AP SU FG<br />
Hyphessobrycon catableptus (Durbin, 1909) GU<br />
Hyphessobrycon copelandi Durbin, 1908 FG<br />
Hyphessobrycon diancistrus Weitzman, 1977 CG VA BO<br />
Hyphessobrycon eos Durbin, 1909 GU<br />
Hyphessobrycon epicharis Weitzman & Palmer, 1997 CG VA BA<br />
Hyphessobrycon eques (Steindachner, 1882) FG<br />
Hyphessobrycon georgettae Géry, 1961 SU<br />
Hyphessobrycon hildae Fernández-Yépez, 1950 VA BO<br />
Hyphessobrycon metae Eigenmann & Henn, 1914 CG VA BO<br />
Hyphessobrycon minimus Durbin, 1909 GU<br />
Hyphessobrycon minor Durbin, 1909 GU<br />
Hyphessobrycon rosaceus Durbin, 1909 GU SU<br />
Hyphessobrycon roseus (Géry, 1960) FG<br />
Hyphessobrycon simulatus (Géry, 1960) FG<br />
Hyphessobrycon sweglesi (Géry, 1961) CG VA BO<br />
Hyphessobrycon takasei Géry, 1964 AP FG<br />
Jupiaba abramoides (Eigenmann, 1909) CG VA BO GU SU FG<br />
Jupiaba atypindi Zanata, 1997 RO<br />
Jupiaba essequibensis (Eigenmann, 1909) GU<br />
Jupiaba keithi (Géry, Planquette & Le Bail, 1996) FG<br />
Jupiaba maroniensis (Géry, Planquette & Le Bail, 1996) FG<br />
Jupiaba meunieri (Géry, Planquette & Le Bail, 1996) SU FG<br />
Jupiaba mucronata (Eigenmann, 1909) GU<br />
Jupiaba ocellata (Géry, Planquette & Le Bail, 1996) AP? FG<br />
Jupiaba pinnata (Eigenmann, 1909) GU SU<br />
Jupiaba polylepis (Günther, 1864) PA GU SU<br />
Jupiaba potaroensis (Eigenmann, 1909) GU<br />
Jupiaba scologaster (Weitzman & Vari, 1986) CG VA BO BA<br />
Knodus heteresthes (Eigenmann, 1908) VA BO<br />
Microschemobrycon callops Böhlke, 1953 VA BO PA
32 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Microschemobrycon casiquiare Böhlke, 1953 VA BA<br />
Microschemobrycon melanotus (Eigenmann, 1912) GU<br />
Microschemobrycon meyburgi Meinken, 1975 RO<br />
Moenkhausia browni Eigenmann, 1909 GU<br />
Moenkhausia chrysargyrea (Günther, 1864) VA BO GU FG<br />
Moenkhausia collettii (Steindachner, 1882) CG VA BO DA GU SU FG<br />
Moenkhausia copei (Steindachner, 1882) CG VA BO<br />
Moenkhausia cotinho Eigenmann, 1908 CG VA BO<br />
Moenkhausia dichroura (Kner, 1858) VA BO DA<br />
Moenkhausia georgiae Géry, 1965 SU FG<br />
Moenkhausia grandisquamis (Müller & Troschel, 1845) VA BO DA GU SU FG<br />
Moenkhausia hemigrammoides Géry, 1965 SU FG<br />
Moenkhausia inrai Géry, 1992 FG<br />
Moenkhausia intermedia Eigenmann, 1908 CG VA BO FG<br />
Moenkhausia lata Eigenmann, 1908 AP FG<br />
Moenkhausia lepidura (Kner, 1858) VA BO DA GU SU<br />
Moenkhausia megalops (Eigenmann, 1907) FG<br />
Moenkhausia miangi Steindachner, 1915 BO RO<br />
Moenkhausia moisae Géry, Planquette & Le Bail, 1995 FG<br />
Moenkhausia oligolepis (Günther, 1864) CG VA BO DA GU SU FG<br />
Moenkhausia rara Zarske, Géry & Isbrücker, 2004 PA SU FG<br />
Moenkhausia shideleri Eigenmann, 1909 GU<br />
Moenkhausia surinamensis Géry, 1965 AP SU FG<br />
Paracheirodon axelrodi (Schultz, 1956) CG VA BO BA<br />
Paracheirodon simulans (Géry, 1963) CG VA BO BA<br />
Paragoniates alburnus Steindachner, 1876 VA BO<br />
Parapristella aubynei (Eigenmann, 1909) GU<br />
Pristella maxillaris (Ulrey, 1894) VA BO GU FG<br />
Salminus hilarii Valenciennes, 1850 CG VA BO RO<br />
Scissor macrocephalus Günther, 1864 SU<br />
Serrabrycon magoi Vari, 1986 CG VA BO BA<br />
Thayeria ifati Géry, 1959 FG<br />
Triportheus angulatus (Spix & Agassiz, 1829) GU?<br />
Triportheus auritus Malabarba, 2004 VA BO DA GU?<br />
Triportheus brachipomus Malabarba, 2004 VA AP? GU SU FG<br />
Triportheus elongatus (Günther, 1864) VA BO<br />
Triportheus venezuelensis Malabarba, 2004 BO<br />
Xenagoniates bondi Myers, 1942<br />
Subfamily: Agoniatinae—Angela M. Zanata<br />
CG VA BO DA<br />
Agoniates anchovia Eigenmann, 1914 PA<br />
Agoniates halecinus Müller & Troschel, 1845<br />
Subfamily: Iguanodectinae—Cristiano R. Moreira<br />
VA BO RO PA AP? GU<br />
Iguanodectes adujai Géry, 1970 VA BA<br />
Iguanodectes geisleri Géry, 1970 VA BA<br />
Iguanodectes spilurus (Günther, 1864) CG VA BO DA BA RO AP GU<br />
Piabucus dentatus (Koelreuter, 1763)<br />
Subfamily: Bryconinae—Flávio C. T. Lima<br />
VA DA GU SU FG<br />
Brycon amazonicus (Spix & Agassiz, 1829) CG VA BO GU
NUMBER 17 33<br />
Brycon bicolor Pellegrin, 1909 VA BO DA<br />
Brycon coquenani Steindachner, 1915 VA BO<br />
Brycon falcatus Müller & Troschel, 1844 CG VA BO RO PA GU SU FG<br />
Brycon pesu Müller & Troschel, 1845<br />
Subfamily: Serrasalminae—Michel Jegú<br />
CG VA BO DA RO PA AP GU SU FG<br />
Acnodon oligacanthus (Müller & Trochel, 1844) GU SU FG<br />
Acnodon senai Jégu & Santos, 1990 PA AP<br />
Catoprion mento (Cuvier, 1819) CG VA BO DA BA RO PA AP GU<br />
Colossoma macropomum (Cuvier, 1818) CG VA BO DA<br />
Metynnis altidorsalis Ahl, 1923 GU SU<br />
Metynnis argenteus Ahl, 1924 VA BO DA GU<br />
Metynnis hypsauchen (Müller & Troschel, 1844) VA BO RO GU<br />
Metynnis lippincottianus (Cope, 1870) CG VA AP? FG<br />
Metynnis luna (Cope, 1878) CG VA BO GU<br />
Metynnis orinocensis (Steindachner, 1908) DA<br />
Mylesinus paraschomburgkii Jégu, Santos & Ferreira, 1989 BA PA AP<br />
Mylesinus schomburgkii Valenciennes, 1850 GU<br />
Myleus knerii (Steindachner, 1881) FG<br />
Myleus setiger Müller & Troschel, 1844 CG VA BO BA RO PA GU SU<br />
Myloplus asterias (Müller & Troschel, 1844) VA BO PA AP GU<br />
Myloplus lobatus (Valenciennes, 1850) PA<br />
Myloplus planquettei Jégu, Keith & Le Bail, 2003 GU SU FG<br />
Myloplus rhomboidalis (Cuvier, 1818) VA RO PA AP GU SU FG<br />
Myloplus rubripinnis (Müller & Troschel, 1844) CG VA BO RO AP GU SU FG<br />
Myloplus schomburgkii (Jardine & Schomburgk, 1841) CG VA BO BA PA<br />
Myloplus ternetzi (Norman, 1929) AP SU FG<br />
Myloplus torquatus (Kner, 1858) VA BO BA RO GU<br />
Mylossoma aureum (Agassiz, 1829) BO DA<br />
Mylossoma duriventre (Cuvier, 1818) VA BO DA AP<br />
Piaractus brachypomus (Cuvier, 1818) VA BO DA GU<br />
Pristobrycon aureus (Spix & Agassiz, 1829) GU<br />
Pristobrycon calmoni (Steindachner, 1908) BO DA BA AP GU<br />
Pristobrycon careospinus Fink & Machado-Allison, 1992 CG VA<br />
Pristobrycon eigenmanni (Norman, 1929) VA BO DA BA RO PA AP GU SU FG<br />
Pristobrycon maculipinnis Fink & Machado-Allison, 1992 VA<br />
Pristobrycon striolatus (Steindachner, 1908) VA BO DA BA PA AP GU SU FG<br />
Pygocentrus cariba (Humboldt & Valenciennes, 1821) CG VA BO DA<br />
Pygocentrus nattereri Kner, 1858 AP GU<br />
Pygopristis denticulata (Cuvier, 1819) VA BO BA PA AP GU SU FG<br />
Serrasalmus altispinis Merckx, Jégu & Santos, 2000 BA PA<br />
Serrasalmus altuvei Ramírez, 1965 VA BO DA<br />
Serrasalmus elongatus Kner, 1858 BO DA BA PA<br />
Serrasalmus gouldingi Fink & Machado-Allison, 1992 VA BO BA PA<br />
Serrasalmus hastatus Fink & Machado-Allison, 2001 BA RO<br />
Serrasalmus irritans Peters, 1877 VA BO DA<br />
Serrasalmus maculatus Kner, 1858 AP<br />
Serrasalmus manueli (Fernández-Yépez & Ramírez, 1967) CG VA BA AP<br />
Serrasalmus medinai Ramírez, 1965 BO DA
34 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Serrasalmus nalseni Fernández-Yépez, 1969 VA BO<br />
Serrasalmus rhombeus (Linnaeus, 1766) CG VA BO DA BA RO PA AP GU SU FG<br />
Serrasalmus serrulatus (Valenciennes, 1850) GU?<br />
Tometes lebaili Jégu, Keith & Belmont-Jégu, 2002 SU FG<br />
Tometes makue Jégu, Santos & Belmont-Jégu, 2002 CG VA BO BA<br />
Tometes trilobatus Valenciennes, 1850<br />
Subfamily: Aphyocharacinae—Rosana S. Lima<br />
AP FG<br />
Aphyocharax avary Souza-Lima, 2003 VA BO GU<br />
Aphyocharax colifax Taphorn & Thomerson, 1991 VA BO<br />
Aphyocharax erythrurus Eigenmann, 1912 GU<br />
Aphyocharax yekwanae Willink, Chern<strong>of</strong>f, Machado-Allison,<br />
Provenzano & Petry, 2003<br />
BO<br />
Subfamily: Characinae—Naércio A. Menezes & Carlos A.S.<br />
Lucena<br />
Acanthocharax microlepis Eigenmann, 1912 GU<br />
Acestrocephalus sardina (Fowler, 1913) VA BO BA GU<br />
Charax apurensis Lucena, 1987 VA BO DA<br />
Charax gibbosus (Linnaeus, 1758) GU SU<br />
Charax hemigrammus (Eigenmann, 1912) GU<br />
Charax metae Eigenmann, 1922 BO<br />
Charax michaeli Lucena, 1989 RO<br />
Charax notulatus Lucena, 1987 VA BO DA<br />
Charax rupununi Eigenmann, 1912 PA GU<br />
Cynopotamus bipunctatus Pellegrin, 1909 VA BO<br />
Cynopotamus essequibensis Eigenmann, 1912 GU SU FG<br />
Gnathocharax steindachneri Fowler, 1913 CG VA BO GU<br />
Heterocharax leptogrammus Toledo-Piza, 2000 VA BA<br />
Heterocharax macrolepis Eigenmann, 1912 CG VA BO GU<br />
Heterocharax virgulatus Toledo-Piza, 2000 VA BA<br />
Lonchogenys ilisha Myers, 1927 CG VA BO BA<br />
Phenacogaster apletostigma Lucena & Gama, 2007 AP<br />
Phenacogaster carteri (Norman, 1934) GU<br />
Phenacogaster megalostictus Eigenmann, 1909 GU<br />
Phenacogaster microstictus Eigenmann, 1909 GU SU<br />
Priocharax ariel Weitzman & Vari, 1987 VA BO<br />
Roeboides affinis (Günther, 1868) VA BO<br />
Roeboides araguaito Lucena, 2003 CG<br />
Roeboides dientonito Schultz, 1944 VA BO DA GU<br />
Roeboides myersii Gill, 1870 VA BO DA<br />
Roeboides numerosus Lucena, 2000 VA BO<br />
Roeboides oligistos Lucena, 2000 RO PA<br />
Roeboides thurni Eigenmann, 1912<br />
Subfamily: Stethaprioninae—Roberto E. Reis<br />
GU SU FG<br />
Brachychalcinus orbicularis (Valenciennes, 1850) GU SU<br />
Poptella brevispina Reis, 1989 RO PA GU SU<br />
Poptella compressa (Günther, 1864) CG VA BO DA GU<br />
Poptella longipinnis (Popta, 1901)<br />
Subfamily: Tetragonopterinae—Roberto E. Reis<br />
CG VA BO DA SU<br />
Tetragonopterus chalceus Spix & Agassiz, 1829 CG VA BO DA GU SU FG
NUMBER 17 35<br />
Tetragonopterus lemniscatus Benine, Pelição & Vari, 2004<br />
Subfamily: Cheirodontinae—Luiz R. Malabarba<br />
SU<br />
Cheirodontops geayi Schultz, 1944 VA BO DA<br />
Odontostilbe gracilis (Géry, 1960) FG<br />
Odontostilbe littoris (Géry, 1960) FG<br />
Odontostilbe pulchra (Gill, 1858) VA BO DA RO<br />
Odontostilbe splendida Bührnheim & Malabarba, 2007<br />
Subfamily: Glandulocaudinae—Stanley H. Weitzman<br />
BO<br />
Ptychocharax rhyacophila Weitzman, Fink, Machado-Allison &<br />
Royero, 1994<br />
VA<br />
Family: Acestrorhynchidae—Naércio A. Menezes<br />
Acestrorhynchus falcatus (Bloch, 1794) VA GU SU FG<br />
Acestrorhynchus falcirostris (Cuvier, 1819) CG VA BA GU<br />
Acestrorhynchus grandoculis Menezes & Géry, 1983 VA BO BA<br />
Acestrorhynchus heterolepis (Cope, 1878) VA<br />
Acestrorhynchus microlepis (Schomburgk, 1841) CG VA BO DA BA GU SU FG<br />
Acestrorhynchus minimus Menezes, 1969 VA BO PA<br />
Acestrorhynchus nasutus Eigenmann, 1912<br />
Family: Cynodontidae—Mônica Toledo-Piza<br />
VA GU<br />
Cynodon gibbus Spix & Agassiz, 1829 CG BO DA GU<br />
Cynodon meionactis Géry, Le Bail & Keith, 1999 SU FG<br />
Cynodon septenarius Toledo-Piza, 2000 VA BO GU<br />
Hydrolycus armatus (Jardine, 1841) CG VA BO DA GU<br />
Hydrolycus tatauaia Toledo-Piza, Menezes & Santos, 1999 VA BO GU<br />
Hydrolycus wallacei Toledo-Piza, Menezes & Santos, 1999 VA BO<br />
Rhaphiodon vulpinus Spix & Agassiz, 1829 VA BO DA GU<br />
Roestes ogilviei (Fowler, 1914)<br />
Family: Erythrinidae—Based on Cl<strong>of</strong>fsca account (Osvaldo O.<br />
Oyakawa)<br />
RO GU<br />
Erythrinus erythrinus (Bloch & Schneider, 1801) CG VA BO DA GU SU FG<br />
Hoplerythrinus gronovii (Valenciennes, 1847) FG<br />
Hoplerythrinus unitaeniatus (Agassiz, 1829) CG VA BO DA AP GU SU FG<br />
Hoplias aimara (Valenciennes, 1847) VA BO FG<br />
Hoplias macrophthalmus (Pellegrin, 1907) CG VA BO DA GU SU FG<br />
Hoplias malabaricus (Bloch, 1794) CG VA BO DA GU SU FG<br />
Hoplias patana (Valenciennes, 1847)<br />
Family: Lebiasinidae—Marilyn Weitzman & Stanley H.<br />
Weitzman<br />
FG<br />
Copella arnoldi (Regan, 1912) PA GU SU FG<br />
Copella carsevennensis (Regan, 1912) AP SU FG<br />
Copella compta (Myers, 1927) CG VA BA<br />
Copella eigenmanni (Regan, 1912) BO DA PA AP GU SU FG<br />
Copella metae (Eigenmann, 1914) CG VA BO DA? BA<br />
Copella nattereri (Steindachner, 1876) CG VA BO BA RO GU<br />
Derhamia h<strong>of</strong>fmannorum Géry & Zarske, 2002 GU<br />
Lebiasina provenzanoi Ardila Rodríguez, 1999 BO<br />
Lebiasina taphorni Ardila Rodríguez, 2004 BO<br />
Lebiasina uruyensis Fernández-Yépez, 1967 BO<br />
Lebiasina yuruaniensis Ardila Rodríguez, 2000 BO
36 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Nannostomus anduzei Fernandez & Weitzman, 1987 VA BO<br />
Nannostomus beckfordi Günther, 1872 AP GU SU FG<br />
Nannostomus bifasciatus Hoedeman, 1954 SU FG<br />
Nannostomus digrammus (Fowler, 1913) BA RO GU<br />
Nannostomus espei (Meinken, 1956) GU<br />
Nannostomus harrisoni (Eigenmann, 1909) GU<br />
Nannostomus marginatus Eigenmann, 1909 CG VA BO DA GU SU<br />
Nannostomus marilynae Weitzman & Cobb, 1975 CG VA BA<br />
Nannostomus minimus Eigenmann, 1909 GU<br />
Nannostomus trifasciatus Steindachner, 1876 RO? GU<br />
Nannostomus unifasciatus Steindachner, 1876 CG VA BO DA GU GU<br />
Piabucina unitaeniata Günther, 1864 BO GU<br />
Pyrrhulina filamentosa Valenciennes, 1847 VA BO DA GU SU FG<br />
Pyrrhulina lugubris Eigenmann, 1922 CG VA BO DA<br />
Pyrrhulina semifasciata Steindachner, 1876 RO GU<br />
Pyrrhulina stoli Boeseman, 1953<br />
Family: Ctenoluciidae—Richard P. Vari<br />
GU SU<br />
Boulengerella cuvieri (Agassiz, 1829) CG VA BO DA BA AP FG<br />
Boulengerella lateristriga (Boulenger, 1895) VA BA<br />
Boulengerella lucius (Cuvier, 1816) CG VA BO BA PA AP<br />
Boulengerella maculata (Valenciennes, 1850) CG VA BO DA<br />
Boulengerella xyrekes Vari, 1995 VA BO<br />
Order: Siluriformes<br />
Family: Cetopsidae—Richard P. Vari<br />
Cetopsidium ferreirai Vari, Ferraris & de Pinna, 2005 PA<br />
Cetopsidium minutum Vari, Ferraris & de Pinna, 2005 GU SU FG<br />
Cetopsidium morenoi Vari, Ferraris & de Pinna, 2005 VA BO<br />
Cetopsidium orientale Vari, Ferraris & de Pinna, 2005 AP GU SU FG<br />
Cetopsidium pemon Vari, Ferraris & de Pinna, 2005 BO RO GU<br />
Cetopsidium roae Vari, Ferraris & de Pinna, 2005 GU<br />
Cetopsis coecutiens (Lichtenstein, 1819) CG VA BO<br />
Cetopsis orinoco Vari, Ferraris & de Pinna, 2005 BO<br />
Denticetopsis iwokrama Vari, Ferraris & de Pinna, 2005 GU<br />
Denticetopsis macilenta Vari, Ferraris & de Pinna, 2005 GU<br />
Denticetopsis praecox Vari, Ferraris & de Pinna, 2005 VA<br />
Denticetopsis royeroi Ferraris, 1996 VA<br />
Denticetopsis sauli Ferraris, 1996 VA<br />
Helogenes castaneus (Dahl, 1960) CG<br />
Helogenes marmoratus Günther, 1863 VA BA GU SU FG<br />
Helogenes uruyensis Fernández-Yépez, 1967<br />
Family: Aspredinidae—John P. Friel<br />
BO<br />
Acanthobunocephalus nicoi Friel, 1995 VA BO<br />
Amaralia hypsiura (Kner, 1855) GU<br />
Aspredinichthys filamentosus (Valenciennes, 1840) DA GU SU FG<br />
Aspredinichthys tibicen (Valenciennes, 1840) DA GU SU FG<br />
Aspredo aspredo (Linnaeus, 1758) DA GU SU FG<br />
Bunocephalus aleuropsis Cope, 1870 VA BO<br />
Bunocephalus amaurus Eigenmann, 1912 VA BO GU SU FG
NUMBER 17 37<br />
Bunocephalus chamaizelus Eigenmann, 1912 GU<br />
Bunocephalus verrucosus (Walbaum, 1792) GU SU<br />
Ernstichthys anduzei Fernández-Yépez, 1953 VA BO<br />
Hoplomyzon sexpapilostoma Taphorn & Marrero, 1990 VA BO<br />
Platystacus cotylephorus Bloch, 1794 DA GU SU FG<br />
Pseudobunocephalus lundbergi Friel, 2008 BO<br />
Pterobunocephalus depressus (Haseman, 1911)<br />
Family: Trichomycteridae—Wolmar Wosiacki<br />
VA BO<br />
Ammoglanis pulex de Pinna & Winemiller, 2000 VA BO<br />
Glanapteryx anguilla Myers, 1927 VA BO BA<br />
Glanapteryx niobium de Pinna, 1998 BA<br />
Haemomaster venezuelae Myers, 1927 VA BO<br />
Henonemus taxistigmus (Fowler, 1914) GU<br />
Henonemus triacanthopomus DoNascimiento & Provenzano, 2006 DA<br />
Ituglanis amazonicus (Steindachner, 1882) FG<br />
Ituglanis gracilior (Eigenmann, 1912) GU<br />
Ituglanis metae (Eigenmann, 1917) CG VA? BO?<br />
Ituglanis nebulosus de Pinna & Keith, 2003 FG<br />
Megalocentor echthrus de Pinna & Britski, 1991 VA BO<br />
Ochmacanthus alternus Myers, 1927 CG VA BO DA<br />
Ochmacanthus flabelliferus Eigenmann, 1912 BO? GU<br />
Ochmacanthus orinoco Myers, 1927 VA BO<br />
Ochmacanthus reinhardtii (Steindachner, 1882) FG<br />
Paracanthopoma parva Giltay, 1935 RO<br />
Pseudostegophilus haemomyzon (Myers, 1942) VA BO<br />
Pygidianops cuao Schaefer, Provenzano, de Pinna & Baskin,<br />
2005<br />
VA<br />
Pygidianops magoi Schaefer, Provenzano, de Pinna & Baskin,<br />
2005<br />
BO DA<br />
Schultzichthys bondi (Myers, 1942) VA BO<br />
Stauroglanis gouldingi de Pinna, 1989 BA<br />
Stegophilus septentrionalis Myers, 1927 VA BO<br />
Trichomycterus celsae Lasso & Provenzano, 2002 BO<br />
Trichomycterus conradi (Eigenmann, 1912) BO? GU<br />
Trichomycterus guianensis (Eigenmann, 1909) VA BO GU FG<br />
Trichomycterus lewi Lasso & Provenzano, 2002 BO<br />
Trichomycterus santanderensis Castellanos-Morales, 2007 CG<br />
Typhlobelus guacamaya Schaefer, Provenzano, de Pinna &<br />
Baskin, 2005<br />
VA<br />
Typhlobelus lundbergi Schaefer, Provenzano, de Pinna & Baskin,<br />
2005<br />
BO DA<br />
Vandellia beccarii Di Caporiacco, 1935 CG VA BO DA GU<br />
Vandellia sanguinea Eigenmann, 1917<br />
Family: Callichthyidae—Roberto E. Reis<br />
VA BO GU<br />
Callichthys callichthys (Linnaeus, 1758) CG VA BO DA GU SU FG<br />
Callichthys serralabium Lehmann & Reis, 2004 VA BA<br />
Corydoras aeneus (Gill, 1858) CG VA BO GU SU FG<br />
Corydoras amapaensis Nijssen, 1972 AP FG<br />
Corydoras approuaguensis Nijssen & Isbrücker, 1983 FG<br />
Corydoras axelrodi Rössel, 1962 CG VA BO
38 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Corydoras baderi Geisler, 1969 PA SU<br />
Corydoras bicolor Nijssen & Isbrücker, 1967 SU<br />
Corydoras blochi Nijssen, 1971 VA BO RO GU<br />
Corydoras boehlkei Nijssen & Isbrücker, 1982 VA BO<br />
Corydoras boesemani Nijssen & Isbrücker, 1967 SU<br />
Corydoras bondi Gosline, 1940 VA BO GU SU<br />
Corydoras breei Isbrücker & Nijssen, 1992 SU<br />
Corydoras brevirostris Fraser-Brunner, 1947 VA BO SU<br />
Corydoras concolor Weitzman, 1961 VA BO<br />
Corydoras condiscipulus Nijssen & Isbrücker, 1980 AP FG<br />
Corydoras coppenamensis Nijssen, 1970 SU<br />
Corydoras delphax Nijssen & Isbrücker, 1983 CG VA<br />
Corydoras ephippifer Nijssen, 1972 AP<br />
Corydoras filamentosus Nijssen & Isbrücker, 1983 SU<br />
Corydoras ge<strong>of</strong>froy La Cepède, 1803 SU FG<br />
Corydoras griseus Holly, 1940 GU<br />
Corydoras guianensis Nijssen, 1970 SU FG<br />
Corydoras habrosus Weitzman, 1960 CG? VA BO<br />
Corydoras heteromorphus Nijssen, 1970 SU<br />
Corydoras melanistius Regan, 1912 GU SU FG<br />
Corydoras melini Lönnberg & Rendahl, 1930 CG VA? BO?<br />
Corydoras metae Eigenmann, 1914 CG VA BO<br />
Corydoras nanus Nijssen & Isbrücker, 1967 SU FG<br />
Corydoras oiapoquensis Nijssen, 1972 AP FG<br />
Corydoras osteocarus Böhlke, 1951 VA BO SU<br />
Corydoras oxyrhynchus Nijssen & Isbrücker, 1967 SU<br />
Corydoras potaroensis Myers, 1927 GU<br />
Corydoras punctatus (Bloch, 1794) SU FG<br />
Corydoras sanchesi Nijssen & Isbrücker, 1967 SU<br />
Corydoras saramaccensis Nijssen, 1970 SU<br />
Corydoras septentrionalis Gosline, 1940 CG? VA BO<br />
Corydoras sipaliwini Hoedeman, 1965 GU SU<br />
Corydoras solox Nijssen & Isbrücker, 1983 AP FG<br />
Corydoras spilurus Norman, 1926 SU FG<br />
Corydoras surinamensis Nijssen, 1970 SU<br />
Corydoras trilineatus Cope, 1872 SU<br />
Hoplosternum littorale (Hancock, 1828) CG VA BO DA GU SU FG<br />
Megalechis picta (Müller & Troschel, 1848) VA BO DA BA GU<br />
Megalechis thoracata (Valenciennes, 1840)<br />
Family: Loricariidae<br />
Subfamily: Lithogeninae—Carl J. Ferraris, Jr.<br />
VA BO GU SU FG<br />
Lithogenes villosus Eigenmann, 1909 GU<br />
Lithogenes wahari Schaefer & Provenzano, 2008<br />
Subfamily: Hypoptopomatinae—Scott A. Schaefer<br />
VA<br />
Acestridium dichromum Retzer, Nico & Provenzano, 1999 VA<br />
Acestridium martini Retzer, Nico & Provenzano, 1999 VA BO<br />
Hypoptopoma guianense Boeseman, 1974 GU SU<br />
Nannoptopoma spectabile (Eigenmann, 1914) CG? VA? BO
NUMBER 17 39<br />
Niobichthys ferrarisi Schaefer & Provenzano, 1998 VA<br />
Otocinclus huaorani Schaefer, 1997 CG<br />
Otocinclus mariae Fowler, 1940 BA<br />
Otocinclus vittatus Regan, 1904 CG? VA BO<br />
Oxyropsis acutirostra Miranda Ribeiro, 1951 CG VA BO<br />
Parotocinclus britskii Boeseman, 1974 VA BO GU SU<br />
Parotocinclus collinsae Schmidt & Ferraris, 1985 GU<br />
Parotocinclus eppleyi Schaefer & Provenzano, 1993 VA BO<br />
Parotocinclus polyochrus Schaefer, 1988<br />
Subfamily: Loricariinae—Carl J. Ferraris, Jr.<br />
VA<br />
Cteniloricaria platystoma (Günther, 1868) SU<br />
Farlowella nattereri Steindachner, 1910 GU<br />
Farlowella oxyrryncha (Kner, 1853) VA BO<br />
Farlowella reticulata Boeseman, 1971 GU SU FG<br />
Farlowella rugosa Boeseman, 1971 GU SU FG<br />
Farlowella vittata Myers, 1942 CG VA BO DA<br />
Harttia fowleri (Pellegrin, 1908) AP FG<br />
Harttia guianensis Rapp Py-Daniel & Oliveira, 2001 SU FG<br />
Harttia maculata (Boeseman, 1971) SU FG<br />
Harttia merevari Provenzano, Machado-Allison, Chern<strong>of</strong>f,<br />
Willink & Petry, 2005<br />
BO<br />
Harttia surinamensis Boeseman, 1971 SU<br />
Harttia trombetensis Rapp Py-Daniel & Oliveira, 2001 PA<br />
Harttiella crassicauda (Boeseman, 1953) SU<br />
Hemiloricaria castroi Isbrücker & Nijssen, 1984 PA<br />
Hemiloricaria eigenmanni (Pellegrin, 1908) VA BO<br />
Hemiloricaria fallax (Steindachner, 1915) RO GU<br />
Hemiloricaria formosa Isbrücker & Njissen, 1979 CG VA BO DA<br />
Hemiloricaria platyura (Müller & Troschel, 1848) GU FG<br />
Hemiloricaria stewarti (Eigenmann, 1909) GU SU FG<br />
Hemiodontichthys acipenserinus (Kner, 1853) AP GU FG<br />
Limatulichthys griseus (Eigenmann, 1909) CG VA BO DA GU<br />
Loricaria cataphracta Linnaeus, 1758 GU SU FG<br />
Loricaria lundbergi Thomas & Rapp Py-Daniel, 2008 VA<br />
Loricaria nickeriensis Isbrücker, 1979 SU FG<br />
Loricaria parnahybae Steindachner, 1907 AP? FG<br />
Loricaria simillima Regan, 1904 VA BO<br />
Loricaria spinulifera Thomas & Rapp Py-Daniel, 2008 RO<br />
Loricariichthys maculatus (Bloch, 1794) SU<br />
Loricariichthys microdon (Eigenmann, 1909) GU<br />
Metaloricaria nijsseni (Boeseman, 1976) SU<br />
Metaloricaria paucidens Isbrücker, 1975 SU FG<br />
Pseudoloricaria laeviuscula (Valenciennes, 1840) RO<br />
Reganella depressa (Kner, 1853) RO<br />
Sturisoma monopelte Fowler, 1914<br />
Subfamily: Hypostominae—Jonathan W. Armbruster & Claude<br />
Weber<br />
GU<br />
Acanthicus hystrix Spix & Agassiz, 1829 VA BO DA<br />
Ancistrus hoplogenys (Günther, 1864) GU
40 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Ancistrus leucostictus (Günther, 1864) GU SU? FG?<br />
Ancistrus lithurgicus Eigenmann, 1912 GU<br />
Ancistrus macrophthalmus (Pellegrin, 1912) VA BO<br />
Ancistrus nudiceps (Müller & Troschel, 1848) GU<br />
Ancistrus temminckii (Valenciennes, 1840) SU<br />
Ancistrus triradiatus Eigenmann, 1918 CG VA BO DA<br />
Baryancistrus beggini Lujan, Arce & Armbruster, 2009 CG VA<br />
Baryancistrus demantoides Werneke, Sabaj, Lujan &<br />
Armbruster, 2005<br />
VA<br />
Baryancistrus niveatus (Castelnau, 1855) PA<br />
Chaetostoma jegui Rapp Py-Daniel, 1991 RO<br />
Chaetostoma vasquezi Lasso & Provenzano, 1998 VA BO<br />
Corymbophanes andersoni Eigenmann, 1909 GU<br />
Corymbophanes kaiei Armbruster & Sabaj, 2000 GU<br />
Dekeyseria niveata (La Monte, 1929) VA<br />
Dekeyseria pulcher (Steindachner, 1915) CG VA BA?<br />
Dekeyseria scaphirhyncha (Kner, 1854) BA<br />
Exastilithoxus fimbriatus (Steindachner, 1915) BO<br />
Exastilithoxus hoedemani Isbrücker & Nijssen, 1985 BA<br />
Hemiancistrus guahiborum Werneke, Armbruster, Lujan &<br />
Taphorn, 2005<br />
VA BO<br />
Hemiancistrus medians (Kner, 1854) SU FG<br />
Hemiancistrus sabaji Armbruster, 2003 VA GU<br />
Hemiancistrus subviridis Werneke, Sabaj, Lujan & Armbruster,<br />
2005<br />
VA<br />
Hypancistrus contradens Armbruster, Lujan & Taphorn, 2007 VA<br />
Hypancistrus debilittera Armbruster, Lujan & Taphorn, 2007 VA<br />
Hypancistrus furunculus Armbruster, Lujan & Taphorn, 2007 VA<br />
Hypancistrus inspector Armbruster, 2002 VA BO<br />
Hypancistrus lunaorum Armbruster, Lujan & Taphorn, 2007 VA<br />
Hypostomus coppenamensis Boeseman, 1969 SU<br />
Hypostomus corantijni Boeseman, 1968 SU<br />
Hypostomus crassicauda Boeseman, 1968 SU<br />
Hypostomus gymnorhynchus (Norman, 1926) SU FG<br />
Hypostomus hemicochliodon Armbruster, 2003 VA BO<br />
Hypostomus hemiurus (Eigenmann, 1912) GU<br />
Hypostomus macrophthalmus Boeseman, 1968 SU<br />
Hypostomus macushi Armbruster & de Souza, 2005 GU<br />
Hypostomus micromaculatus Boeseman, 1968 SU<br />
Hypostomus nematopterus Isbrücker & Nijssen, 1984 FG<br />
Hypostomus nickeriensis Boeseman, 1969 SU<br />
Hypostomus occidentalis Boeseman, 1968 SU<br />
Hypostomus paucimaculatus Boeseman, 1968 SU<br />
Hypostomus plecostomoides (Eigenmann, 1922) BO<br />
Hypostomus plecostomus (Linnaeus, 1758) GU SU<br />
Hypostomus pseudohemiurus Boeseman, 1968 SU<br />
Hypostomus rhantos Armbruster, Tansey & Lujan, 2007 VA<br />
Hypostomus saramaccensis Boeseman, 1968 SU<br />
Hypostomus sculpodon Armbruster, 2003 VA
NUMBER 17 41<br />
Hypostomus sipaliwini Boeseman, 1968 SU<br />
Hypostomus squalinus (Jardine, 1841) RO GU<br />
Hypostomus surinamensis Boeseman, 1968 SU<br />
Hypostomus tapanahoniensis Boeseman, 1969 SU<br />
Hypostomus taphorni (Lilyestrom, 1984) BO GU<br />
Hypostomus tenuis (Boeseman, 1968) SU?<br />
Hypostomus ventromaculatus Boeseman, 1968 SU FG<br />
Hypostomus villarsi (Lütken, 1874) VA BO<br />
Hypostomus watwata Hancock, 1828 DA GU SU FG<br />
Lasiancistrus schomburgkii (Günther, 1864) GU<br />
Lasiancistrus tentaculatus Armbruster, 2005 VA BO<br />
Leporacanthicus galaxias Isbrücker & Nijssen, 1989 VA BO<br />
Leporacanthicus triactis Isbrücker, Nijssen & Nico, 1992 CG VA BO<br />
Lithoxus boujardi Muller & Isbrücker, 1993 FG<br />
Lithoxus bovallii (Regan, 1906) GU<br />
Lithoxus jantjae Lujan, 2008 VA<br />
Lithoxus lithoides Eigenmann, 1910 GU SU<br />
Lithoxus pallidimaculatus Boeseman, 1982 SU<br />
Lithoxus planquettei Boeseman, 1982 FG<br />
Lithoxus stocki Nijssen & Isbrücker, 1990 FG<br />
Lithoxus surinamensis Boeseman, 1982 SU<br />
Neblinichthys pilosus Ferraris, Isbrücker & Nijssen, 1986 VA<br />
Neblinichthys roraima Provenzano, Lasso & Ponte, 1995 BO<br />
Neblinichthys yaravi (Steindachner, 1915) BO<br />
Panaque maccus Schaefer & Stewart, 1993 VA BO<br />
Panaque nigrolineatus (Peters, 1877) CG VA BO<br />
Peckoltia braueri (Eigenmann, 1912) GU<br />
Peckoltia brevis (La Monte, 1935) CG?<br />
Peckoltia cavatica Armbruster & Werneke, 2005 GU<br />
Peckoltia lineola Armbruster, 2008 CG VA<br />
Peckoltia vittata (Steindachner, 1881) CG? VA PA<br />
Pseudacanthicus fordii (Günther, 1868) SU<br />
Pseudacanthicus leopardus (Fowler, 1914) GU<br />
Pseudacanthicus serratus (Valenciennes, 1840) SU FG<br />
Pseudancistrus barbatus (Valenciennes, 1840) GU SU FG<br />
Pseudancistrus brevispinis (Heitmans, Nijssen & Isbrücker, 1983) SU FG<br />
Pseudancistrus coquenani (Steindachner, 1915) BO<br />
Pseudancistrus corantijniensis De Chambrier & Montoya-<br />
Burgos, 2008<br />
SU<br />
Pseudancistrus depressus (Günther, 1868) SU<br />
Pseudancistrus guentheri (Regan, 1904) GU<br />
Pseudancistrus longispinis (Heitmans, Nijssen & Isbrücker, 1983) FG<br />
Pseudancistrus macrops (Lütken, 1874) SU<br />
Pseudancistrus megacephalus (Günther, 1868) GU SU<br />
Pseudancistrus niger (Norman, 1926) FG<br />
Pseudancistrus nigrescens Eigenmann, 1912 GU<br />
Pseudancistrus orinoco Isbrücker, Nijssen & Cala, 1988 CG VA BO<br />
Pseudancistrus pectegenitor Lujan, Armbruster & Sabaj, 2007 VA<br />
Pseudancistrus reus Armbruster & Taphorn, 2008 BO
42 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Pseudancistrus sidereus Armbruster, 2004 VA<br />
Pseudancistrus yekuana Lujan, Armbruster & Sabaj, 2007 VA<br />
Pseudolithoxus anthrax (Armbruster & Provenzano, 2000) VA BO<br />
Pseudolithoxus dumus (Armbruster & Provenzano, 2000) VA BO<br />
Pseudolithoxus nicoi (Armbruster & Provenzano, 2000) VA<br />
Pseudolithoxus tigris (Armbruster & Provenzano, 2000) VA BO<br />
Pterygoplichthys gibbiceps (Kner, 1854) CG VA BO<br />
Pterygoplichthys multiradiatus (Hancock, 1828)<br />
Family: Pseudopimelodidae—Oscar A. Shibbata<br />
VA BO GU?<br />
Batrochoglanis raninus (Valenciennes, 1840) GU SU FG<br />
Batrochoglanis villosus (Eigenmann, 1912) VA BO GU SU<br />
Cephalosilurus albomarginatus (Eigenmann, 1912) GU<br />
Cephalosilurus nigricaudus (Mees, 1974) SU<br />
Microglanis poecilus Eigenmann, 1912 GU FG<br />
Microglanis secundus Mees, 1974 CG? VA BO GU SU<br />
Pseudopimelodus bufonius (Valenciennes, 1840) CG? VA BO GU SU FG<br />
Family: Heptapteridae—Updated from Cl<strong>of</strong>fsca account (Flávio<br />
A. Bockmann & Gizelani M. Guazzeli)<br />
Brachyglanis frenata Eigenmann, 1912 VA BO BA GU<br />
Brachyglanis magoi Fernández-Yépez, 1967 VA BO<br />
Brachyglanis melas Eigenmann, 1912 GU<br />
Brachyglanis microphthalmus Bizerril, 1991 PA<br />
Brachyglanis phalacra Eigenmann, 1912 GU<br />
Brachyrhamdia heteropleura (Eigenmann, 1912) GU SU<br />
Brachyrhamdia imitator Myers, 1927 VA BO<br />
Cetopsorhamdia insidiosa (Steindachner, 1917) RO<br />
Cetopsorhamdia orinoco Schultz, 1944 BO<br />
Chasmocranus brevior Eigenmann, 1912 GU SU FG<br />
Chasmocranus chimantanus Inger, 1956 BO<br />
Chasmocranus longior Eigenmann, 1912 VA BO BA GU SU<br />
Chasmocranus surinamensis (Bleeker, 1862) SU<br />
Gladioglanis machadoi Ferraris & Mago-Leccia, 1989 VA BO BA?<br />
Goeldiella eques (Müller & Troschel, 1848) CG VA BO BA? GU<br />
Heptapterus bleekeri Boeseman, 1953 AP? SU FG<br />
Heptapterus tapanahoniensis Mees, 1967 SU FG<br />
Heptapterus tenuis Mees, 1986 FG<br />
Imparfinis hasemani Steindachner, 1917 RO<br />
Imparfinis pijpersi (Hoedeman, 1961) SU<br />
Imparfinis pristos Mees & Cala, 1989 CG VA BO<br />
Imparfinis pseudonemacheir Mees & Cala, 1989 CG VA BO<br />
Leptorhamdia essequibensis (Eigenmann, 1912) GU<br />
Leptorhamdia marmorata Myers, 1928 CG? VA BO BA<br />
Mastiglanis asopos Bockmann, 1994 VA BO PA<br />
Medemichthys guayaberensis (Dahl, 1961) CG<br />
Myoglanis aspredinoides DoNascimiento & Lundberg, 2005 VA<br />
Myoglanis potaroensis Eigenmann, 1912 GU<br />
Nemuroglanis pauciradiatus Ferraris, 1988 CG VA BO BA<br />
Phenacorhamdia anisura (Mees, 1987) VA BO<br />
Phenacorhamdia macarenensis Dahl, 1961 CG
NUMBER 17 43<br />
Phenacorhamdia provenzanoi DoNascimiento & Milani, 2008 VA BO<br />
Pimelodella altipinnis (Steindachner, 1864) GU<br />
Pimelodella cristata (Müller & Troschel, 1848) GU FG<br />
Pimelodella cruxenti Fernández-Yépez, 1950 VA BO<br />
Pimelodella figueroai Dahl, 1961 CG<br />
Pimelodella geryi Hoedeman, 1961 SU<br />
Pimelodella linami Schultz, 1944 VA BO<br />
Pimelodella macturki Eigenmann, 1912 GU SU FG<br />
Pimelodella megalops Eigenmann, 1912 GU FG<br />
Pimelodella pallida Dahl, 1961 CG<br />
Pimelodella procera Mees, 1983 FG<br />
Pimelodella wesselii (Steindachner, 1877) GU<br />
Rhamdella leptosoma Fowler, 1914 GU<br />
Rhamdia foina (Müller & Troschel, 1848) RO PA GU<br />
Rhamdia laukidi Bleeker, 1858 CG VA BO GU<br />
Rhamdia muelleri (Günther, 1864) VA BO DA PA GU<br />
Rhamdia quelen (Quoy & Gaimard, 1824)<br />
Family: Pimelodidae—John G. Lundberg<br />
CG VA BO DA GU SU FG<br />
Brachyplatystoma filamentosum (Lichtenstein, 1819) CG VA BO DA GU SU FG<br />
Brachyplatystoma juruense (Boulenger, 1898) CG? VA BO DA<br />
Brachyplatystoma platynemum (Boulenger, 1898) CG VA BO DA<br />
Brachyplatystoma rousseauxii (Castelnau, 1855) BO DA FG<br />
Brachyplatystoma vaillantii (Valenciennes, 1840) CG VA BO DA GU SU FG<br />
Calophysus macropterus (Lichtenstein, 1819) CG VA BO DA PA<br />
Duopalatinus peruanus Eigenmann & Allen, 1942 BO<br />
Exallodontus aguanai Lundberg, Mago-Leccia & Nass, 1991 BO DA<br />
Hemisorubim platyrhynchos (Valenciennes, 1840) CG VA BO DA GU SU FG<br />
Hypophthalmus edentatus Spix & Agassiz, 1829 VA BO DA GU SU<br />
Hypophthalmus fimbriatus Kner, 1858 BO<br />
Hypophthalmus marginatus Valenciennes, 1840 SU FG<br />
Leiarius marmoratus (Gill, 1870) CG VA BO<br />
Leiarius pictus (Müller & Troschel, 1849) CG? VA BO GU<br />
Megalonema amaxanthum Lundberg & Dahdul, 2008 GU<br />
Megalonema platycephalum Eigenmann, 1912 CG VA BO GU<br />
Phractocephalus hemioliopterus (Bloch & Schneider, 1801) CG VA BO DA GU<br />
Pimelodina flavipinnis Steindachner, 1877 BO DA<br />
Pimelodus alb<strong>of</strong>asciatus Mees, 1974 CG VA BO BA SU<br />
Pimelodus blochii Valenciennes, 1840 CG VA BO DA GU SU FG<br />
Pimelodus ornatus Kner, 1858 CG VA BO GU SU FG<br />
Pimelodus pictus Steindachner, 1877 BO<br />
Pinirampus pirinampu (Spix & Agassiz, 1829) CG VA BO DA GU<br />
Platynematichthys notatus (Jardine, 1841) CG VA BO DA<br />
Platysilurus mucosus (Vaillant, 1880) CG VA BO DA<br />
Propimelodus eigenmanni (van der Stigchel, 1946) AP FG<br />
Pseudoplatystoma fasciatum (Linnaeus, 1766) GU SU FG<br />
Pseudoplatystoma metaense Buitrago-Suárez & Burr, 2007 BO DA<br />
Pseudoplatystoma orinocoense Buitrago-Suárez & Burr, 2007 VA BO DA<br />
Sorubim elongatus Littmann, Burr, Schmidt & Isern, 2001 CG VA BO GU
44 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Sorubim lima (Bloch & Schneider, 1801) CG VA BO DA<br />
Sorubimichthys planiceps (Spix & Agassiz, 1829) CG VA BO DA<br />
Zungaro zungaro (Humboldt, 1821)<br />
Family: Doradidae—Mark H. Sabaj Pérez<br />
CG VA BO DA BA GU<br />
Acanthodoras cataphractus (Linnaeus, 1758) CG VA BO? BA RO PA AP GU SU FG<br />
Acanthodoras spinosissimus (Eigenmann & Eigenmann, 1888) CG VA BO? DA? BA RO GU<br />
Agamyxis albomaculatus (Peters, 1877) CG VA BO DA BA?<br />
Amblydoras affinis (Kner, 1855) RO GU<br />
Amblydoras bolivarensis (Fernández-Yépez, 1968) CG VA BO DA? BA?<br />
Amblydoras gonzalezi (Fernández-Yépez, 1968) CG VA BO DA? BA?<br />
Anadoras regani (Steindachner, 1908) PA AP FG<br />
Anduzedoras oxyrhynchus (Valenciennes, 1821) CG VA BA<br />
Centrodoras hasemani (Steindachner, 1915) CG? VA BA<br />
Doras carinatus (Linnaeus, 1766) BO AP GU SU FG<br />
Doras micropoeus (Eigenmann, 1912) GU SU FG<br />
Doras phlyzakion Sabaj Pérez & Birindelli, 2008 RO<br />
Hassar orestis (Steindachner, 1875) CG VA BO DA BA RO? PA? GU<br />
Leptodoras cataniai Sabaj, 2005 VA BA RO?<br />
Leptodoras copei (Fernández-Yépez, 1968) CG VA BO BA RO? PA?<br />
Leptodoras hasemani (Steindachner, 1915) CG VA BO DA BA? RO GU<br />
Leptodoras linnelli Eigenmann, 1912 VA BO BA? RO GU<br />
Leptodoras praelongus (Myers & Weitzman, 1956) CG VA BA RO PA?<br />
Leptodoras rogersae Sabaj, 2005 CG VA BO DA<br />
Megalodoras guayoensis (Fernández-Yépez, 1968) CG? VA? BO DA<br />
Megalodoras uranoscopus (Eigenmann & Eigenmann, 1888) CG? VA? BA? RO? PA? GU<br />
Nemadoras leporhinus (Eigenmann, 1912) CG VA BO BA? RO PA GU<br />
Nemadoras trimaculatus (Boulenger, 1898) CG VA BO BA RO GU<br />
Opsodoras morei (Steindachner, 1881) CG VA BO? BA RO? PA? GU<br />
Opsodoras ternetzi Eigenmann, 1925 CG VA BO DA BA RO PA GU<br />
Orinocodoras eigenmanni Myers, 1927 BO DA<br />
Oxydoras niger (Valenciennes, 1821) CG? VA? BA? RO? GU<br />
Oxydoras sifontesi Fernández-Yépez, 1968 CG? VA? BO DA<br />
Physopyxis ananas Sousa & Rapp Py-Daniel 2005 CG VA BA RO PA GU<br />
Physopyxis cristata Sousa & Rapp Py-Daniel 2005 BA RO?<br />
Platydoras armatulus (Valenciennes 1840) BO DA<br />
Platydoras costatus (Linnaeus, 1758) SU FG<br />
Platydoras hancockii (Valenciennes 1840) CG VA BO? BA RO GU<br />
Pterodoras granulosus (Valenciennes, 1821) CG? VA? BA? RO? PA? GU SU<br />
Pterodoras rivasi (Fernández-Yépez, 1950) CG? VA? BO DA<br />
Rhinodoras armbrusteri Sabaj, Taphorn & Castillo, 2008 GU<br />
Rhynchodoras woodsi Glodek, 1976 RO GU<br />
Scorpiodoras heckelii (Kner, 1855) CG VA BO BA<br />
Trachydoras brevis (Kner, 1853) BA RO GU<br />
Trachydoras microstomus (Eigenmann, 1912)<br />
Family: Auchenipteridae—Carl J. Ferraris, Jr.<br />
CG VA BO DA? BA RO PA GU<br />
Ageneiosus inermis (Linnaeus, 1766) CG VA BO DA GU SU FG<br />
Ageneiosus marmoratus Eigenmann, 1912 GU SU<br />
Ageneiosus piperatus (Eigenmann, 1912) RO GU
NUMBER 17 45<br />
Ageneiosus polystictus Steindachner, 1915 RO<br />
Ageneiosus ucayalensis Castelnau, 1855 VA BO DA GU SU<br />
Ageneiosus vittatus Steindachner, 1908 BO DA<br />
Asterophysus batrachus Kner, 1858 VA BO BA?<br />
Auchenipterus ambyiacus Fowler, 1915 CG? VA BO DA GU<br />
Auchenipterus brevior Eigenmann, 1912 GU<br />
Auchenipterus demerarae Eigenmann, 1912 BO GU<br />
Auchenipterus dentatus Valenciennes, 1840 SU FG<br />
Auchenipterus nuchalis (Spix & Agassiz, 1829) RO? FG<br />
Centromochlus concolor (Mees, 1974) SU<br />
Centromochlus existimatus Mees, 1974 VA BO<br />
Centromochlus punctatus (Mees, 1974) SU<br />
Centromochlus reticulatus (Mees, 1974) GU<br />
Entomocorus gameroi Mago-Leccia, 1984 VA BO DA<br />
Gelanoglanis nanonocticolus Soares-Porto, Walsh, Nico & Netto,<br />
1999<br />
VA BO<br />
Glanidium leopardum (Hoedeman, 1961) GU SU FG<br />
Liosomadoras oncinus (Jardine, 1841) RO<br />
Pseudepapterus gracilis Ferraris & Vari, 2000 BO<br />
Tatia brunnea Mees, 1974 SU FG<br />
Tatia creutzbergi (Boeseman, 1953) SU<br />
Tatia galaxias Mees, 1974 VA BO DA<br />
Tatia gyrina (Eigenmann & Allen, 1942) SU<br />
Tatia intermedia (Steindachner, 1877) GU SU FG<br />
Tatia meesi Sarmento-Soares & Martins-Pinheiro, 2008 GU<br />
Tatia musaica Royero, 1992 VA BO<br />
Tatia nigra Sarmento-Soares & Martins-Pinheiro, 2008 PA<br />
Tatia strigata Soares-Porto, 1995 VA BO BA<br />
Tetranematichthys wallacei Vari & Ferraris, 2006 VA BO BA<br />
Trachelyichthys decaradiatus Mees, 1974 GU<br />
Trachelyopterichthys anduzei Ferraris & Fernandez, 1987 VA BO<br />
Trachelyopterichthys taeniatus (Kner, 1858) CG VA BO BA?<br />
Trachelyopterus ceratophysus (Kner, 1858) RO<br />
Trachelyopterus coriaceus Valenciennes, 1840 FG<br />
Trachelyopterus galeatus (Linnaeus, 1766) SU FG<br />
Trachycorystes trachycorystes Valenciennes, 1840 GU<br />
Order: Gymnotiformes<br />
Family: Gymnotidae—Ricardo Campos-da-Paz<br />
Electrophorus electricus (Linnaeus, 1766) CG VA BO DA GU SU FG<br />
Gymnotus anguillaris Hoedeman, 1962 SU FG<br />
Gymnotus carapo Linnaeus, 1758 VA BO DA FG<br />
Gymnotus cataniapo Mago-Leccia, 1994 BO SU<br />
Gymnotus coropinae Hoedeman, 1962 GU SU<br />
Gymnotus pedanopterus Mago-Leccia, 1994 VA BO BA<br />
Gymnotus stenoleucus Mago-Leccia, 1994<br />
Family: Sternopygidae—James S. Albert<br />
VA BO DA<br />
Archolaemus blax Korringa, 1970 RO AP? FG<br />
Distocyclus conirostris (Eigenmann & Allen, 1942) BO DA
46 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Eigenmannia limbata (Schreiner & Miranda Ribeiro, 1903) BO<br />
Eigenmannia macrops (Boulenger, 1897) BA GU<br />
Eigenmannia nigra Mago-Leccia, 1994 CG VA BO BA GU<br />
Eigenmannia virescens (Valenciennes, 1842) CG VA BO DA BA AP GU SU FG<br />
Rhabdolichops caviceps (Fernández-Yépez, 1968) BO<br />
Rhabdolichops eastwardi Lundberg & Mago-Leccia, 1986 BO DA<br />
Rhabdolichops electrogrammus Lundberg & Mago-Leccia, 1986 BO DA BA RO<br />
Rhabdolichops jegui Keith & Meunier, 2000 FG<br />
Rhabdolichops stewarti Lundberg & Mago-Leccia, 1986 VA BO<br />
Rhabdolichops troscheli (Kaup, 1856) BO DA<br />
Rhabdolichops zareti Lundberg & Mago-Leccia, 1986 VA BO DA<br />
Sternopygus astrabes Mago-Leccia, 1994 CG? VA BO BA<br />
Sternopygus macrurus (Bloch & Schneider, 1801)<br />
Family: Rhamphichthyidae—Carl J. Ferraris, Jr.<br />
CG VA BO DA BA? RO? PA? AP? GU SU FG<br />
Gymnorhamphichthys hypostomus Ellis, 1912 CG VA DA<br />
Gymnorhamphichthys rondoni (Miranda Ribeiro, 1920) VA BO GU SU<br />
Iracema caiana Triques, 1996 BA RO<br />
Rhamphichthys apurensis (Fernández-Yépez, 1968) VA BO DA<br />
Rhamphichthys rostratus (Linnaeus, 1766)<br />
Family: Hypopomidae—James S. Albert<br />
GU SU<br />
Brachyhypopomus beebei (Schultz, 1944) CG VA BO DA GU SU<br />
Brachyhypopomus brevirostris (Steindachner, 1868) CG VA BO DA GU SU<br />
Brachyhypopomus pinnicaudatus (Hopkins, 1991) BO GU SU<br />
Hypopomus artedi (Kaup, 1856) PA? AP? GU SU FG<br />
Hypopygus lepturus Hoedeman, 1962 CG VA BO DA BA? SU FG<br />
Hypopygus neblinae Mago-Leccia, 1994 CG VA BO DA<br />
Microsternarchus bilineatus Fernández-Yépez, 1968 CG VA BO BA<br />
Racenisia fimbriipinna Mago-Leccia, 1994 VA BO<br />
Steatogenys duidae (La Monte, 1929) VA BO<br />
Steatogenys elegans (Steindachner, 1880)<br />
Family: Apteronotidae—James S. Albert<br />
CG VA BO DA<br />
Adontosternarchus clarkae Mago-Leccia, Lundberg & Baskin,<br />
1985<br />
CG VA BA RO<br />
Adontosternarchus devenanzii Mago-Leccia, Lundberg & Baskin,<br />
1985<br />
BO DA<br />
Adontosternarchus sachsi (Peters, 1877) BO DA<br />
Apteronotus albifrons (Linnaeus, 1766) VA BO DA GU SU FG<br />
Apteronotus leptorhynchus (Ellis, 1912) VA BO DA BA GU SU FG<br />
Compsaraia compsus (Mago-Leccia, 1994) BO DA<br />
Megadontognathus cuyuniense Mago-Leccia, 1994 BO<br />
Platyurosternarchus crypticus de Santana &Vari, 2009 RO GU<br />
Platyurosternarchus macrostomus (Günther, 1870) VA BO DA GU<br />
Porotergus gymnotus Ellis, 1912 GU FG<br />
Sternarchella orthos Mago-Leccia, 1994 VA BO DA<br />
Sternarchella sima Starks, 1913 BO DA<br />
Sternarchorhamphus muelleri (Steindachner, 1881) BO DA<br />
Sternarchorhynchus gnomus de Santana & Taphorn, 2006 BO<br />
Sternarchorhynchus oxyrhynchus (Müller & Troschel, 1849) BO GU FG<br />
Sternarchorhynchus roseni Mago-Leccia, 1994 BO DA
NUMBER 17 47<br />
Order: Cyprinodontiformes<br />
Family: Rivulidae—Wilson J. E. M. Costa<br />
Austr<strong>of</strong>undulus rupununi Hrbek, Taphorn &Thomerson, 2005 GU<br />
Austr<strong>of</strong>undulus transilis Myers, 1932 VA BO<br />
Gnatholebias hoignei (Thomerson, 1974) VA BO<br />
Gnatholebias zonatus (Myers, 1935) VA BO<br />
Kryptolebias sepia Vermeulen & Hrbek, 2005 SU<br />
Micromoema xiphophora (Thomerson & Taphorn, 1992) VA BO<br />
Rachovia maculipinnis (Radda, 1964) BO DA<br />
Rachovia stellifer (Thomerson & Turner, 1973) VA BO<br />
Renova oscari Thomerson & Taphorn, 1995 VA BO<br />
Rivulus agilae Hoedeman, 1954 GU SU FG<br />
Rivulus altivelis Huber, 1992 CG<br />
Rivulus amphoreus Huber, 1979 SU<br />
Rivulus breviceps Eigenmann, 1909 GU<br />
Rivulus caurae Radda, 2004 BO<br />
Rivulus cladophorus Huber, 1991 FG<br />
Rivulus corpulentus Thomerson & Taphorn, 1993 CG<br />
Rivulus deltaphilus Seegers, 1983 VA BO DA<br />
Rivulus frenatus Eigenmann, 1912 GU SU<br />
Rivulus gaucheri Keith, Nandrin & Le-Bail 2006 FG<br />
Rivulus geayi Vaillant, 1899 AP FG<br />
Rivulus gransabanae Lasso, Taphorn & Thomerson, 1992 BO<br />
Rivulus holmiae Eigenmann, 1909 GU<br />
Rivulus igneus Huber, 1991 AP FG<br />
Rivulus immaculatus Thomerson, Nico & Taphorn, 1991 BO<br />
Rivulus lanceolatus Eigenmann, 1909 GU<br />
Rivulus lungi Berkenkamp, 1984 FG<br />
Rivulus lyricauda Thomerson, Berkenkamp & Taphorn, 1991 VA BO<br />
Rivulus mahdiaensis Suijker & Collier, 2006 GU<br />
Rivulus manaensis Hoedeman, 1961 FG<br />
Rivulus mazaruni Myers, 1924 GU<br />
Rivulus nicoi Thomerson & Taphorn, 1992 VA BO<br />
Rivulus sape Lasso-Alcalá, Taphorn, Lasso & León-Mata, 2006 BO<br />
Rivulus stagnatus Eigenmann, 1909 GU SU<br />
Rivulus tecminae Thomerson, Nico & Taphorn, 1992 VA BO<br />
Rivulus torrenticola Vermeulen & Isbrücker, 2000 GU<br />
Rivulus waimacui Eigenmann, 1909 GU<br />
Rivulus xiphidius Huber, 1979 AP FG<br />
Terranatos dolichopterus (Weitzman & Wourms, 1967)<br />
Family: Poeciliidae—Paulo H. J. Lucinda<br />
VA BO<br />
Fluviphylax palikur Costa & Le Bail, 1999 AP FG<br />
Fluviphylax pygmaeus (Myers & Carvalho, 1955) CG VA BA PA<br />
Fluviphylax simplex Costa, 1996 PA<br />
Micropoecilia bifurca (Eigenmann, 1909) GU SU FG<br />
Micropoecilia parae (Eigenmann, 1894) AP? GU SU FG<br />
Micropoecilia picta (Regan, 1913) GU SU FG<br />
Poecilia reticulata Peters, 1859 BO DA AP? GU SU FG
48 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Poecilia vivipara Bloch & Schneider, 1801 DA GU SU FG<br />
Tomeurus gracilis Eigenmann, 1909 DA AP GU SU FG<br />
Order: Beloniformes<br />
Family: Belonidae—Bruce B. Collette<br />
Belonion dibranchodon Collette, 1966 VA BA<br />
Potamorrhaphis guianensis (Jardine, 1843) CG VA BA RO PA GU SU FG<br />
Potamorrhaphis petersi Collette, 1974 CG VA BO BA<br />
Pseudotylosurus microps (Günther, 1866)<br />
Family: Hemiramphidae—Bruce B. Collette<br />
VA BO DA GU SU<br />
Hyporhamphus brederi (Fernández-Yépez, 1948) VA BO DA<br />
Order: Synbranchiformes<br />
Family: Synbranchidae—Sven O. Kullander<br />
Synbranchus marmoratus Bloch, 1795 CG VA BO DA BA RO AP GU SU FG<br />
Order: Perciformes<br />
Family: Sciaenidae—Ning Labbish Chao & Lilian Casatti<br />
Pachypops fourcroi (Lacpède, 1802) VA BO DA AP GU SU FG<br />
Pachypops pigmaeus Casatti, 2002 RO<br />
Pachypops trifilis (Müller & Troschel, 1849) BA RO GU SU<br />
Pachyurus gabrielensis Casatti, 2001 VA BO BA<br />
Pachyurus schomburgkii Günther, 1860 VA BO<br />
Petilipinnis grunniens (Jardine, 1843) BO GU<br />
Plagioscion auratus (Castelnau, 1855) BO DA GU SU FG<br />
Plagioscion squamosissimus (Heckel, 1840) CG VA BO DA SU FG<br />
Plagioscion surinamensis (Bleeker, 1973)<br />
Family: Polycentridae—Richard P. Vari<br />
CG DA SU FG<br />
Monocirrhus polyacanthus Heckel, 1840 BO RO<br />
Polycentrus schomburgkii Müller & Troschel, 1849<br />
Family: Cichlidae—Sven O. Kullander<br />
DA AP GU SU FG<br />
Acaronia nassa (Heckel, 1840) RO AP GU FG<br />
Acaronia vultuosa Kullander, 1989 CG VA BO BA<br />
Aequidens chimantanus Inger, 1956 BO<br />
Aequidens diadema (Heckel, 1840) CG VA BO BA<br />
Aequidens paloemeuensis Kullander & Nijssen, 1989 SU<br />
Aequidens potaroensis Eigenmann, 1912 GU<br />
Aequidens tetramerus (Heckel, 1840) CG VA BO DA PA AP GU SU FG<br />
Aequidens tubicen Kullander & Ferreira, 1991 PA<br />
Apistogramma angayuara Kullander & Ferreira, 2005 PA<br />
Apistogramma diplotaenia Kullander, 1987 VA BO BA<br />
Apistogramma gibbiceps Meinken, 1969 RO<br />
Apistogramma gossei Kullander, 1982 AP FG<br />
Apistogramma hoignei Meinken, 1965 VA BO<br />
Apistogramma hongsloi Kullander, 1979 VA<br />
Apistogramma iniridae Kullander, 1979 CG<br />
Apistogramma inornata Staeck, 2003 BO<br />
Apistogramma ortmanni (Eigenmann, 1912) BO? GU SU<br />
Apistogramma rupununi Fowler, 1914 RO GU<br />
Apistogramma salpinction Kullander & Ferreira, 2005 PA<br />
Apistogramma steindachneri (Regan, 1908) GU SU
NUMBER 17 49<br />
Apistogramma velifera Staeck, 2003 VA<br />
Apistogramma viejita Kullander, 1979 CG<br />
Apistogramma wapisana Römer, Hahn & Conrad, 2006 RO<br />
Astronotus ocellatus (Agassiz, 1831) FG<br />
Biotodoma cupido (Heckel, 1840) GU<br />
Biotodoma wavrini (Gosse, 1963) CG VA BO BA<br />
Biotoecus dicentrarchus Kullander, 1989 CG VA BO DA<br />
Caquetaia spectabilis (Steindachner, 1875) BA RO AP GU<br />
Chaetobranchus flavescens Heckel, 1840 CG VA BO AP GU SU FG<br />
Cichla intermedia Machado-Allison, 1971 VA BO<br />
Cichla jariina Kullander & Ferreira, 2006 PA<br />
Cichla monoculus Spix & Agassiz, 1831 PA AP FG<br />
Cichla nigromaculata Jardine, 1843 VA<br />
Cichla ocellaris Schneider, 1801 RO GU SU FG<br />
Cichla orinocensis Humboldt, 1821 CG VA BO DA<br />
Cichla temensis Humboldt, 1821 CG VA BO DA BA<br />
Cichla thyrorus Kullander & Ferreira, 2006 PA<br />
Cichlasoma amazonarum Kullander, 1983 AP FG<br />
Cichlasoma bimaculatum (Linnaeus, 1758) BO RO GU SU FG<br />
Cichlasoma orinocense Kullander, 1983 CG VA BO<br />
Cleithracara maronii (Steindachner, 1881) DA GU SU FG<br />
Crenicara punctulatum (Günther, 1863) GU<br />
Crenicichla albopunctata Pellegrin, 1904 GU SU FG<br />
Crenicichla alta Eigenmann, 1912 RO GU<br />
Crenicichla coppenamensis Ploeg, 1987 SU<br />
Crenicichla heckeli Ploeg, 1989 PA<br />
Crenicichla hummelincki Ploeg, 1991 PA<br />
Crenicichla johanna Heckel, 1840 CG VA BO AP GU FG<br />
Crenicichla lugubris Heckel, 1840 BA RO GU SU<br />
Crenicichla multispinosa Pellegrin, 1903 SU FG<br />
Crenicichla nickeriensis Ploeg, 1987 SU<br />
Crenicichla pydanielae Ploeg, 1991 PA<br />
Crenicichla reticulata (Heckel, 1840) GU<br />
Crenicichla saxatilis (Linnaeus, 1758) DA GU SU FG<br />
Crenicichla sipaliwini Ploeg, 1987 SU<br />
Crenicichla ternetzi Norman, 1926 FG<br />
Crenicichla tigrina Ploeg, Jégu & Ferreira, 1991 PA<br />
Crenicichla vaillanti Pellegrin, 1903 GU FG<br />
Crenicichla virgatula Ploeg, 1991 RO<br />
Crenicichla wallacii Regan, 1905 GU<br />
Crenicichla zebrina Montaña, López-Fernández & Taphorn,<br />
2008<br />
VA<br />
Dicrossus filamentosus (Ladiges, 1958) CG VA BO<br />
Dicrossus gladicauda Schindler & Staeck, 2008 CG<br />
Geophagus abalios López-Fernández & Taphorn, 2004 VA BO<br />
Geophagus brachybranchus Kullander & Nijssen, 1989 GU SU<br />
Geophagus brokopondo Kullander & Nijssen, 1989 SU<br />
Geophagus camopiensis Pellegrin, 1903 AP FG<br />
Geophagus dicrozoster López-Fernández & Taphorn, 2004 VA BO
50 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Geophagus gottwaldi Schindler & Staeck, 2006 VA<br />
Geophagus grammepareius Kullander & Taphorn, 1992 VA BO<br />
Geophagus harreri Gosse, 1976 SU FG<br />
Geophagus surinamensis (Bloch, 1791) SU FG<br />
Geophagus taeniopareius Kullander & Royero, 1992 VA BO<br />
Geophagus winemilleri López-Fernández & Taphorn, 2004 VA<br />
Guianacara (Guianacara) cuyunii López-Fernández, Taphorn, &<br />
Kullander, 2006<br />
BO<br />
Guianacara (Guianacara) geayi (Pellegrin, 1902) AP FG<br />
Guianacara (Guianacara) owroewefi Kullander & Nijssen, 1989 SU FG<br />
Guianacara (Guianacara) sphenozona Kullander & Nijssen, 1989 GU SU<br />
Guianacara (Guianacara) stergiosi López-Fernández, Taphorn,<br />
& Kullander, 2006<br />
BO<br />
Guianacara (Oelemaria) oelemariensis Kullander & Nijssen,<br />
1989<br />
SU<br />
Heros severus Heckel, 1840 CG VA BO BA<br />
Hoplarchus psittacus (Heckel, 1840) CG VA BO BA<br />
Hypselecara coryphaenoides (Heckel, 1840) CG VA BO BA<br />
Hypselecara temporalis (Günther, 1862) AP<br />
Ivanacara adoketa (Kullander & Prada-Pedreros, 1993) BA<br />
Ivanacara bimaculata Eigenmann, 1912 GU<br />
Krobia guianensis (Regan, 1905) GU SU<br />
Krobia itanyi (Puyo, 1943) SU FG<br />
Laetacara fulvipinnis Staeck & Schindler, 2007 CG VA BA<br />
Mazarunia mazarunii Kullander, 1990 GU<br />
Mesonauta egregius Kullander & Silfvergrip, 1991 CG<br />
Mesonauta guyanae Schindler, 1998 RO GU<br />
Mesonauta insignis (Heckel, 1840) CG VA BO BA<br />
Nannacara anomala Regan, 1905 GU SU<br />
Nannacara aureocephalus Allgayer, 1983 FG<br />
Pterophyllum altum Pellegrin, 1903 CG VA<br />
Pterophyllum leopoldi (Gosse, 1963) GU<br />
Pterophyllum scalare (Schultze, 1823) BA AP GU SU FG<br />
Retroculus septentrionalis Gosse, 1971 AP FG<br />
Satanoperca acuticeps (Heckel, 1840) RO<br />
Satanoperca daemon (Heckel, 1840) CG VA BO BA<br />
Satanoperca jurupari (Heckel, 1840) BA PA AP<br />
Satanoperca leucosticta (Müller & Troschel, 1849) GU SU<br />
Satanoperca lilith Kullander & Ferreira, 1988 BA RO PA<br />
Satanoperca mapiritensis (Fernández-Yépez, 1950) VA BO DA<br />
Uaru fernandezyepezi Stawikowski, 1989<br />
Family: Gobiidae—Sven O. Kullander<br />
VA<br />
Awaous flavus Valenciennes, 1837 DA AP GU SU FG<br />
Ctenogobius claytonii (Meek, 1902) DA<br />
Dormitator maculatus (Bloch, 1792) GU SU FG<br />
Eleotris amplyopsis (Cope, 1871) DA GU SU FG<br />
Eleotris pisonis (Gmelin, 1789) DA GU SU FG<br />
Gobioides broussonnetii La Cepède, 1800 DA FG<br />
Gobioides grahamae (Palmer & Wheeler, 1995) GU SU FG
NUMBER 17 51<br />
Gobiomorus dormitor La Cepède, 1800 GU SU<br />
Microphilypnus ternetzi Myers, 1927 VA BO DA<br />
Order: Pleuronectiformes<br />
Family: Achiridae—Robson T. C. Ramos<br />
Achirus achirus (Linnaeus, 1758) GU FG<br />
Achirus novoae Cervigón, 1982 BO DA<br />
Apionichthys dumerili Kaup, 1858 DA PA AP GU SU FG<br />
Apionichthys finis Eigenmann, 1912 RO GU<br />
Apionichthys menezesi Ramos, 2003 VA BO<br />
Hypoclinemus mentalis (Günther, 1862) VA BO BA RO AP GU<br />
Order: Tetraodontiformes<br />
Family: Tetraodontidae—Carl J. Ferraris, Jr. & Sven O.<br />
Kullander<br />
Colomesus asellus (Müller & Troschel, 1849) BO DA GU<br />
Order: Lepidosireniformes<br />
Family: Lepidosirenidae—Based on Cl<strong>of</strong>fsca account (Gloria<br />
Arratia)<br />
Lepidosiren paradoxa Fitzinger, 1837 CG? VA? BA? FG
PHOTOGRAPHIC ATLAS OF FISHES OF THE GUIANA SHIELD<br />
Introduction<br />
The last decade or so has witnessed a surge in<br />
expeditions to both ichthyologically familiar and virgin<br />
waters in southeastern Venezuela, Guyana, Suriname, and<br />
French Guiana. Included are surveys <strong>of</strong> the Iwokrama<br />
Forest in west-central Guyana (Watkins et al. 2005),<br />
retracing Carl Eigenmann’s 1908 collecting route up the<br />
Essequibo to the Potaro River above Kaiteur Falls<br />
(Hardman et al. 2002), and rapid assessments targeting<br />
species-rich waters such as the upper Essequibo Basin,<br />
Guyana (Lasso et al. 2008), Coppename Basin, Suriname<br />
(Berrenstein 2005, Alonso & Berrenstein 2006, and<br />
references therein), and Venezuelan states <strong>of</strong> Amazonas<br />
(Lasso et al. 2006, and references therein), and Bolívar<br />
(Machado-Allison et al. 2003). Systematic fish inventories<br />
<strong>of</strong> French Guiana began over 50 years ago (see references<br />
in Vari & Ferraris, this volume), and have been recently<br />
expanded by French and Swiss ichthyologists to include<br />
ecological (e.g., Lord et al. 2007) and molecular data, the<br />
latter to investigate the origins <strong>of</strong> the Guianas’ highly<br />
diversified fish fauna (Cardoso & Montoya-Burgos 2009).<br />
Explorations <strong>of</strong> remote Shield regions in search <strong>of</strong><br />
undescribed catfishes (Sabaj Pérez et al. 2009) have<br />
assembled a parade <strong>of</strong> new taxa led by the sucker-mouth<br />
armored siluriforms in the family Loricariidae. Fifteen<br />
new loricariid species from Guyana, Suriname, and<br />
Amazonas, Venezuela, have been described in the last<br />
five years (e.g., Werneke et al. 2005, Armbruster et al.<br />
2007, de Chambrier & Montoya-Burgos 2008, Lujan et al.<br />
2009) with many more discoveries awaiting description.<br />
This impressive amount <strong>of</strong> fieldwork has significantly<br />
advanced our taxonomic understanding <strong>of</strong> fishes in the<br />
Guianas; nevertheless, much must still be accomplished.<br />
Expeditions to remote, previously unsampled waters,<br />
particularly headwater systems above waterfalls or large<br />
cataracts, routinely yield new and sometimes enigmatic<br />
ichthy<strong>of</strong>aunas (Taphorn et al. 2008; Lujan, pers. comm.;<br />
pers. obs.). More comprehensive collecting efforts (e.g.,<br />
night sampling) in relatively well-sampled waters have<br />
uncovered new species that escaped prior efforts (e.g.,<br />
Armbruster et al. 2000; pers. obs.). Fieldwork aside,<br />
there exists in museums a wealth <strong>of</strong> specimens <strong>of</strong><br />
Guianas fishes that require critical evaluation. The rich<br />
and complex diversity <strong>of</strong> fishes in the Guianas, and their<br />
systematic placement in the greater context <strong>of</strong> the<br />
Neotropical fauna, will remain a lodestone for ichthyological<br />
studies in decades to come.<br />
Scope<br />
The plates present 130 individuals representing 127<br />
species <strong>of</strong> 46 families. Fishes were collected in Guyana<br />
MARK H. SABAJ PÉREZ<br />
(53 species), Suriname (36) and Amazonas State,<br />
Venezuela (38) from 1985 to 2008. Most <strong>of</strong> the species<br />
occur on or immediately peripheral to the Guiana<br />
Shield, with a few species restricted to lowland, coastal<br />
habitats in fresh and/or estuarine waters (i.e., Rhinosardinia<br />
amazonica, Sciades parkeri, Tomeurus gracilis,<br />
Anableps anableps, Polycentrus schomburgkii).<br />
Fishes were imaged live or shortly after death (89<br />
species), or from specimens purchased at market (2),<br />
preserved in formalin (2), or stored in alcohol (34).<br />
Each image is identified in the plate description by<br />
taxon, condition <strong>of</strong> specimen at time <strong>of</strong> photo, museum<br />
and catalog number, size and sex (if so determined),<br />
current status <strong>of</strong> voucher if other than preserved whole<br />
in alcohol, and complete locality data. Depositories are<br />
The Academy <strong>of</strong> Natural Sciences, Philadelphia<br />
(ANSP), Auburn University Natural History Museum<br />
(AUM), Field Museum <strong>of</strong> Natural History (FMNH),<br />
Illinois Natural History Survey (INHS), Museo de<br />
Ciencias Naturales de la UNELLEZ, Guanare<br />
(MCNG), National Zoological Collection <strong>of</strong> Suriname<br />
(NZCS), and University <strong>of</strong> Guyana, Center for the<br />
Study <strong>of</strong> <strong>Biological</strong> Diversity (UG/CSBD). Photos are<br />
by author unless credited otherwise. Abbreviations in<br />
the text are: LEA – length to end <strong>of</strong> anal fin; SL –<br />
standard length; and TL – total length. Scale bars are<br />
presented only for those species in which that indicator<br />
was included in the original photograph.<br />
Fish Photography<br />
There is a variety <strong>of</strong> techniques for capturing highquality<br />
color images <strong>of</strong> fishes, all <strong>of</strong> which have been<br />
vastly simplified and in many ways improved by the<br />
advent <strong>of</strong> digital technology. Most <strong>of</strong> the images<br />
presented here are <strong>of</strong> live (or recently so) and alcohol<br />
preserved specimens immersed in water in a glass<br />
phototank. Materials and methods are largely the same<br />
whether taken streamside <strong>of</strong> live specimens (Figs. 3, 4) or<br />
in the lab <strong>of</strong> preserved specimens (Fig. 5), except for the<br />
light source: ambient sunlight in the field vs. incandescent<br />
light in-doors. Other photographers have used electronic<br />
flashes (e.g., Jenkins & Burkhead 1994:129, Planquette et<br />
al. 1996:17) to produce stunning photos <strong>of</strong> live fishes in<br />
phototanks. I have not tried such techniques, but consider<br />
a cooperative sun to be equally effective and in some ways<br />
less burdensome. In any event, phototank-immersion<br />
remains the gold standard for ex-situ fish photography.<br />
Phototank-immersion Method<br />
This method involves three stages: equipment set up,<br />
specimen preparation, and image capture and editing.
54 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Figure 3. Author using phototank-immersion method to photograph fish streamside in Mongolia. Photo by C. Sabaj Pérez.<br />
The techniques described below follow a minimalist<br />
approach with some advice limited to the specific<br />
cameras and conditions involved. For a more sophisticated<br />
system and additional tips on fish photography<br />
see Jenkins & Burkhead (1994:127–130).<br />
Equipment set up.—The phototank is made <strong>of</strong><br />
ordinary plate glass bonded together with clear silicone<br />
adhesive. Outside dimensions (in inches) <strong>of</strong> the tanks<br />
used for the photos in this section are: 13.5 length 3<br />
10.25 height 3 2.75 width (field and lab) and 15.75<br />
length 3 12.25 height 3 3.5 width (lab only). Both are<br />
made from one-quarter inch thick glass, except oneeighth<br />
inch glass is used for the front plate <strong>of</strong> smaller<br />
tank. These dimensions are well suited for lateral and<br />
<strong>of</strong>ten dorsal/ventral views <strong>of</strong> small to medium-sized<br />
fishes up to about 300 mm total length and 63 mm<br />
width for smaller tank, and 370 mm total length and 75<br />
mm width for larger. Two important factors compromise<br />
field utility <strong>of</strong> larger phototanks: the volume <strong>of</strong><br />
water necessary to fill it and size <strong>of</strong> carrying case (see<br />
below). Each tank requires a separate glass plate to<br />
immobilize the subject. The free plate can be oneeighth<br />
(smaller tank) or one-quarter (larger) inch thick<br />
and is slightly shorter and deeper than the inside<br />
dimensions <strong>of</strong> the tanks (e.g., 13 3 10.25 and 15 3 12<br />
inches for smaller and larger tank, respectively).<br />
Having smooth edges <strong>of</strong> all plates is recommended.<br />
The tank should be filled with clear bottled or<br />
filtered/deionized tap water to minimize formation <strong>of</strong><br />
air bubbles on specimen and glass. Stream or lake<br />
water is unsuitable because it lacks the desired clarity<br />
and suspended debris is a significant distraction in an<br />
otherwise good photo. Any water will accumulate<br />
debris over an extended photo session, and an ample<br />
supply <strong>of</strong> clean photo water must accompany long<br />
forays to remote locations.<br />
In the lab the phototank is stationed between two<br />
pairs <strong>of</strong> incandescent bulbs positioned to the side and<br />
slightly above the top <strong>of</strong> the tank (Fig. 5). Polarizing<br />
filters are useful for reducing glare or overexposed hot<br />
spots on the specimen, particularly on the snout. When<br />
using sunlight, the tank is oriented to maximize the<br />
even distribution <strong>of</strong> light and minimize glare and<br />
shadows on the subject.<br />
Selection <strong>of</strong> a camera is important, but the rapid<br />
pace <strong>of</strong> digital technology soon outdistances specific<br />
recommendations on make or model. By current<br />
standards a digital camera with a good optical zoom<br />
(6X and higher) that records images at or above<br />
resolutions <strong>of</strong> 12 Megapixels (MP) is generally a safe
NUMBER 17 55<br />
Figure 4. Author photographing fish streamside in Guyana. Photo by J.W. Armbruster.<br />
choice. Most <strong>of</strong> the photos herein were taken with a<br />
Nikon Coolpix E8700 (8 MP); others with this model’s<br />
predecessors, the E4500 (4 MP) and older E995 (3.1<br />
MP). The most recent photos, all <strong>of</strong> alcohol preserved<br />
specimens, were taken in the lab with a Nikon D90 D-<br />
SLR (12.3 MP) fitted with a 60 mm f/2.8G micro lens.<br />
Images taken with the E8700 contain a high level <strong>of</strong><br />
sharp detail that is slightly exceeded by the D90 (or<br />
other cameras <strong>of</strong>fering greater MP), particularly for<br />
small specimens. The differences, however, are only<br />
visible at high magnification or extremely large print<br />
sizes. The greatest advantage <strong>of</strong> the D-SLR design and<br />
micro lens is the enhanced ability to reliably focus on<br />
very small specimens. Any camera and lens should be<br />
thoroughly vetted by comparing published reviews<br />
(many available on-line), and then personally tested<br />
with the phototank-immersion method. A few digital<br />
cameras apparently have difficulties rendering a sharp<br />
specimen image through glass and water.<br />
Additional essentials for basic set up are a tripod<br />
(mini-tripods are handy in the field; Figs. 3–5), 4-ply<br />
mat board in several background colors (e.g., flat<br />
black, dull light blue) and 3/16 th inch foam board with<br />
flat black surface for camera blind (sizes <strong>of</strong> all boards<br />
ideally fitted to carrying case), glass cleaner, and paper<br />
towels or lint-free cloth, both long and small forceps,<br />
large metal binder clips, 12-inch plastic metal rulers,<br />
stiff wire, an assortment <strong>of</strong> needles and insect pins,<br />
calipers, a system for tagging individual specimens<br />
(e.g., dymo-tags in pre-punched number series tied to<br />
strong twine), extra camera batteries and charger,<br />
memory cards and reader, and laptop computer for<br />
image storage. These essentials are best stored with the<br />
phototank in a crushpro<strong>of</strong> and watertight carrying<br />
case. The smaller tank is ideal for field use as it requires<br />
less water and allows for co-storage <strong>of</strong> accessories and<br />
laptop in a small case suitable for carry-on luggage (see<br />
Fig. 3). Cameras are better stored separately to<br />
facilitate other uses and avoid residual moisture in<br />
the phototank.<br />
Specimen preparation.—The overarching strategy<br />
when photographing fishes for identification purposes<br />
is to maximize the content and accuracy <strong>of</strong> information<br />
in the image. This aim determines which among<br />
multiple specimens is photographed, how it is illuminated<br />
and arranged for display, and which color<br />
background is used. Most striking are photographs <strong>of</strong><br />
the most impressive specimens (i.e., in peak coloration<br />
and with fins and scales intact), but even the image <strong>of</strong><br />
an impressive fish may be rendered less informative if<br />
the photograph is poorly composed.<br />
Once a live or alcohol specimen is selected it is<br />
carefully inspected and cleaned <strong>of</strong> foreign debris.<br />
Mucous-laden skin and fins <strong>of</strong>ten attract distracting grit<br />
or other suspended particles, and cheesecloth fibers may<br />
adhere to preserved specimens. An anesthetized fish (e.g.,<br />
with a few drops <strong>of</strong> clove oil) is quickly euthanized in a<br />
container <strong>of</strong> strong (30–50%) formalin. This <strong>of</strong>ten causes<br />
the body to straighten and fins to become completely<br />
erect. Otherwise the anesthetized specimen may be
56 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Figure 5. Kyle Luckenbill photographing small alcohol-preserved specimen (above ruler) while holding polarizing filter in lab. Photo<br />
by author.<br />
removed to a tray <strong>of</strong> shallow formalin wherein small<br />
forceps are carefully used to hold the fins erect without<br />
damaging them. The most important consideration<br />
when photographing live specimens is time; bright colors<br />
and iridescences are soon lost in formalin. Fatty skin, as<br />
in pseudopimelodid catfishes, also becomes opaque in<br />
formalin, obscuring any underlying color.<br />
Once the specimen is flat with fins erect, it is carefully<br />
wedged between the front plate <strong>of</strong> the phototank<br />
and free plate <strong>of</strong> glass, the latter set at an angle and<br />
braced against metal binder clips either attached to the<br />
sides <strong>of</strong> the tank or loosely set between the free plate<br />
and back <strong>of</strong> tank (Fig. 3). Positioning laterally compressed<br />
fishes in this manner is easy. Dorsoventrally<br />
depressed specimens, particularly those with pectoral<br />
spines, require more attention to achieve a vertical<br />
lateral view. Maintaining pectoral spines folded against<br />
the body as one wedges the specimen between the two<br />
glass plates requires practice and patience. Long<br />
forceps, a metal ruler and stiff wire are useful tools<br />
for fine-tuning a specimen’s posture, arranging long<br />
delicate features such as barbels, and dislodging air<br />
bubbles that form on the fish. Information content <strong>of</strong> a<br />
fish photo is diminished when the specimen is tilted or<br />
otherwise poorly positioned.<br />
Preserved specimens <strong>of</strong>fer fewer options for achieving<br />
an ideal photo-friendly posture. Laterally contorted<br />
specimens <strong>of</strong>ten can be made to appear more linear<br />
when tightly wedged between the two plates <strong>of</strong> glass.<br />
Issues that are more difficult arise with partial or<br />
complete folding <strong>of</strong> fins. In some cases insect pins<br />
(carefully inserted in the body opposite the side to be<br />
imaged) may be used to prop up the anterior most<br />
portions <strong>of</strong> fins. This technique, however, may cause<br />
small tears in the fin membranes.<br />
Next is selection <strong>of</strong> an appropriate background.<br />
Many specimens, particularly dark ones with opaque<br />
fins, <strong>of</strong>ten render best with more dramatic effect<br />
against flat black backgrounds. This may pose a<br />
serious drawback for specimens with relatively transparent<br />
fins. Black pigment in fin membranes or along<br />
distal fin margins disappears against dark backgrounds.<br />
In such cases, a light blue background<br />
provides better contrast and will highlight dark<br />
pigmentation in fins. Conversely, transparent fins<br />
lacking pigmentation and with clear margins, particularly<br />
in live specimens, are <strong>of</strong>ten lost against light<br />
backgrounds. This can be alleviated to a certain degree<br />
by adjusting the tank relative to light source to achieve<br />
a small measure <strong>of</strong> direct side or back lighting. While it<br />
is true that graphics editing s<strong>of</strong>tware (e.g., Vertus Fluid<br />
Mask) can virtually affect any color background,<br />
specimens may not appear natural if the new background<br />
deviates sharply from the original (i.e., black to
NUMBER 17 57<br />
white and vice versa). Choice <strong>of</strong> background color<br />
<strong>of</strong>ten involves trade-<strong>of</strong>fs, and is ultimately a reflection<br />
<strong>of</strong> personal taste determined via trial and error.<br />
The final step is placement <strong>of</strong> a scale bar. This is<br />
accomplished by cutting out a 10+ mm portion <strong>of</strong> a<br />
plastic ruler, dipping it in water and adhering it to the<br />
outside front <strong>of</strong> the phototank beneath the specimen<br />
and within the photographic field.<br />
Image capture and editing.—The camera is mounted<br />
on a tripod, as most exposures are too long to permit<br />
hand-held use, and positioned behind a black foam<br />
board with central circular aperture fitted to lens. The<br />
blind prevents the phototank glass from reflecting the<br />
images <strong>of</strong> camera and photographer. Whether horizontal<br />
or angled the specimen should occupy about<br />
90% <strong>of</strong> the length <strong>of</strong> the digital image recorded. To<br />
preserve detail in extremely long and slender fishes<br />
(e.g., belonids; Pl. 14, Fig. E) the specimen is imaged in<br />
two aligned and overlapping parts (anterior and<br />
posterior halves) that are digitally combined. The<br />
shutter is placed on a timer delay and white balance<br />
set appropriately (e.g., sunlight vs. incandescent).<br />
Digital photography frees one from limits imposed<br />
by the amount <strong>of</strong> available film and developing costs.<br />
In the field, particularly while the sun is dodging<br />
clouds, it is advisable to take multiple photos for<br />
each <strong>of</strong> several combinations <strong>of</strong> exposures and apertures<br />
(f-stops). Full sunlight <strong>of</strong>ten highlights fine<br />
structures (e.g., odontodes in loricariids), but at the<br />
same time may wash out bright colors or result in<br />
overexposed hot spots on the snout or dorsum. The<br />
phototank should be carefully oriented with respect<br />
to the light source, and extra mat boards used to<br />
shadow harsh sunlight and maintain vibrant colors<br />
(Fig. 3).<br />
For the Coolpix E8700 in manual mode, the shutter<br />
speed is set such that the target aperture (i.e., lower<br />
f-stops) lies between f-stops 5 and 7; larger apertures<br />
reduce depth <strong>of</strong> field, and smaller apertures tend to<br />
reduce resolution. The Nikon D90 D-SLR better<br />
accommodates smaller apertures (f-stop fixed at 16<br />
with ISO set to 200), and the shutter speed is manually<br />
adjusted for the best exposure. Aut<strong>of</strong>ocus generally<br />
works fine as long as the active area <strong>of</strong> focus includes<br />
important features on the fish, not the scale bar or<br />
background. Digital cameras typically have a setting<br />
whereby the user determines the active area <strong>of</strong><br />
aut<strong>of</strong>ocus. Depending on specimen size, the camera<br />
may need to be manually set to macro mode, and some<br />
cameras (e.g., Nikon CoolPix) also require one to<br />
slightly zoom in on subject for sharp aut<strong>of</strong>ocus. Nikon<br />
images presented here are <strong>of</strong> Fine quality (recorded as<br />
JPEGs with compression ratio <strong>of</strong> roughly 1:4) and<br />
maximum size (3264 3 2448 and 4288 3 2848 pixels for<br />
E8700 and D90, respectively). Higher quality settings<br />
record either uncompressed TIFF or RAW (NEF)<br />
images, the latter requiring extra s<strong>of</strong>tware and com-<br />
puting time for conversion to TIFF files (Nikon D90<br />
allows one to record NEF and Fine JPEG images<br />
concurrently). TIFF and RAW files retain the full<br />
quality <strong>of</strong> the image and the latter maximizes allowable<br />
post exposure processing, whereas JPEGs are compressed<br />
<strong>of</strong>ten with some visual quality permanently lost<br />
in the process (the loss, however, is barely perceptible).<br />
Larger image files (NEF, RAW, TIFF) do <strong>of</strong>fer<br />
slightly higher resolution, but the improvement is <strong>of</strong>ten<br />
negligible, except at high magnification. For any<br />
camera, there is no substitute for testing a variety <strong>of</strong><br />
settings and image qualities to optimize the desired<br />
effect and protocol.<br />
While photographing a specimen it is difficult to<br />
know which image will optimize the desired effect; so,<br />
it is best to have ample images from which to choose.<br />
The number <strong>of</strong> images I generally take is directly<br />
proportional to the impressive and unique nature <strong>of</strong><br />
the specimen added to the amount <strong>of</strong> time expended to<br />
pose it properly in the phototank. It is easy to<br />
accumulate many photos <strong>of</strong> numerous species, thus it<br />
is critical to have a system for later identification and<br />
management <strong>of</strong> images. Failure to do so guarantees<br />
extra time and <strong>of</strong>ten frustration when attempting to<br />
match images to specimens long after capture. The best<br />
field solution is to take a final photo <strong>of</strong> the specimen<br />
together with a uniquely numbered tag that is then<br />
secured to the fish. In the case <strong>of</strong> museum specimens,<br />
the jar label is photographed immediately after imaging<br />
the fish. A photo-log is useful for recording the<br />
standard length <strong>of</strong> the specimen. Such practices greatly<br />
facilitate subsequent annotation <strong>of</strong> images with catalog<br />
and measurement data. A new and much welcomed<br />
trend in digital cameras is a built in or accessory global<br />
positioning system (GPS) receiver that records and<br />
embeds latitude, longitude, altitude and universal time<br />
as image metadata.<br />
The final step is image editing, all <strong>of</strong> which was<br />
performed on the photos in this section using Adobe<br />
Photoshop. This program <strong>of</strong>fers a seemingly endless<br />
myriad <strong>of</strong> simple to advanced tools for graphics<br />
manipulation. Only a few <strong>of</strong> the more basic tools and<br />
techniques are mentioned briefly here.<br />
Once an image is selected the background (original)<br />
layer is immediately duplicated and subsequent edits<br />
are made to the duplicate layer. A third blank layer is<br />
added to mask the specimen with a uniform background.<br />
Masking color (e.g., solid black or white, or a<br />
color shade taken from the original background using<br />
the eyedropper tool) is first added as a rough outline<br />
using a large diameter pencil tool, and then completed<br />
with a fine-tipped brush (1–10 pixels) under magnification<br />
(e.g., $300%) to carefully trace the specimen’s<br />
precise contours. The magic wand and/or magnetic<br />
lasso are more expedient, yet less precise, tools for<br />
masking the specimen with a uniform background.<br />
Next, the duplicate layer is automatically and manually
58 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
adjusted for levels (tonal range and color balance),<br />
brightness/contrast, and hue/saturation. The auto<br />
options <strong>of</strong>ten render extreme values that are manually<br />
faded to desired opacity before additional manual finetuning.<br />
The cloning stamp tool is useful for removing<br />
small bubbles or debris on the specimen, while the<br />
dodge and burn tools help lighten or darken localized<br />
regions (brush size/shape and exposure/opacity <strong>of</strong> such<br />
tools are manually adjusted). Under the Filter menu<br />
‘Sharpen.Unsharp Mask’ can sharpen an image with<br />
s<strong>of</strong>t focus, and ‘Noise.Despeckle’ removes graininess,<br />
particularly for images scanned from 35 mm slides.<br />
One final trick is to render a solid scale bar by: 1)<br />
rotating the entire image so the ruler piece in the<br />
photograph is horizontal, 2) using the rectangular<br />
marquee tool to select and copy a 5 or 10 mm long<br />
portion, 3) rotating the entire image back to its final<br />
intended position, 4) pasting the copied selection,<br />
thereby creating a new layer, and 5) adjusting the<br />
brightness/contrast <strong>of</strong> this layer to extreme values to<br />
render a black or white bar that is then labeled<br />
accordingly with the text tool.<br />
The final edited version <strong>of</strong> the specimen image (i.e.,<br />
duplicate layer) can be quickly compared to the<br />
original by using the layers window and clicking the<br />
‘eye’ icon to hide or display the corresponding layer.<br />
Creation <strong>of</strong> additional layers requires the new image to<br />
be saved as an uncompressed TIFF file that is suitable<br />
for archiving and any additional post processing. A<br />
copy <strong>of</strong> the final edited image is flattened to a single<br />
layer and resaved as a TIFF for print publication (with<br />
Image.Mode set to Grayscale, RGB, or CMYK color<br />
based upon printer specifications) and separately as a<br />
JPEG (with Mode set to 8 bits/channel) for use in<br />
presentations or easy transmission and accession via<br />
the Internet.<br />
In sum, a high-quality fish photo is the product<br />
<strong>of</strong> preparation, practice, and patience all committed<br />
with keen attention to detail. Additional factors in<br />
the field are perseverance under suboptimal conditions<br />
and a bit <strong>of</strong> luck with respect to weather and finding<br />
the ideal specimen. The amount and quality <strong>of</strong> the<br />
images presented herein had more to do with will than<br />
skill.<br />
Acknowledgments<br />
Numerous colleagues contributed valuable assistance<br />
both before and after the shutter click. For<br />
assistance in the field I wish to thank in particular:<br />
Mariangeles Arce, Jonathan Armbruster, Michael<br />
Hardman, Oscar J. León-Mata, Nathan Lujan, John<br />
Lundberg, Jan Mol, Matthew Thomas, David Werneke,<br />
and Philip Willink. For loans <strong>of</strong> specimens I<br />
thank Michael Retzer, and for help tracking down<br />
specimens and taking measurements I thank Osvaldo<br />
Oyakawa and especially Nathan Lujan. For help<br />
cleaning backgrounds I thank Kyle Luckenbill and<br />
Rebecca Meyer. Special thanks to Pierre-Yves le Bail,<br />
Raphaël Covain and Juan Montoya Burgos for<br />
comments on the introduction, and to Jonathan<br />
Armbruster, Hernán López-Fernández, Nathan Lujan,<br />
Donald Stewart, and Donald Taphorn for graciously<br />
contributing their images to this atlas. Finally I thank<br />
my ichthyological friends for years <strong>of</strong> help identifying<br />
many <strong>of</strong> the fishes presented here. Support for this<br />
project was received in part from the All Catfish<br />
Species Inventory (NSF DEB-0315963).<br />
Literature Cited<br />
Alonso, L. E., & H. J. Berrenstein (eds.). 2006. A rapid biological<br />
assessment <strong>of</strong> the aquatic ecosystems <strong>of</strong> the Coppename River<br />
Basin, Suriname. RAP Bulletin <strong>of</strong> <strong>Biological</strong> Assessment 39.<br />
Conservation International, <strong>Washington</strong>, D.C., 119 pp.<br />
Armbruster, J. W., N. K. Lujan, & D. C. Taphorn. 2007. Four new<br />
Hypancistrus (Siluriformes: Loricariidae) from Amazonas,<br />
Venezuela.—Copeia 2007(1):62–79.<br />
———, M. H. Sabaj, M. Hardman, L. M. Page, & J. H. Knouft.<br />
2000. Catfish genus Corymbophanes (Loricariidae: Hypostominae)<br />
with description <strong>of</strong> one new species: Corymbophanes<br />
kaiei.—Copeia 2000(4):997–1006.<br />
Berrenstein, H. J. 2005. Field guide to the freshwater fishes <strong>of</strong> the<br />
Central Suriname Nature Reserve (CSNR), (Coppename<br />
River Basin, Suriname). Conservation International Suriname,<br />
Paramaribo, 96 pp.<br />
Cardoso, Y. P., & J. I. Montoya-Burgos. 2009. Unexpected diversity<br />
in the catfish Pseudancistrus brevispinis reveals dispersal routes<br />
in a Neotropical center <strong>of</strong> endemism: the Guyanas Region.—<br />
Molecular Ecology 18(5):947–964.<br />
de Chambrier, S., & J. I. Montoya-Burgos. 2008. Pseudancistrus<br />
corantijniensis, a new species from the Guyana Shield<br />
(Siluriformes: Loricariidae) with a molecular and morphological<br />
description <strong>of</strong> the Pseudancistrus barbatus group.—<br />
Zootaxa 1918:45–58.<br />
Hardman, M., L. M. Page, M. H. Sabaj, J. W. Armbruster, & J. H.<br />
Knouft. 2002. A comparison <strong>of</strong> fish surveys made in 1908 and<br />
1998 <strong>of</strong> the Potaro, Essequibo, Demerara, and coastal river<br />
drainages <strong>of</strong> Guyana.—Ichthyological Exploration <strong>of</strong> Freshwaters<br />
13:225–238.<br />
Jenkins, R. E., & N. M. Burkhead. 1994. Freshwater fishes <strong>of</strong><br />
Virginia. American Fisheries <strong>Society</strong>, Bethesda, Maryland,<br />
1080 pp.<br />
Lasso, C. A., J. C. Señarìs, L. E. Alonso, & A. L. Flores (eds.). 2006.<br />
Evaluación rápida de la biodiversidad de los ecosistemas<br />
acuáticos en la confluencia de los ríos Orinoco y Ventuari,<br />
Estado Amazonas (Venezuela). Boletín RAP de Evaluación<br />
Biológica 30. Conservation International, <strong>Washington</strong>, D.C.,<br />
240 pp.<br />
Lasso, C. A., J. Hernández-Acevedo, E. Alexander, J. C. Señaris, L.<br />
Mesa, H. Samudio, J. Mora-Day, C. Magalhaes, A. Shushu,<br />
E. Mauruwanaru, & R. Shoni. 2008. Aquatic biota: fishes,<br />
decapod crustaceans and mollusks <strong>of</strong> the upper Essequibo<br />
Basin (Konashen COCA), Southern Guyana. Pp. 43–54 in<br />
L. E. Alonso, J. McCullough, P. Naskrecki, E. Alexander, &<br />
H. E. Wright, eds., A rapid biological assessment <strong>of</strong> the<br />
Konashen Community-Owned Conservation Area, Southern<br />
Guyana. RAP Bulletin <strong>of</strong> <strong>Biological</strong> Assessment 51, Conservation<br />
International, <strong>Washington</strong>, D.C.<br />
Lord, C., Y. Fermon, F. J. Meunier, M. Jégu, & P. Keith. 2007.<br />
Croissance et longévité du Watau yaike, Tometes lebaili
NUMBER 17 59<br />
(Serrasalminae), dans le bassin du Haut Maroni (Guyane<br />
française). Résultats préliminaires.—Cybium, 31(3):359–367.<br />
Lujan, N. K., M. Arce, & J. W. Armbruster. 2009. A new black<br />
Baryancistrus with blue sheen from the upper Orinoco<br />
(Siluriformes: Loricariidae).—Copeia 2009(1):50–56.<br />
Machado-Allison, A., B. Chern<strong>of</strong>f, F. Provenzano, P. W. Willink, A.<br />
Marcano, P. Petry, B. Sidlauskas, & T. Jones. 2003.<br />
Inventory, relative abundance and importance <strong>of</strong> fishes in<br />
the Caura River basin, Bolívar State, Venezuela. Pp. 64–74 in<br />
B. Chern<strong>of</strong>f, A. Machado-Allison, K. Riseng, & J. R.<br />
Montambault, eds., A biological assessment <strong>of</strong> the aquatic<br />
Ecosystems <strong>of</strong> the Caura River Basin, Bolívar State,<br />
Venezuela. RAP Bulletin <strong>of</strong> <strong>Biological</strong> Assessment 28.<br />
Conservation International, <strong>Washington</strong> D.C. 284 pp.<br />
Planquette, P., P. Keith, & P.-Y. Le Bail. 1996. Atlas des poissons<br />
d’eau douce de Guyane. Volume 1, Collection du Patrimoine<br />
Naturel, vol. 22. Institut d’Ecologie et de Gestion de la<br />
Biodiversité du Muséum National d’Histoire Naturelle,<br />
Institut national de la Recherche Agronomique, Conseil<br />
Supérior de la Pêche, Paris, 429 pp.<br />
Sabaj Pérez, M. H, J. W. Armbruster, C. J. Ferraris, Jr., J. P. Friel, J.<br />
G. Lundberg, & L. M. Page (eds.). 2009. All Catfish Species<br />
Inventory. Internet address: http://silurus.acnatsci.org.<br />
Taphorn, B. D. C, H. López-Fernández, & C. R. Bernard. 2008.<br />
Apareiodon agmatos, a new species from the upper Mazaruni<br />
river, Guyana (Teleostei: Characiformes: Parodontidae).—<br />
Zootaxa 1925:31–38.<br />
Watkins, G., W. Saul, E. Holm, C. Watson, D. Arjoon, & J. Bicknell.<br />
2005. The fish fauna <strong>of</strong> the Iwokrama Forest.—Proceedings <strong>of</strong><br />
the Academy <strong>of</strong> Natural Sciences <strong>of</strong> Philadelphia 154(1):39–53.<br />
Werneke, D. C., M. H. Sabaj, N. K. Lujan, & J. W. Armbruster. 2005.<br />
Baryancistrus demantoides and Hemiancistrus subviridis,twonew<br />
uniquely colored species <strong>of</strong> catfishes from Venezuela (Siluriformes:<br />
Loricariidae).—Neotropical Ichthyology 3(4):533–542.
APPENDIX: PLATES
62 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Potamotrygonidae<br />
Plate 1<br />
A. Paratrygon aiereba (live). AUM 43646 (154 mm maximum disk width). Venezuela,<br />
Amazonas, Río Negro (Amazonas drainage), left bank sandy beach and small adjacent<br />
backwater 7.2 km NW <strong>of</strong> San Carlos de Rio Negro, 01u589110N, 067u069100W, 19 Mar<br />
2005, M. Sabaj, D. Werneke et al.<br />
B. Potamotrygon orbignyi (live). AUM 43201 (171 mm maximum disk width). Venezuela,<br />
Amazonas, Río Orinoco ca. 60 km E <strong>of</strong> San Fernando de Atabapo, 03u589260N,<br />
067u099460W, 3 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
C. Potamotrygon marinae (live). ANSP 187098 (400 mm maximum disk width). Suriname,<br />
Sipaliwini, Lawa River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan<br />
(airstrip), 03u199310N, 054u039480W, 18 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P.<br />
Willink, & K. Wan.<br />
D. Potamotrygon schroederi (live). AUM 44507 (423 mm maximum disk width). Venezuela,<br />
Amazonas, Río Orinoco, island W <strong>of</strong> Puerto Venado, 4.5 km S <strong>of</strong> Samariapo, 56.5 km<br />
SW <strong>of</strong> Puerto Ayacucho, 05u129250N, 067u489320W, 28 Feb 2005, M. Sabaj, N. Lujan, D.<br />
Werneke et al.
NUMBER 17 63
64 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Arapaimidae<br />
Plate 2<br />
A. Arapaima sp. (live). UG/CSBD uncataloged (174 cm SL, male, skeleton). Guyana, Grass<br />
Pond, Rewa River basin (Rupununi drainage), near Rewa village, Aug 2006, D. Stewart et<br />
al. Photo by D. J. Stewart.<br />
Osteoglossidae<br />
B. Osteoglossum bicirrhosum (live). No voucher (ca. 500 mm TL). Guyana, Crane Pond,<br />
southwestern part <strong>of</strong> Rupununi basin near Karanambu Ranch, 2007, D. Stewart et al.<br />
Photo by D. J. Stewart.
NUMBER 17 65
66 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Clupeidae<br />
Plate 3<br />
A. Rhinosardinia amazonica (alcohol). INHS 49009 (33.7 mm SL). Guyana, East Demerara,<br />
Demerara River (Atlantic drainage), Land <strong>of</strong> Canaan, 11.8 mi S-SW <strong>of</strong> Georgetown at<br />
bearing 213u, 06u389390N, 058u119460W, 14 Oct 1998, M. Sabaj, J. Armbruster, M.<br />
Hardman et al.<br />
Pristigasteridae<br />
B. Pellona castelnaeana (live). MCNG 51957. Venezuela, Amazonas, Río Casiquiare at<br />
mouth <strong>of</strong> Caño Caripo, 37 km W-SW <strong>of</strong> La Esmeralda, 03u069500N, 065u529380W, 5 Mar<br />
2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
Engraulidae<br />
C. Amazonsprattus scintilla (live). ANSP 181134. Guyana, Pirara River (Ireng-Takutu-<br />
Branco drainage), 3.5 km N-NW <strong>of</strong> Pirara, 03u389550N, 059u419200W, 2 Nov 2002, M.<br />
Sabaj, J. Armbruster, M. Thomas et al.<br />
D. Anchovia surinamensis (alcohol). ANSP 189252 (69.7 mm SL). Guyana, Rupununi River<br />
(Essequibo drainage), at Massara’s Landing, 1.1 km NE village <strong>of</strong> Massara, 03u539410N,<br />
059u179370W, 26 Oct 2002. M. Sabaj, J. Armbruster, M. Thomas et al.<br />
E. Lycengraulis batesii (alcohol). ANSP 189251 (52.4 mm SL). Guyana, Cuyuni-Mazaruni,<br />
Mazaruni River (Essequibo drainage) long partially exposed sandy shoal between two<br />
islands in main channel, 6.9 km SW <strong>of</strong> Bartica, 06u229470N, 058u409320W, 12 Nov 2002,<br />
M. Sabaj, J. Armbruster, M. Thomas et al.<br />
F. Anchoviella sp. (alcohol). ANSP 189234 (29.5 mm SL). Suriname, Sipalawini, Lawa River<br />
(Marowini drainage), small sand beach below cataract upstream from base camp, ca. 9 km<br />
S-SW <strong>of</strong> Anapaike. 03u199120N, 054u039410W, 19–23 Apr 2007, J. Lundberg, J. Mol, M.<br />
Sabaj, P. Willink, & K. Wan.<br />
Parodontidae<br />
G. Apareiodon agmatos (alcohol). ROM 83755 (52.6 mm SL). Guyana, Mazaruni River<br />
(Essequibo drainage), sandy beach and embayment on right bank, upstream from village<br />
<strong>of</strong> Jawalla, 05u41935.40N, 060u28911.80 W, 18 Apr 2008, H. López-Fernández, D.<br />
Taphorn, E. Liverpool, K. Kramer, & C. Thierens. Photo by D. C. Taphorn Baechle & H.<br />
López-Fernández.<br />
H. Apareiodon orinocensis (live). ANSP 185045. Venezuela, Amazonas, Río Orinoco at<br />
Puerto Venado, 4.3 km S <strong>of</strong> Samariapo, 56.4 km SW <strong>of</strong> Puerto Ayacucho, 05u129380N,<br />
067u489180W, 26 Feb 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
I. Parodon guyanensis (live). ANSP 189204 (ca. 70 mm SL). Suriname, Sipaliwini, Lawa River<br />
(Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip), 03u199310N,<br />
054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.<br />
J. Parodon guyanensis (alcohol). ANSP 189204 (73 mm SL). Same locality as I.
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68 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Curimatidae<br />
Plate 4<br />
A. Curimatopsis crypticus (live). ANSP 189091 (32.4 mm SL, female). Suriname, Para,<br />
Coropinae Creek (Suriname drainage), vicinity <strong>of</strong> Republiek, 05u299570N, 055u129520W,<br />
28 Apr 2007, M. Sabaj & P. Willink.<br />
B. Curimatopsis crypticus (live). ANSP 189091 (28.8 mm SL, male). Same locality as A.<br />
Prochilodontidae<br />
C. Prochilodus rubrotaeniatus (alcohol). ANSP 175495 (160 mm SL). Guyana, Essequibo<br />
River, sandbar ca. 800 m downstream <strong>of</strong> Essequibo (Maipuri) campsite, 04u459430N,<br />
058u459520W, 29 Jan 1997, W. Saul et al.<br />
D. Semaprochilodus varii (live). ANSP 187435 (210 mm SL). Suriname, Sipaliwini, Lawa<br />
River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
Crenuchidae<br />
E. Leptocharacidium omospilus (live). ANSP 189272 (60.5 mm SL). Venezuela, Amazonas, 1<br />
km upstream from mouth <strong>of</strong> left bank tributary <strong>of</strong> Río Siapa, mouth below Salto Oso and<br />
above Salto Sardinas on Río Siapa, 01u269240N, 065u409010W, 14 Mar 2005, M. Sabaj, N.<br />
Lujan, M. Arce, & T. Wesley.<br />
F. Poecilocharax bovalii (alcohol). INHS 49600 (28.6 mm SL). Guyana, first N bank creek<br />
tributary <strong>of</strong> Potaro River (Essequibo drainage) downstream from Waratuk Cataract, 27<br />
Oct 1998, L. Page, M. Sabaj, J. Armbruster et al.<br />
Hemiodontidae<br />
G. Argonectes longiceps (alcohol). ANSP 189151 (184 mm SL). Same locality data as D.<br />
H. Bivibranchia bimaculata (live). ANSP 189149 (111 mm SL). Same locality data as D.<br />
Gasteropelecidae<br />
I. Carnegiella strigata (alcohol). INHS 49173 (26 mm SL). Guyana, Mazaruni–Potaro,<br />
‘‘Himarakus’’ Creek, tributary <strong>of</strong> Essequibo River (Atlantic drainage) at Rockstone,<br />
05u599080N, 058u339030W, 19 Oct 1998, M. Sabaj, J. Armbruster, M. Hardman.<br />
J. Gasteropelecus sternicla (alcohol). ANSP 189193 (46.6 mm SL). Same locality as A.
NUMBER 17 69
70 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Anostomidae<br />
Plate 5<br />
A. Anostomus brevior (live). FMNH ex ANSP 189141 (51.6 mm SL). Suriname, Sipaliwini,<br />
Lawa River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
B. Anostomus anostomus (live). ANSP 180172 (72.5 mm SL). Guyana, Essequibo River<br />
(Atlantic drainage) at Yukanopito Falls, 44.5 km SW <strong>of</strong> mouth <strong>of</strong> Kuyuwini River,<br />
01u549530N, 058u319140W, 9 Nov 2003, M. H. Sabaj, J. Armbruster, N. Lujan et al.<br />
C. Synaptolaemus cingulatus (live). AUM 43269 (78.8 mm SL). Venezuela, Amazonas, Río<br />
Orinoco, 147 km SE <strong>of</strong> San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar<br />
2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.<br />
D. Leporellus vittatus (formalin). ANSP 189270 (photo voucher). Guyana, Rupununi River<br />
(Essequibo drainage), road crossing 5.9 km W-SW <strong>of</strong> village <strong>of</strong> Sand Creek, 02u579000N,<br />
059u349100W, 4 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
E. Leporinus fasciatus (live). ANSP 189158 (81.7 mm SL). Same locality as A.<br />
F. Leporinus maculatus (live). FMNH ex ANSP 189041 (123.4 mm SL). Same locality as A.<br />
G. Hypomasticus megalepis (live). AUM 37999 (68.3 mm SL). Guyana, Essequibo River at<br />
Kassi-Attae Rapids, 5.5 km SE <strong>of</strong> mouth <strong>of</strong> Kuyuwini River, 02u139360N, 058u179380W, 8<br />
Nov 2003, M. Sabaj, J. Armbruster, M. Hardman et al.<br />
H. Leporinus ortomaculatus (live). AUM 43262 (72.9 mm SL). Same locality as C.<br />
I. Leporinus lebaili (live). FMNH ex ANSP 189043 (56.7 mm SL). Same locality as A.<br />
J. Caenotropus maculosus (live). ANSP 189147 (70 mm SL). Suriname, Sipalawini, Litanie<br />
River, side channel behind Theo’s bakery, just upstream <strong>of</strong> confluence with Marowini<br />
River and village <strong>of</strong> Konya Kondre, 03u179240N, 054u049380W, 23 Apr 2007, J. Lundberg,<br />
J. Mol, M. Sabaj, P. Willink, & K. Wan.
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Characidae<br />
Plate 6<br />
A. Brittanichthys myersi (formalin). INHS 49053 (22.4 mm SL). Guyana, East Demerara,<br />
Maduni River and Conservancy Canal (Mahaica-Atlantic drainage), Maduni Stop-<strong>of</strong>f,<br />
22.3 mi S <strong>of</strong> Georgetown bearing 176u,06u309010N, 58u029140W, 15 Oct 1998, M. Sabaj, J.<br />
Armbruster, M. Hardman et al.<br />
B. Bryconops melanurus (live). ANSP 189268 (75.5 mm SL). Suriname, Sipalawini, Litanie<br />
River at mouth and confluence with Marowini River, just upstream from settlement <strong>of</strong><br />
Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P.<br />
Willink, J. Mol et al.<br />
C. Chalceus epakros (live). AUM 43073 (75 mm SL). Venezuela, Amazonas, Río Orinoco,<br />
inlet between 2 islands in channel, 84.5 km N <strong>of</strong> San Fernando de Atabapo, 9 km S<strong>of</strong><br />
Monduapo, 1 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
D. Exodon paradoxus (live). AUM 36843 (61 mm SL). Guyana, Rupununi River (Essequibo<br />
drainage), sand beach and inlet at Karanambo Ranch, 03u459000N, 059u189300W, 30 Oct<br />
2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
E. Myleus rubripinnis (live). ANSP 189267 (34 mm SL). Guyana, Suriname, Sipaliwini, Lawa<br />
River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18-22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
F. Myleus schomburgkii (live). ANSP 180812 (160 mm SL). Venezuela, Amazonas, Río<br />
Orinoco, bedrock outcrop, 52.9 km SE <strong>of</strong> San Antonio, 102 km W <strong>of</strong> La Esmerelda,<br />
03u069010N, 066u279460W, 4 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
G. Myloplus cf. planquettei (live). ANSP 179808 (425 mm SL). Guyana, Essequibo River<br />
(Atlantic drainage) at Yukanopito Falls, 44.5 km SW <strong>of</strong> mouth <strong>of</strong> Kuyuwini River,<br />
01u549530N, 058u319140W, 9 Nov 2003, M. H. Sabaj, J. Armbruster, N. Lujan et al.<br />
H. Serrasalmus rhombeus (live). ANSP 180287 (ca. 260 mm SL). Venezuela, Amazonas, Río<br />
Ventuari (Orinoco drainage), large rock outcrop, 97 km NE <strong>of</strong> Macuruco, 163 km E-NE<br />
<strong>of</strong> San Fernando de Atabapo, 7 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.
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Acestrorhynchidae<br />
Plate 7<br />
A. Acestrorhynchus falcatus (live). INHS 48983 (153.9 mm SL). Guyana, East Demerara,<br />
Madewini River (Demerara drainage), 21.5 mi S-SW <strong>of</strong> Georgetown, bearing 207u, at<br />
Linden highway bridge, 06u 309 05.00 N, 58u 129 44.90 W, 14 Oct 1998, M. Sabaj, J.<br />
Armbruster, M. Hardman et al.<br />
B. Acestrorhynchus microlepis (live). AUM 36753 (123 mm SL). Guyana, Circle West Creek,<br />
tributary <strong>of</strong> Pirara River (Ireng-Takutu drainage), 26.6 km SW <strong>of</strong> Karanambo Ranch,<br />
03u399140N, 059u319430W, 30 Oct 2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
Cynodontidae<br />
C. Cynodon meionactis (alcohol). ANSP 189129 (190 mm SL). Suriname, Sipalawini, Litanie<br />
River at mouth and confluence with Marowini River, just upstream from settlement <strong>of</strong><br />
Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P.<br />
Willink, J. Mol et al.<br />
D. Hydrolycus armatus (live). MCNG 51983 (ca. 400 mm SL). Venezuela, Amazonas, Rio<br />
Orinoco (Atlantic drainage), Pasaganado, 38 km N <strong>of</strong> San Fernando de Atabapo,<br />
04u239040N, 067u469280W, 1 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
Lebiasinidae<br />
E. Copella compta (live). AUM 41327 (37 mm SL). Venezuela, Amazonas, Caño Carmen,<br />
tributary <strong>of</strong> Río Orinoco, 1.5 km S-SE <strong>of</strong> Manaka, 70 km E <strong>of</strong> San Fernando de Atabapo,<br />
4 Apr 2004, N. Lujan & D. Werneke.<br />
F. Copella cf. arnoldi (live). ANSP 189192 (30.4 mm SL). Suriname, Para, Coropinae Creek<br />
(Suriname drainage), vicinity <strong>of</strong> Republiek, 05u299570N, 055u129520W, 28 Apr 2007, M.<br />
Sabaj & P. Willink.<br />
Erythrinidae<br />
G. Hoplias aimara (alcohol). ANSP 176723 (172 mm SL). Guyana, Tumble Down Creek,<br />
tributary <strong>of</strong> Siparuni River (Essequibo drainage), 04u489390N, 058u519110W, 8 Dec 1997,<br />
G. Watkins et al.<br />
H. Hoplias lacerdae group (live). AUM 44674 (152 mm SL). Guyana, Pirara River (Ireng-<br />
Takutu drainage), at Pirara Ranch, 03u379310, 059u409360, 26 Nov 2005, N. Lujan, D.<br />
Taphorn et al. Photo by N. K. Lujan.<br />
I. Erythrinus erythrinus (alcohol). ANSP 175537 (121 mm SL). Guyana, Culvert creek<br />
(Essequibo drainage) crossing Kurupukari-Surama River road, 04u199570N, 058u519130W,<br />
5 Feb 1997, W. Saul et al.<br />
Ctenoluciidae<br />
J. Boulengerella cuvieri (live). AUM 40987 (560 mm SL). Venezuela, Amazonas, Río Ventuari<br />
(Orinoco drainage), large rock outcrop, 97 km NE <strong>of</strong> Macuruco, 163 km E-NE <strong>of</strong> San Fernando<br />
de Atabapo, 04u259100N, 066u179080W,7Apr2004,M.Sabaj,N.Lujan,D.Wernekeetal.
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Cetopsidae<br />
Plate 8<br />
A. Cetopsis cf. montana (live). ANSP 178782 (78.3 mm SL). Guyana, Ireng River (Takutu-<br />
Branco-Negro drainage), 6.9 km W-SW <strong>of</strong> village <strong>of</strong> Karasabai, 04u019100N,<br />
059u369060W, 1 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
B. Helogenes marmoratus (live). AUM 27960 (56 mm SL). Guyana, West Demerara,<br />
Catabuly Creek (Demerara-Atlantic drainage), at Wismar, 1.87 mi. N-NW <strong>of</strong> Linden<br />
bearing 335u, 06u01937.20N, 58u19925.30W, 18 Oct 1998, L. Page, M. Sabaj, J. Armbruster<br />
et al. Photo by J. W. Armbruster.<br />
Aspredinidae<br />
C. Ernstichthys sp. (alcohol). ANSP 180002 (26.4 mm SL). Same locality as A.<br />
D. Bunocephalus verrucosus (alcohol). ANSP 180015 (36.6 mm SL). Guyana, Hassar Pond<br />
and outlet (Rupununi drainage), 5.4 km S-SE <strong>of</strong> Massara, 03u509400N, 059u179090W, 27<br />
Oct 2002, J. Armbruster, D. Werneke, C. Allison et al.<br />
Ariidae<br />
E. Sciades parkeri (live). ANSP 178741 (445 mm SL). Guyana, purchased at Georgetown fish<br />
market, 8–15 Nov 2002, M. Sabaj.<br />
Auchenipteridae<br />
F. Ageneiosus inermis (live). FMNH ex ANSP 187115 (ca. 200 mm SL). Suriname,<br />
Sipalawini, Litanie River at mouth and confluence with Marowini River, just upstream<br />
from settlement <strong>of</strong> Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg,<br />
M. Sabaj, P. Willink, J. Mol et al.<br />
G. Auchenipterichthys punctatus (live). AUM 43416 (131 mm SL). Venezuela, Amazonas, Río<br />
Casiquiare, bedrock riffle and outcrop, 74.6 km NE <strong>of</strong> San Carlos de Rio Negro,<br />
02u219460N, 066u339530W, 9 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
H. Trachycorystes trachycorystes (live). AUM 35933 (111 mm SL). Guyana, unnamed stream<br />
(Rupununi drainage) at crossing on road between Massara and Karanambo, 10.3 km NW<br />
<strong>of</strong> Karanambo Ranch, 03u489270N, 059u239060W, 28 Oct 2002, M. Sabaj, J. Armbruster,<br />
M. Thomas et al.<br />
I. Gelanoglanis sp. (live). AUM 35908 or ANSP 178738 (male). Guyana, Ireng River<br />
(Takutu-Branco drainage), 6.9 km WSW village <strong>of</strong> Karasabai, 04u019100N, 059u369060W, 1<br />
Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
J. Glanidium leopardum (live). ANSP 189104 (48.5 mm SL). Same locality as F.
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Doradidae<br />
Plate 9<br />
A. Acanthodoras cataphractus (alcohol). NZCS 1618–1619 (84.5 mm SL). Suriname, Para,<br />
tributary <strong>of</strong> lower Suriname River at Republiek, 14 Nov 1989, P. Outboter et al.<br />
B. Scorpiodoras heckelii (live). ANSP 182790 (82.5 mm SL, specimen with simple secondary<br />
gas bladder). Venezuela, Amazonas, Río Orinoco at Puerto Samariapo, 05u159N,<br />
067u489W, 25 Feb 2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.<br />
C. Leptodoras copei (live). ANSP 182225 (85.8 mm SL). Venezuela, Amazonas, Río Ventuari<br />
(Orinoco drainage), beach across river from Picua village, 34 km E-NE <strong>of</strong> Macuruco, 104<br />
km E <strong>of</strong> San Fernando de Atabapo, 04u069550N, 066u459520W, 5 Apr 2004, M. Sabaj, N.<br />
Lujan, D. Werneke et al.<br />
D. Leptodoras linnelli (live). AUM 41038 (171 mm SL). Venezuela, Amazonas, Río Ventuari<br />
(Orinoco drainage), village <strong>of</strong> Marueta at landing, 91 km<br />
E-NE <strong>of</strong> Macuruco, 159 km E-NE <strong>of</strong> San Fernando de Atabapo, 04u189510N,<br />
066u179320W, 6 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.<br />
E. Centrodoras hasemani (live). ANSP 182227 (211 mm SL). Venezuela, Amazonas, Río<br />
Casiquiare, bedrock outcrop 59.5 km SW <strong>of</strong> La Esmerelda, 02u499070N, 065u579190W, 8<br />
Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
F. Oxydoras niger (live). AUM 35508 (340 mm SL). Pirara River, tributary <strong>of</strong> Ireng River<br />
(Takutu-Branco drainage), beach at Pirara Ranch on road to Lethem, 03u379170N,<br />
059u409290W, 2 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
G. Doras micropoeus (alcohol). ANSP 187110 (225 mm SL). Suriname, Sipaliwini, Lawa<br />
River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
H. Hassar orestis (live). ANSP 180294 (161 mm SL). Venezuela, Amazonas, Río Ventuari<br />
(Orinoco drainage), beach at village <strong>of</strong> Moriche, 116 km NE <strong>of</strong> Macuruco, 169 km NE <strong>of</strong><br />
San Fernando de Atabapo, 04u459N, 066u219130W, 7 Apr 2004, M. Sabaj, N. Lujan, D.<br />
Werneke et al.<br />
I. Opsodoras morei (live). ANSP 182836 (157 mm SL). Venezuela, Amazonas, Río Orinoco<br />
near confluence with Río Atabapo, long narrow beach between channel and laguna, San<br />
Fernando de Atabapo, 04u029480N, 067u429170W, 2 Apr 2004, M. Sabaj, N. Lujan, D.<br />
Werneke et al.<br />
J. Rhinodoras armbrusteri (live). ANSP 179096 (66.1 mm SL). Guyana, Takutu River (Rio<br />
Branco-Negro drainage), ca. 2.75 km W <strong>of</strong> Saint Ignatius, 03u219180N, 059u499510W, 5-6<br />
Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
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Pseudopimelodidae<br />
Plate 10<br />
A. Pseudopimelodus bufonius (live). ANSP 189098 (87.4 mm SL). Suriname, Sipalawini,<br />
Litanie River at mouth and confluence with Marowini River, just upstream from<br />
settlement <strong>of</strong> Konya Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M.<br />
Sabaj, P. Willink, J. Mol et al.<br />
Heptapteridae<br />
B. Mastiglanis sp. (live). FMNH ex ANSP 189106 (43.2 mm SL). Same locality as A.<br />
C. Leptorhamdia sp. (live). AUM 43261 (51 mm SL). Venezuela, Amazonas, Río Orinoco,<br />
147 km SE <strong>of</strong> San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar 2005, M.<br />
Sabaj, N. Lujan, M. Arce, & T. Wesley.<br />
D. Pimelodella geryi (live). ANSP 189109 (114 mm SL). Suriname, Sipaliwini, Lawa River<br />
(Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip), 03u199310N,<br />
054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.<br />
Pimelodidae<br />
E. Hypophthalmus marginatus (market). ANSP 187103 (330 mm SL). Suriname, Paramaribo,<br />
purchased at main fish market in Paramaribo, presumably from vicinity in Surinam River,<br />
17 Apr 2007, J. Lundberg, M. Sabaj, & P. Willink.<br />
F. Megalonema platycephalum (live). AUM 36018 (97.2 mm SL). Guyana, Rupununi River<br />
(Essequibo drainage), sand beach and inlet at Karanambo Ranch, 03u459000N,<br />
059u189300W, 30 Oct 2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
G. Brachyplatystoma filamentosum (live). ANSP 187070 (540 mm SL). Venezuela, Amazonas,<br />
Río Casiquiare, bedrock outcrop 59.5 km SW <strong>of</strong> La Esmerelda, 02u499070N,<br />
065u579190W, 8 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
H. Pseudoplatystoma fasciatum (market). ANSP 187106 (ca. 640 mm SL, skeleton). Same locality<br />
as E.<br />
I. Pimelodus ornatus (live). ANSP 187113 (169 mm SL). Same locality as D.<br />
J. Pinirampus pirinampu (live). AUM 37964. Guyana, Essequibo River, along beach 12.9 km<br />
SE (upstream) <strong>of</strong> mouth <strong>of</strong> Kuyuwini River, 02u099430, 058u169350, 10 Nov 2003, M.<br />
Sabaj, J. Armbruster, M. Hardman et al.
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Callichthyidae<br />
Plate 11<br />
A. Callichthys callichthys (live). ANSP 179110 (68.2 mm SL). Guyana, Orokang River<br />
(Mazaruni drainage), 1.2 km S <strong>of</strong> Chi Chi Falls airstrip, 21.1 km S-SW <strong>of</strong> Imbaimadai,<br />
05u319310N, 060u139560W, 13 Nov 2002, J. Armbruster, M. Sabaj, M. Thomas et al.<br />
B. Megalechis thoracata (alcohol). ANSP 179795 (62 mm SL). Guyana, Hassar Pond and<br />
outlet (Rupununi drainage), 5.4 km S-SE <strong>of</strong> Massara, 03u509400N, 059u179090W, 27 Oct<br />
2002, J. Armbruster, D. Werneke, C. Allison et al.<br />
Trichomycteridae<br />
C. Ituglanis cf. metae (live). AUM 43074 (61 mm SL). Venezuela, Amazonas, Río Orinoco,<br />
inlet between two islands in channel, 84.5 km N <strong>of</strong> San Fernando de Atabapo, 9 km S <strong>of</strong><br />
Monduapo, 04u479540N, 067u499160W, 1 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et<br />
al.<br />
D. Trichomycterus hasemani (alcohol, top 5 lateral view, bottom 5 dorsal view). ANSP<br />
179154 (13.5 mm SL). Guyana, Rupununi River (Essequibo drainage), 3.7 km S-SE <strong>of</strong><br />
village <strong>of</strong> Massara, 03u519440N, 059u179040W, 27 Oct 2002, M. Sabaj, J. Armbruster, M.<br />
Thomas et al.<br />
E. Henonemus taxistigmus (alcohol). ANSP 179953 (90.6 mm SL). Guyana, Rupununi River<br />
(Essequibo drainage) at Kwatamang, 4 km SE <strong>of</strong> Annai, 03u559030N, 059u069010W, 25 Oct<br />
2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
F. Trichomycterus guianensis (alcohol). ANSP 179109 (52.5 mm SL). Guyana, Orokang<br />
River (Mazaruni drainage), at Chi Chi Falls airstrip, 20.1 km S-SW <strong>of</strong> Imbaimadai,<br />
05u329060N, 060u139590W, 13 Nov 2002, J. Armbruster, M. Sabaj, M. Thomas et al.<br />
G. Typhlobelus sp. (alcohol, dorsal view). AUM 35802 (22.3 mm SL). Guyana, Ireng River<br />
(Takutu-Branco-Negro drainage), 6.9 km W-SW <strong>of</strong> village <strong>of</strong> Karasabai, 04u019100N,<br />
059u369060W, 1 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.<br />
H. Typhlobelus sp. (alcohol, left 5 dorsal view, right 5 ventral view <strong>of</strong> head). Same data as<br />
G.<br />
I. Pygidianops sp. (live). ANSP 179820 (23.1 mm SL). Guyana, Takutu River (Branco-Negro<br />
drainage), 3.77 km S-SW <strong>of</strong> Lethem, 03u219180N, 059u499510W, 16 Nov 2003, M. Sabaj, J.<br />
Armbruster, M. Hardman et al.<br />
J. Sarcoglanis simplex (live). ANSP 179212 (17 mm SL). Same locality as G.
NUMBER 17 83
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Loricariidae<br />
Plate 12<br />
A. Leporacanthicus triactis (live). AUM 39243 (116 mm SL). Venezuela, Amazonas, Río<br />
Ventuari (Orinoco drainage), 23 km NE <strong>of</strong> Macaruco, 94 km E <strong>of</strong> San Fernando de<br />
Atabapo, 04u049500N, 066u519540W, 5 Apr 2004, N. Lujan, D. Werneke, M. Sabaj et al.<br />
B. Leporacanthicus cf. galaxias (live). AUM 39226 (93 mm SL). Venezuela, Amazonas, Río<br />
Orinoco at Cucue Amerindian village, opposite Trapichote village, 60 km E <strong>of</strong> San<br />
Fernando de Atabapo, 03u589260N, 067u099300W, 3 Apr 2004, N. Lujan, D. Werneke, M.<br />
Sabaj et al.<br />
C. Hemiancistrus medians (live). ANSP 187122 (156 mm SL). Suriname, Sipaliwini, Lawa<br />
River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
D. Peckoltia sabaji (live). ANSP 185094 (109.7 mm SL). Guyana, Takutu River (Branco-<br />
Negro drainage), 3.77 km S-SW <strong>of</strong> Lethem, 03u219180N, 059u499510W, 1 Nov 2003, M.<br />
Sabaj, J. Armbruster, M. Hardman et al.<br />
E. Peckoltia braueri (live). AUM 38882 (103 mm SL). Same locality as D.<br />
F. Lithoxus stocki (live). ANSP 189131 (66.5 mm SL). Suriname, Sipalawini, Litanie River at<br />
mouth and confluence with Marowini River, just upstream from settlement <strong>of</strong> Konya<br />
Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P. Willink, J.<br />
Mol et al.<br />
G. Pseudolithoxus dumus (live). AUM 43267 (87 mm SL). Venezuela, Amazonas, Río<br />
Orinoco, 147 km SE <strong>of</strong> San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar<br />
2005, M. Sabaj, N. Lujan, M. Arce, & T. Wesley.<br />
H. Pseudancistrus pectegenitor (alcohol). MCNG 54797 (241.6 mm SL). Venezuela,<br />
Amazonas, Río Casiquiare, bedrock in stream, 73 km NE <strong>of</strong> San Carlos de Río Negro,<br />
02u219090N, 066u349310W, 9 Mar 2005, N. Lujan, M. Sabaj, D. Werneke et al.<br />
I. Metaloricaria paucidens (live). ANSP 187325 (146.5 mm SL). Suriname, Sipaliwini, Lawa<br />
River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
J. Rhadinoloricaria macromystax (live). ANSP 182349 (100 mm SL). Guyana, Ireng River<br />
(Takutu-Branco-Negro drainage), 6.9 km W-SW <strong>of</strong> village <strong>of</strong> Karasabai, 04u019100N,<br />
059u369060W, 1 Nov 2002, M. Sabaj, J. Armbruster, M. Thomas et al.
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Gymnotidae<br />
Plate 13<br />
A. Electrophorus electricus (live). MCNG 51982. Venezuela, Amazonas, Río Orinoco, 147 km<br />
SE <strong>of</strong> San Fernando de Atabapo, 03u189240N, 066u369120W, 4 Mar 2005, M. Sabaj, N.<br />
Lujan, M. Arce, & T. Wesley.<br />
Sternopygidae<br />
B. Sternopygus sp. (live). ANSP 189018 (146 mm TL). Suriname, Sipaliwini, Lawa River<br />
(Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip), 03u199310N,<br />
054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, & K. Wan.<br />
Rhamphichthyidae<br />
C. Gymnorhamphichthys hypostomus (alcohol). INHS 49454 (161 mm LEA). Guyana, Potaro<br />
River (Essequibo drainage), beach on N bank, downstream <strong>of</strong> Tumatumari Cataract,<br />
05u21948.40N, 59u00904.40W, 22 Oct 1998, M. Sabaj, J. Armbruster, & M. Hardman.<br />
Hypopomidae<br />
D. Hypopomus artedi (alcohol). ANSP 189266 (152 mm TL). Suriname, Sipaliwini, Lawa<br />
River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
Apteronotidae<br />
E. Apteronotus albifrons (live). AUM 40678 (109 mm SL). Venezuela, Amazonas, Río<br />
Manapiare (Ventuari-Orinoco drainage), at San Juan de Manapiare landing, 15 Apr 2004,<br />
J. Para.
NUMBER 17 87
88 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Rivulidae<br />
Plate 14<br />
A. Rivulus waimacui (alcohol). INHS 49635 (55.5 mm SL). Guyana, tributary <strong>of</strong> Potaro River<br />
(Essequibo drainage) near Tukeit Cataract, 05u12916.50N, 059u279090W, 27-28 Oct 1998,<br />
L. Page, J. Armbruster, M. Hardman et al.<br />
Poeciliidae<br />
B. Tomeurus gracilis (alcohol). INHS 49017 (27 mm SL). Guyana, East Demerara, Demerara<br />
River (Atlantic drainage), Land <strong>of</strong> Canaan, 11.8 mi S-SW <strong>of</strong> Georgetown bearing 213u,<br />
06u389390N, 058u119460W, 14 Oct 1998, M. Sabaj, J. Armbruster, M. Hardman et al.<br />
Anablepidae<br />
C. Anableps anableps (alcohol). INHS 49016 (140 mm SL). Same locality as B.<br />
Belonidae<br />
D. Potamorrhaphis guianensis (live). ANSP 179480 (228 mm SL). Guyana, Yuora River<br />
(Ireng-Takutu-Branco drainage), 6.7 km NE <strong>of</strong> village <strong>of</strong> Karasabai on road to Tiger<br />
Creek village, 04u039140N, 059u299070W, 31 Oct 2002, M. Sabaj, J. Armbruster, M.<br />
Thomas et al.<br />
E. Potamorrhaphis petersi (alcohol). ANSP 163026 (231 mm SL). Venezuela, Amazonas, Río<br />
Sipapo (Orinoco drainage), backwater channel behind sandbar 6-7 km above Pendare,<br />
04u519N, 67u439W, 12 Nov 1985, B. Chern<strong>of</strong>f et al.<br />
F. Pseudotylosurus microps (alcohol). ANSP 179633 (178 mm SL). Guyana, Rupununi River<br />
(Essequibo drainage) at Kwatamang, 4 km SE <strong>of</strong> Annai, 03u559030N, 059u069010W, 25 Oct<br />
2002, M. Sabaj, J. Armbruster, M. Thomas et al.
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90 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Cichlidae<br />
Plate 15<br />
A. Crenicichla multispinosa (live). ANSP 187101 (223 mm SL). Suriname, Sipaliwini, Lawa<br />
River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
B. Geophagus harreri (live). ANSP 187136 (170 mm SL). Suriname, Sipalawini, Litanie River<br />
at mouth and confluence with Marowini River, just upstream from settlement <strong>of</strong> Konya<br />
Kondre, 03u179240N, 054u049380W, 21 Apr 2007, J. Lundberg, M. Sabaj, P. Willink, J.<br />
Mol et al.<br />
C. Geophagus sp. (live). AUM 38940 (103 mm SL). Guyana, Kuyuwini River, main channel<br />
and backwater 19.5 km W <strong>of</strong> confluence with Essequibo River (Atlantic drainage).<br />
02u149280N, 058u309030W, 11 Nov 2003, M. Sabaj, J. Armbruster, M. Hardman et al.<br />
D. Hoplarchus psittacus (live). AUM 41425 (147 mm SL). Venezuela, Río Ventuari (Orinoco<br />
drainage), bedrock outcrops 83 km E-NE <strong>of</strong> Macuruco, 153 km E-NE <strong>of</strong> San Fernando<br />
de Atabapo, 04u159120N, 066u209410W, 6 Apr 2004, M. Sabaj, N. Lujan, D. Werneke<br />
et al.<br />
E. Mesonauta insignis (live). AUM 40590 (82 mm SL). Venezuela, Río Orinoco near mouth<br />
<strong>of</strong> Río Ventuari, Macuruco Landing, 75 km E <strong>of</strong> San Fernando de Atabapo, 03u579290N,<br />
067u019560W, 4 Apr 2004, M. Sabaj, N. Lujan, D. Werneke et al.<br />
F. Cichla intermedia (live). ANSP 189269 . Venezuela, Amazonas, Río Orinoco, bedrock<br />
outcrop, 52.9 km SE <strong>of</strong> San Antonio, 102 km W <strong>of</strong> La Esmerelda, 03u069010N,<br />
066u279460W, 4 Mar 2005, M. Sabaj, N. Lujan, D. Werneke et al.<br />
G. Pterophyllum altum (live). AUM 40584 (59 mm SL). Same locality as E.
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92 BULLETIN OF THE BIOLOGICAL SOCIETY OF WASHINGTON<br />
Synbranchidae<br />
Plate 16<br />
A. Synbranchus marmoratus (live). ANSP 187334 (492 mm TL). Suriname, Sipaliwini, Lawa<br />
River (Marowini drainage), ca. 8 km S-SW <strong>of</strong> Anapaike/Kawemhakan (airstrip),<br />
03u199310N, 054u039480W, 18–22 Apr 2007, J. Lundberg, J. Mol, M. Sabaj, P. Willink, &<br />
K. Wan.<br />
B. Synbranchus marmoratus (live, ventral view). Same data as A.<br />
Sciaenidae<br />
C. Pachyurus schomburgkii (alcohol). ANSP 162800 (195 mm SL). Venezuela, Amazonas,<br />
Río Iguapo approximately 1 hour above its confluence with Río Orinoco, 03u099N,<br />
065u289W, 13 Mar 1987, H. Lopez et al.<br />
D. Plagioscion squamosissimus (alcohol). ANSP 177421 (257 mm SL). Guyana, Siparuni<br />
River (Essequibo drainage), Black Water camp, 04u449N, 058u599W, 6 Dec 1997, G.<br />
Watkins et al.<br />
Eleotridae<br />
E. Microphilypnus sp. (live). ANSP 180643. Venezuela, Amazonas, Río Negro (Amazonas<br />
drainage), left bank sandy beach and small adjacent backwater 7.2 km NW <strong>of</strong> San Carlos<br />
de Rio Negro, 01u589110N, 067u069100W, 19 Mar 2005, M. Sabaj, D. Werneke et al.<br />
Achiridae<br />
F. Hypoclinemus mentalis (alcohol). ANSP 163857 (80.5 mm SL). Venezuela, Amazonas,<br />
caño <strong>of</strong> Río Orinoco separating island and beach just downstream <strong>of</strong> Quiratare, 02u599N,<br />
066u049W, 11 Mar 1987, B. Chern<strong>of</strong>f et al.<br />
G. Soleonasus finis (alcohol). ANSP 179510 (75.5 mm TL). Guyana, Essequibo River<br />
(Atlantic drainage), E bank at Kurukupari, 04u399410N, 058u409310W, 24 Oct 2002, M.H.<br />
Sabaj et al.<br />
Polycentridae<br />
H. Polycentrus schomburgkii (live). FMNH ex ANSP 189014 (22.3 mm SL). Suriname, Para,<br />
Coropinae Creek (Suriname drainage), vicinity <strong>of</strong> Republiek, 05u299570N, 055u129520W,<br />
28 Apr 2007, M. Sabaj & P. Willink.<br />
Tetraodontidae<br />
I. Colomesus asellus (live). AUM 37945 (35 mm SL). Guyana, Essequibo River (E bank) at<br />
Kurukupari, 04u399410N, 058u409310W, 17 Nov 2003, M. Sabaj, M. Hardman, N. Lujan<br />
et al.
NUMBER 17 93
INDEX TO ORDERS, FAMILIES, AND SUBFAMILIES<br />
Acestrorhynchidae<br />
Achiridae<br />
Agoniatinae<br />
Anguilliformes<br />
Anostomidae<br />
Aphyocharacinae<br />
Apteronotidae<br />
Arapaimidae<br />
Aspredinidae<br />
Auchenipteridae<br />
Belonidae<br />
Beloniformes<br />
Bryconinae<br />
Callichthyidae<br />
Cetopsidae<br />
Characidae<br />
Characiformes<br />
Characinae<br />
Cheirodontinae<br />
Chilodontidae<br />
Cichlidae<br />
Clupeidae<br />
Clupeiformes<br />
Crenuchidae<br />
Ctenoluciidae<br />
Curimatidae<br />
Cynodontidae<br />
Cyprinodontiformes<br />
Doradidae<br />
Engraulidae<br />
Erythrinidae<br />
Gasteropelecidae<br />
Genera Incerta Sedis, family<br />
Characidae<br />
Glandulocaudinae<br />
Gobiidae<br />
Gymnotidae<br />
Gymnotiformes<br />
Hemiodontidae<br />
Hemiramphidae<br />
Heptapteridae<br />
Hypopomidae<br />
Hypoptopomatinae<br />
Hypostominae<br />
Iguanodectinae<br />
Lebiasinidae<br />
Lepidosirenidae<br />
Lepidosireniformes<br />
Lithogeninae<br />
Loricariidae<br />
Loricariinae<br />
Myliobatiformes<br />
Ophichthidae<br />
Osteoglossidae<br />
Osteoglossiformes<br />
Parodontidae<br />
Perciformes<br />
Pimelodidae<br />
Pleuronectiformes<br />
Poeciliidae<br />
Polycentridae<br />
Potamotrygonidae<br />
Pristidae<br />
Pristiformes<br />
Pristigasteridae<br />
Prochilodontidae<br />
Pseudopimelodidae<br />
Rhamphichthyidae<br />
Rivulidae<br />
Sciaenidae<br />
Serrasalminae<br />
Siluriformes<br />
Sternopygidae<br />
Stethaprioninae<br />
Synbranchidae<br />
Synbranchiformes<br />
Tetragonopterinae<br />
Tetraodontidae<br />
Tetraodontiformes<br />
Trichomycteridae
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