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P Cyrino Digital Instant Download
Author(s): J E P Cyrino
ISBN(s): 9781578083756, 1578083753
Edition: 1
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Year: 2008
Language: english
Feeding and Digestive
Functions of Fishes
Feeding and Digestive
Functions of Fishes
Editors
J.E.P. Cyrino
USP-ESALQ
Department of Animal Science (Zootecnia)
University of Sao Paulo
Piracicaba, SP
Brazil
D.P. Bureau
Fish Nutrition Research Laboratory
Department of Animal and Poultry Science
University of Guelph
Guelph, Ontario
Canada
B.G. Kapoor
Formerly Professor of Zoology
The University of Jodhpur
Jodhpur, India
Science Publishers
Enfield (NH) Jersey Plymouth
CRC Press
Taylor & Francis Group
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No claim to original U.S. Government works
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Version Date: 20131029
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Preface
Understanding the biology of the innumerable number of aquatic species

on our planet is the focus of sustained research efforts. Environmental
degradation, management or rehabilitation of wild stocks, and the
forecasted climatic changes are fueling interest in the study of the ecology,
feeding behavior, and nutrition of aquatic animals in their natural habitat.
In parallel, the rapid expansion of aquaculture in many parts of the world
has supported vigorous research programs on nutrition and feeding of
cultivated aquatic organisms. More than 250 aquatic species are
cultivated on a commercial scale using a wide variety of production
systems. This great diversity results in great opportunities and tremendous
challenges. The number of scientific publications on nutrition and feeding
of aquatic animals is ever increasing and our knowledge base is expanding
rapidly.
The diversity of species and issues studied and the large number of
papers published each year makes it very difficult to keep up with progress.
It is highly desirable to occasionally review the state-of-the-art and
progresses, identify gaps in our understanding, and pinpoint emerging
issues. There are numerous books providing a good overview of state-of-
the-art in fish nutrition. Numerous authors have reviewed in details the
nutritional requirements of fish, notably from an aquaculture perspective.
However, few books have specifically focused on feeding and digestion in
fish in captivity or in their natural habitat. This book, therefore, attempts
to fill what we think was a significant gap in the reference literature.
Scientists actively involved in research on feeding and digestion in fish
were invited by the editorial team on the basis of their expertise and track
record. These investigators were given "carte blanche" as to the content
and style of their contribution. The twelve chapters contributed by these
vi Preface
investigators cover topics as diverse as feeding ecology of fish in their

natural habitat, feeding behavior, digestive anatomy, biochemistry and
physiology of different fish species at different life stages, the impact of diet
on gastro-intestinal development and health, nutrition and disease
resistance, and modeling conversion of food or feed inputs into biomass.
We believe this book contains a wealth of information and will be
valuable to a wide variety of readers. However, it does not pretend to be
exhaustive or perfect. It is mainly an attempt to allow sharing of
knowledge, experience, and ideas. In the various chapters, a number of
interesting and rather unique scientific analyses are presented and these
may capture the interest of scientists and graduate students. Aquaculture
specialists, fisheries biologists, students in natural and applied science
programs, and fish feed manufacturers will find in this book relevant
information suited to their diverse needs. We invite readers to approach
the material in this book with an inquisitive mind. This book should
initiate learning and rather complete it.
J.E.P. Cyrino, D.P. Bureau and B.G. Kapoor

Contents
Preface v
List of Contributors ix
1. Feeding Ecology of Fishes in the Brazilian Amazon - 1

A Naturalistic Approach
Jansen Zuanon and Efrem Ferreira
2. Environmental Changes, Habitat Modifications and 35
Feeding Ecology of Freshwater Fish
Norma Segatti Hahn and Rosemara Fug
3. Feeding Activity in Teleost Fish: Influence of Biotic and 67
Abiotic Factors
Ann L. Gannam
4. Classical and Modern Concepts in Fish Digestion 85

Victoria V. Kuz'mina
5. Temperature Adaptation of Digestive Enzymes in Fish 155
Alexander Gelman, Victoria Kuz'mina, Vladimir Drabkin
and Larisa Glatman
6. Diets, Physiology, Biochemistry and Digestive Tract 227
Development of Freshwater Fish Larvae
Maria Cé lia Portella and Konrad Dabrowski
7. Ontogeny and Physiology of the Digestive System of 281

Marine Fish Larvae
José Luiz Zambonino Infante, Enric Gisbert, Carmen Sarasquete,
Isabel Navarro, Joachim Gutiérrez and Chantal L. Cahu
viii Contents
8. Tracer Studies in Fish Larvae 349

Luís E.C. Conceiçã o, Sofia Morais, Maria Teresa Dinis
and Ivar Rø nnestad
9. An Overview of the Feeding Ecology and Physiology of 393
Elasmobranch Fishes
Enric Corté s, Yannis P. Papastamatiou, John K. Carlson,
Lara Ferry-Graham and Bradley M. Wetherbee
10. Alternative Protein Sources and Digestive Function 445
Alterations in Teleost Fishes
Anne Marie Bakke-McKellep and Stå le Refstie
11. Nutrition and Disease Resistance in Fish 479
Chhorn Lim, Mediha Yildirim-Aksoy and Phillip H. Klesius
12. Efficiency of Conversion of Feed Inputs into Animal 547
Biomass: The Usefulness of Bioenergetics Models and
Need for a Transition to Nutrient-flow Models
Dominique P Bureau, Katheline Hua and Paula A. Azevedo
Index 569
Color Plate Section 577
List of Contributors
Azevedo Paula A.
Fish Nutrition Research Laboratory, Dept. of Animal and Poultry
Science, University of Guelph, Guelph, Ontario N1G 2W1 Canada.
E-mail: [email protected]
Bakke-McKellep Anne Marie
Aquaculture Protein Centre, Department of Basic Sciences
and Aquatic Medicine, Norwegian School of Veterinary Science,
P.O. Box 8146, N-0033 Oslo Norway.
Bureau Dominique P.
Science, University of Guelph, Guelph, Ontario N1G 2W1 Canada.
Cahu Chantal L.
IFREMER Centre de Brest, UMR 1067-Nutrition Aquaculture
Génomique des Poissons, BP 70, 29280 Plouzané France.
Carlson John K.
NOAA Fisheries, Panama City Laboratory, 3500 Delwood Beach
Road, Panama City, FL 32408 USA.
x List of Contributors
Conceiçã o Luís E.C.

Centro de Ciências do Mar - CCMAR, Universidade do Algarve,
Campus de Gambelas, 8005-139 Faro Portugal.
Cortés Enric
NOAA Fisheries, Panama City Laboratory, 3500 Delwood Beach Rd,
Panama City, FL 32408 USA.
Dabrowski Konrad
School of Natural Resources, College of Food, Agricultural and
Environmental Sciences, Ohio State University, 473D Kottman Hall,
2021 Coffey Rd., Columbus, OH 43210 USA.
Dinis Maria Teresa

Centro de Ciéncias do Mar-CCMAR, Universidade do Algarve,
Drabkin Vladimir
Fishery Products Laboratory, Kimron Veterinary Institute, P.O. Box 12,
Bet Dagan 50250 Israel.
Ferreira Efrem
Coordenaçã o de Pesquisas em Biologia Aquática, Instituto Nacional
de Pesquisas da Amazônia-INPA, Av. André Araújo, 2936 - Aleixo,
CP 478, 69083-970 Manaus, AM Brazil.
Ferry-Graham Lara A.
California State University, Moss Landing Marine Laboratories, 8272
Moss Landing Rd, Moss Landing, CA 95039 USA.
Fugi Rosemara
Universidade Estadual de Maringá, Núcleo de Pesquisas em
Limnologia, Ictiologia e Aquicultura - Nupélia, Av. Colombo, 5790,
Campus Universitário, 87020-900 Maringá PR Brazil.
List of Contributors xi
Gannam Ann L.
Abernathy Fish Technology Center, 1440 Abernathy Creek Road,
Longview, WA 98632 USA.
Gelman Alexander
. Box
Fishery Products Laboratory, Kimron Veterinary Institute, PO.
12, Bet Dagan 50250 Israel.
Gisbert Enric
Centre d'Aqüicultura, Institut de Recerca i Tecnologia
Agroalimentaries (IRTA), Aptat. Correus 200, 43540 Sant Carles de
la Ràpita Tarragona, España.
Glatman Larisa
Fishery Products Laboratory, Kimron Veterinary Institute, P.O. Box
12, Bet Dagan 50250 Israel.
Gutierrez Joachim
Department of Physiology, Faculty of Biology, University of Barcelona,
Ave. Diagonal 645, 08028, Barcelona, Spain.
Hahn Norma Segatti

Universidade Estadual de Maringá, Núcleo de Pesquisas em
Limnologia, Ictiologia e Aquicultura - Nupélia, Av. Colombo, 5790,
Campus Universitário, 87020-900 Maringá, PR Brazil.
Hua Katheline
Science, University of Guelph, Guelph, Ontario, N1G 2W1 Canada.
Klesius Phillip H.
Aquatic Animal Health Research Unit, USDA-ARS, P.O. Box 952,
Auburn, AL 36831 USA.
xii List of Contributors
Kuz'mina Victoria V.
Laboratory of Fish Ecology, I.D. Papanian Institute for Biology of
Inland Waters, Russian Academy of Sciences, 152742 Borok Nekouz
Yaroslavl Russia.
Lim Chhorn
Aquatic Animal Health Research Unit, USDA-ARS, P.O. Box 952,
Auburn, AL 36831 USA.
Morais Sofia
Centro de Ciências do Mar - CCMAR, Universidade do Algarve,
Navarro Isabel
Department of Physiology, Faculty of Biology, University of Barcelona,
Ave. Diagonal 645, 08028, Barcelona, Spain.
Papastamatiou Yannis P.
Department of Zoology, Hawaii Institute of Marine Biology, University
of Hawaii at Manoa, Honolulu, HI 98822, USA.
Portella Maria Célia

Universidade Estadual Paulista, Centro de Aqüicultura da UNESP,
Via de Acesso Prof. Paulo Donato Castellane s/n, 14884-900
Jaboticabal, SP Brasil.
Refstie Ståle
Aquaculture Protein Centre, Nofima Akvaforsk-Fiskeriforskning,
N-6600 Sunndals0ra, Norway.
R0nnestad Ivar
Department of Biology, University of Bergen, PB 7800, 5020 Bergen,
Norway.
List of Contributors xiii
Sarasquete Carmen
Instituto de Ciencias Marinas de Andalucía - CSIC, Polígono Rio San
Pedro s/n, Apdo. Oficial 11510, Puerto Real Cádiz, España.
Wetherbee Bradley M.
Department of Biological Sciences, University of Rhode Island, 100
Flagg Road, Kingston, RI 02881 USA.
Yildirim-Aksoy Mediha
Aquatic Animal Health Research Unit, USDA-ARS, 990 Wire Road,
P. O. Box 952, Auburn, AL 36831 USA.
Zambonino-Infante José Luis

IFREMER Centre de Brest, UMR 1067-Nutrition Aquaculture
Génomique des Poissons, BP 70, 29280 Plouzané, France.
Zuanon Jansen
Coordenaçã o de Pesquisas em Biologia Aquática, Instituto Nacional
de Pesquisas da Amazônia-INPA, Av. André Araújo, 2936 - Aleixo,
CP 478, 69083-970 Manaus, AM Brazil.
CHAPTER
1
Feeding Ecology of Fishes in

the Brazilian Amazon -
A Naturalistic Approach
Jansen Zuanon* and Efrem Ferreira #
INTRODUCTION
The Amazon system is composed by a multitude of rivers with varying

water types and strikingly different productivities. This environmental
diversity is accompanied by the presence of diverse fish species
assemblages and ecological characteristics. Although a considerable
amount of information about the trophic relations of the Amazonian
fishes and their natural habitats have been generated in the previous few
decades, most of the published literature deals only with descriptions of
the main food items of the commercially most important species, and is
based mainly on stomach contents analyses. We present here a quick look
into the trophic interactions of the fishes and their natural environments,
based on the diversity of the Amazonian fish fauna and its aquatic
Authors' address: Coordenaçã o de Pesquisas em Biologia Aquática, Instituto Nacional de
Pesquisas da Amazônia-INPA, Av. André Araújo, 2936 - Aleixo, CP 478, 69083-970
Manaus, AM Brazil.
Corresponding authors: E-mail: *[email protected]; #[email protected]
2 Feeding and Digestive Functions of Fishes
habitats. This chapter focuses on the main aquatic environments found in

the Brazilian Amazon: the huge river-floodplain systems and their
different water types; the multitude of small clear and black water streams
that drains the "terra firme" (upland, non-floodable) forests, including
their marginal temporary ponds; and the riffles and rapids zones of the
rivers that drain the Brazilian and Guyana Shields and harbor a very rich
and trophically specialized fish fauna.
White water river-floodplain systems sustain high primary
productivity that is responsible for the major portion of the biomass
produced in the aquatic environments in the Amazon. The flood pulse, a
monomodal seasonal rising of the river level, is the most important factor
influencing the life of fishes in the large rivers of the Amazon. The cyclic
rising of the water level inundates a large area of riparian forest, and in the
process makes available new habitats and an enormous variety of food
items to the fish fauna. It is during this time of the year that the fishes can
enter the flooded forests and feed on fruits and invertebrates that fall from
the trees. The large black water rivers present an extremely low aquatic
primary productivity but contain a surprisingly rich and diverse
ichthyofauna. A low abundance of most species and a strong dependence
on allochthonous food sources are the remarkable characteristics of the
fish fauna of Amazonian black waters.
The Amazon forests are also drained by an endless number of small
streams of clear or black waters which are characteristically oligotrophic
and apparently sustain a low biomass of fishes. The nutrient-limited
environment of the first-order streams is occupied almost exclusively by
fishes that feed on allochthonous sources of food, such as terrestrial
insects and other small invertebrates that fall from the trees. These fragile,
low-productivity aquatic environments are being submitted to strong
environmental disturbance, mainly derived from the deforestation of large
areas for agriculture, pasture, and urban growth.
The clear water rivers that drain the Brazilian and Guiana Shields
show highly variable chemical characteristics and primary productivity,
and usually have a series of rapids and waterfalls along their courses. These
turbulent zones are the habitat of many endemic fish species that rely on
the local periphytic production as their main food source, which is
exploited in a variety of specialized ways. The high periphytic productivity
coupled with variations in mouth and teeth morphology and different
feeding strategies allow for the presence of highly diverse fish fauna. The
Jansen Zuanon and Efrem Ferreira 3
presence of many endemic species in the rapids also points out to the need
for special conservation strategies since those areas are highly threatened
by the construction of hydropower plants in the Brazilian Amazon.
The relationships between fishes and flooded forests probably
represent the most remarkable biological interaction in the aquatic
environments of the Amazon, and represent an important energy pathway
between the terrestrial and aquatic systems. The knowledge of these
intricate ecological interactions and the measurement of their impact on
the regional economy may represent an effective way to preserve the
magnificent flooded forests and the amazing diversity of aquatic
environments in the Amazon.
The most remarkable ecological characteristic of the Amazonian
ichthyofauna probably is its close relationship with the forest. The seminal
work of Goulding (1980) in the Madeira River system depicted in detail
the intimate relations of the fishes and the flooded forest, where fish acted
as fruit and seed predators and simultaneously, as seed dispersers that help
to maintain the high plant diversity of the riparian forests. Nevertheless,
the Amazon system is composed of a multitude of rivers with varying types
of water bodies and with strikingly different productivities. White water
river-floodplain systems sustain high primary productivity, both by
phytoplankton production in the floodplain lakes and the rapidly growing
aquatic macrophytes that abound in the shallow marginal areas. These
floodplains are responsible for the major portion of the biomass produced
in the aquatic environments in the Amazon and also sustain intense
commercial fisheries. The black waters of the Rio Negro and other
Amazon tributaries represent the opposite situation, with an extremely
low aquatic primary productivity but holding a surprisingly rich and
diverse ichthyofauna (Goulding et al., 1988). The clear water rivers that
drain the Brazilian and Guiana Shield lie between these extremes,
showing highly variable chemical characteristics and primary productivity
(Sioli, 1984). Furthermore, these rivers present a series of rapids and
waterfalls along the transition region between the Guianas and Central
Brazilian plateaus and the main Amazon sedimentary basin. These riffle
zones are the habitat of many endemic fish species that rely on the local
periphytic production as their main food source, which is exploited in a
variety of specialized ways.
This environmental diversity is accompanied by the presence of
different fish species assemblages and ecological characteristics. Although
the ecological interactions in the aquatic environments of the Amazon are
still far from well known, a considerable amount of information about the
trophic relations of the fishes and their natural habitats have been
generated in the last few decades (e.g. Marlier, 1967, 1968; Knöppel,
1970; Soares, 1979; Goulding, 1980; Ferreira, 1981, 1993; Goulding and
Carvalho, 1982, 1984; Santos, 1982; Carvalho and Goulding, 1985; Silva,
1993; Santos et al., 1997; Sabino and Zuanon, 1998; Santos and Rosa,
1998; Zuanon, 1999; Walker, 2004). Most of the published data
concerning the diet of Amazonian fishes deals only with descriptions of
the primary food items of the commercially most important species, and
were based mainly on stomach contents analyses. Furthermore, very few
studies present information on the nutritional contribution of the various
kinds of foods consumed by the fishes, which do not allow for a precise
analysis of their relative importance on the fish diets. Although important,
these approaches to the study of the feeding ecology of fishes usually do
not allow for a comprehensive view of the diverse and complex
interactions of the fish fauna in different biotopes. The available
information in the literature usually presents generalized views of the
trophic characteristics of the regional fish fauna, or comprises somewhat
loose data regarding the diet of single species. In other instances, fish
community studies end up with a multitude of data about a large number
of species, so deeply transformed by multivariate statistical techniques
that the beautiful trophic interactions that occur in the natural
surroundings are not readily evident.
In view of these limitations, we aim to present a different look at the
trophic interactions of the fishes and their natural environments, based on
the diversity of the Amazonian fish fauna and its aquatic habitats. This
chapter focuses on the main aquatic environments found in the Brazilian
Amazon: the huge river-floodplain systems and their different water types;
the multitude of small clear- and black-water streams that drain the 'terra
firme' (upland, non-floodable) forests, including their marginal temporary
ponds, and the riffles and rapids zones of the rivers that drain the Brazilian
and Guyana Shields and harbor a very rich and trophically specialized fish
fauna.
Most information presented herein is derived from the authors'
personal experiences gathered during field activities and along several
research projects developed in the Brazilian Amazon. We hope that this
chapter will provide some new perspectives on the feeding ecology of
freshwater tropical fishes, and stimulate a much-needed revival of the
naturalistic approach to the study of the Amazon and its fishes.
RIVER-FLOODPLAIN SYSTEMS
The flood pulse, a monomodal seasonal rising of the river level, is the most
important factor influencing the life of fishes in the large rivers of the
Amazon (Junk et al., 1989). The cyclic rising of the water level inundates
a large area of the riparian forest, making available new habitats and an
enormous variety of food items to the fish fauna. It is during this time of
the year that fishes can enter the flooded forests and feed on fruits and
invertebrates that fall from the trees. The extravasation of the water over
the banks of the rivers also floods lateral depressions along the river
channel, thus constituting more lentic habitats that function temporarily
as true lakes, especially when the water recedes and these water bodies
become isolated from the river channel (Plate 1.1).
In white water river basins, these lakes constitute the most productive
natural aquatic environment of the Amazon (Junk, 1970, 1973; Sioli,
1984; Junk and Piedade, 1993), but the situation is markedly different in
black water rivers. The low autochthonous productivity of black waters
does not sustain a large biomass of fishes. A low abundance of most species
and a strong dependence on allochthonous food sources are remarkable
characteristics of the fish fauna of black waters such as the Negro River
in the Brazilian Amazon (Goulding et al., 1988). Even the reduction of
adult body size ( = miniaturization) has been hypothesized to represent an
evolutionary strategy to minimize the energetic demand among
Amazonian fishes (Weitzman and Vari, 1988). Clear water rivers are
somewhat more productive and apparently sustain a larger biomass of
fishes, mainly dependent of the periphyton production (e.g. Ferreira,
1993; Zuanon, 1999).
Floodplains have long been recognized as critically important habitats
for the ichthyofauna, whereas the river channel itself has been supposed
to serve mainly as migration routes for most Amazonian fishes (Junk et al.,
1989). Recent studies have revealed that the river channels harbor a rich
fish fauna dominated by species of catfishes and electric knife fishes, but
still poorly known with respect to trophic characteristics (e.g. Lundberg
et al., 1987; Cox-Fernandes et al., 2004). In order to allow for a better
understanding of such diverse and complex trophic characteristics and
interrelationships, we choose to present information for each of the main
habitat types recognized in these river-floodplain systems: the river
channels, including the steep banks, beaches, and the channel itself (open
water and bottom habitats); and the floodplain lakes, including the open
Plate 1.1
(Upper left) General view of a floodplain lake along the Amazon river margin. Note the
floating meadows bordering the lake margins. Photo: E. Ferreira.
Plate 1.1 Contd.

water habitats, floating meadows and the flooded forests. Although

present both at river and lake margins, the flooded forest is discussed
among the floodplain habitats in function of its predominantly lentic
characteristics.
River Channels
Probably one of the most obvious habitats along a river is its banks. The
steep river margin usually constitutes a highly dynamic habitat that is
composed of many different features, such as backwaters, eddies, small
bays, and submerged woods. The calmer zones are commonly occupied by
shoals of the pike characin Boulengerella spp. (Plate 1.1) and other
opportunistic piscivorous species that ambush and pursue small characins
moving along the river margins. The woody shores are the habitat of some
large armored, sucker mouth catfishes like Panaque spp. and Acanthicus
hystrix (Loricariidae). Panaque catfishes are known to feed on wood
fragments scraped of submerged tree trunks and branches, although how
the fish digests the cellulose is not well known (Nelson et al., 1999).
Acanthicus hystrix, the largest loricariid catfish (up to one meter long), is
also frequently found foraging on the periphyton that grows in the
submerged trunks. Large doradid catfishes like adult Pterodoras granulosus
and Megalodoras uranoscopus congregate in the deep and calm bays to feed
on insects, mollusks, small fishes and almost any edible organic particles.
Shoals of the pimelodid catfish Sorubim lima are usually found in these
habitats searching for small fishes, shrimps and worms close to the
Plate 1.1 Contd.

(Upper right) A sandy beach in the Amazon River near Santarém, Pará State, Brazil.
Photo: E. Ferreira.
(Second row, left) A small school of the pike characin Boulengerella maculata ambushing
small prey fishes close to the banks of Araguaia River, Tocantins State, Brazil. Photo:
J. Zuanon.
(Second row, right) The boldly patterned scale-eating characin Exodon paradoxus
(Characidae) in field aquarium. Photo: E. Ferreira.
(Third row, left) A hematophagous candiru (Vandellia cirrhosa, Trichomycteridae)
attacking a young tambaqui (Colossoma macropomum, Characidae) in aquarium. Photo:
I. Sazima.
(Third row, right) A large Tambaqui (Colossoma macropomum, Characidae) taking a
rubber tree fruit (Hevea spruceana) at the water surface. Photo: M. Goulding.
(Bottom left) A stingray (Potamotrygon leopoldi, Potamotrygonidae) partially buried in the
sand in a beach of Xingu River, Brazil. Photo: J. Zuanon.
(Bottom right) The needlefish Potamorrhaphis guianensis staying motionless near the
marginal vegetation in Araguaia River, Brazil. Needlefishes usually ambush small fishes
and catch drifting invertebrates along river and lake banks. Photo: E. Ferreira.
substrate (Goulding and Ferreira, 1984), whereas groups of the widespread

catfish Pimelodus blochii forage on a variety of food items that include
fruits, invertebrates, fishes and organic debris. Zungaro zungaro, one of the
giant pimelodid catfishes stays in the deeper portions close to the banks
where it preys on medium-sized fishes, such as the migratory
prochilodontids Prochilodus and Semaprochilodus (Goulding, 1979, 1980;
Barthem and Goulding, 1997; authors' pers. obs.).
During the dry season, a large portion of the river margins appears in
the form of sandy or muddy beaches (Plate 1.1). These gently sloping
margins may form long extensions of shallow waters and represent rather
simple habitats, with plenty of light scattered by the sand and clay particles
suspended in the water. The highly seasonal nature of the beaches and its
low structural complexity are supposed to result in rather unpredictable
species assemblages (e.g. Goulding et al., 1988; Ibarra and Stewart, 1989;
Jepsen, 1997). Nevertheless, some field evidences indicate that the
presence of a few "core species" that are rather frequent and numerically
dominant in the fish fauna of the beaches. These species can be
characterized by their small to medium size, drab and light color, and
generalized feeding habits. Up to one hundred species of small characins,
hemiodontids, loricariinae catfishes, shoaling engraulidids and cichlids
(mostly geophagine "earth-eaters") comprise most of the fish fauna that
congregate in the shallow beaches in the Amazon. Streamlined
piscivorous characins such as species of Acestrorhynchus, Boulengerella,
Rhaphiodon vulpinus and Cynodon patrol the open waters pursuing small
fishes. Needlefishes (mainly Pseudotylosurus sp.) also hunt for small fishes
close to the surface, biting off the head of their prey in the process
(Goulding and Carvalho, 1984). The plain and well-lit sandy beaches are
home to several hemiodontid fishes, such as Argonectes longiceps,
A. robertsi, Hemiodus immaculatus, and Bivibranchia spp., that take
mouthfuls of substrate and sift the insect larvae, microcrustaceans, and
any small edible particles found among the sand. The odd-looking
freshwater flatfishes Hypoclinemus mentalis and Apionichthys spp.
(Achiridae) occupy the same habitat, burying themselves in the substrate
in order to ambush invertebrates and small fishes.
Another group of fishes commonly found in the beaches are the scale-
eaters. Several species of the characid genera Roeboides, Roeboexodon,
Bryconexodon, Exodon and the wimple piranha Catoprion mento ambush
their prey close to the shore, hidden from their prospective prey among
submerged tree branches or grass leaves. Some scale eaters of the genus
Roeboides are supposed to rely on their body translucency and scattered

light in the beaches to avoid being detected by their prey (Sazima and
Machado, 1982), while others like Exodon paradoxus (Plate 1.1) are boldly
marked and forage unobtrusively in large schools (Sazima, 1983).
At night-time a different set of fishes emerge from deeper river
portions to forage in the shallow beaches, including predators that
approach the shore to hunt for fish and other preys. The large and flat-
headed pimelodid catfishes Sorubimichthys planiceps, Pseudoplatystoma
fasciatum and P. tigrinum forage solitarily in the shallows, engulfing their
prey (mainly resting diurnal fishes) with a loud, shot-like noise. Species of
Potamotrygon stingrays (Plate 1.1) dig the soft substrate with their
undulating pectoral fins in search for buried worms and insects, while large
Paratrygon aiereba stingrays (up to 1.1 m disk diameter and over 70 kg)
chases the resting diurnal fishes in the shallows. Long, flat-bodied and
cryptic-colored loricariinae catfishes (e.g. Loricariichthys spp., Loricaria
spp., Pseudoloricaria laeviuscula) that stayed partially buried in the
substrate during the day emerge in the dark to feed on organic detritus and
small insect larvae.
Although small fishes constitute one of the main prey items at night
in the beaches, medium to large-sized fishes resting in the shallows are not
safe either. Some highly specialized predators, the strictly hematophagous
catfishes of the genus Vandellia (Trichomycteridae) (Plate 1.1) abound in
some places, and feed on the blood of their host by piercing its arteries at
the corner of branchial arches (Zuanon and Sazima, 2004a). The beach-
dwelling Stegophilinae trichomycterids Pseudostegophilus nemurus,
Henonemus punctatus and Stegophilus spp. are also highly specialized
feeders. They apparently subsist exclusively on the mucus and scales of
other fishes (Winemiller, 1989) that are rasped off by attaching the mouth
to the host's body and widely swinging its body like a pendulum.
The river channels are supposed to serve mainly as migration routes
for several groups of fishes, notably the long-ranging pimelodid goliath
catfishes of the genera Brachyplatystoma (Junk et al., 1989; Lundberg and
Akama, 2005), as well as passive dispersion pathways to larval stages of
many fish species. In fact, few fish species permanently inhabit the water
column of the main river channel. Large individuals of the pristigasterid
Pellona castelnaeana represents one of the few fishes that live in the open
waters of white water river channels, pursuing smaller prey fishes (Le
Guennec and Loubens, 2004). Some species of wide-mouthed catfishes of
the genus Ageneiosus also live in the river channels and hunt at night for
fishes and shrimps that are gulped down whole.
The deeper portions of the river channels present a very distinct
situation, with a high diversity of fishes living permanently in the dark
river bottom. This fish fauna is composed mainly by catfishes (especially
pimelodids, doradids, and loricariids) and by gymnotiform electric knife
fishes (Lundberg et al., 1987). The deep river channels do not sustain an
important autochthonous primary productivity since light penetration is
scarce and never reaches the bottom. The food utilized by the fishes is
carried by the current from upstream locations after being produced in the
lakes or in the flooded forests. Most of the fish fauna is composed by
opportunistic foragers that depend on small animals, plant debris, organic
detritus and even carcasses brought by the current to feed. The whale
catfishes Cetopsis coecutiens and C. candiru (Cetopsidae), and Pareiodon
microps (Trichomycteridae) are examples of voracious carrion-feeders that
forage on dead (or dying) animals in very large groups (just like the well-
known piranhas in lakes). A high diversity of pimelodid catfishes (e.g.
Platystomatichthys sturio, Exallodontus aguanai, Aguarunichthys inpai,
Propimelodus eigenmanni) live permanently in the river channels and feed
on animal matter (invertebrates, small fishes, and carrion) but their
trophic interrelationships are largely unknown.
The diets and trophic relations of the gymnotiform knife fishes, which
show an impressive variety of snout sizes, mouth shapes and teeth
arrangements, remains scarcely studied. Some long-snouted species (e.g.
Sternarchorhynchus spp. and Sternarchorhamphus muelleri, Apteronotidae)
feed on insect aquatic larvae taken from small holes and channels in the
mud bottom, in sponge beds or in conglomerates of pebbles or laterite
(Marrero and Winemiller, 1993). One of the most bizarre feeding habits
among knife fishes is shown by Magosternarchus duccis and M. raptor that
seem to include the tail of other gymnotiform fishes as a staple item on
their diets (Lundberg et al., 1996). Despite such pinpointed examples,
much remains to be studied about the trophic relationships of the deep
channel fish assemblages of Amazonian rivers.
Floodplain Lakes
Among the different habitats recognized in the floodplain lakes, some are
available throughout the year such as the open waters and the lake
bottoms, whereas others are highly seasonal, like the floating meadows
and flooded forests. The open waters are dominated by wandering
predators and plankton-eating schooling fishes. Among the commonest

predators are the clupeiform Pellona flavipinnis and the characiforms
Acestrorhynchus falcirostris and A. microlepis, which pursue small fishes
(mainly young-of-the-year of medium to large-sized species) in the
water column. The fish- and invertebrate-eating Ilisha amazonica
(Pristigasteridae) also forages in open water, where it feeds on small
schooling engraulidid fishes (e.g. Anchoviella spp.), pelagic shrimps (Acetes
paraguayensis, Sergestidae), and swimming insects like notonectid bugs
(Heteroptera) and mayfly larvae (Ephemeroptera). Its large and dorsally
positioned eyes possibly allow the capture of fishes and invertebrates that
swim close to the surface, detected from bellow and evidenced against the
sky light (JZ, pers. obs.).
Planktivorous fishes compose another feeding guild that forage in the
open waters of lakes. Large schools of the pelagic catfish Hypophthalmus
spp. (Pimelodidae) swim in the water column filtering mostly zooplankton
(Carvalho and Goulding, 1985). Similarly, schools of the hemiodids
Anodus elongatus and Hemiodus spp. consume mainly phytoplankton
(although the later also frequently browse on periphyton). Some cichlids
are also specialized plankton feeders, notably Chaetobranchus flavescens, C.
semifasciatus and Chaetobranchopsis orbicularis. Close to the lake margin,
small characids (mainly Moenkhausia intermedia, Hemigrammus spp., and
Tetragonopterus spp.) feed on plankton and insects fallen in the water. The
truly omnivorous Triportheus albus, T. angulatus and T. elongatus also scan
the water surface searching for insects, small flowers, seeds of aquatic
grasses, fruits, and organic debris carried by the wind or by the waves.
The lake bottom presents low dissolved oxygen content throughout
most of the hydrological cycle. This oxygen depletion results from the
microbial decomposition of large amounts of organic matter that
accumulates in the bottom, and limits the presence of numerous fish
species. The fish fauna in the lake bottom is dominated by air-breathing
detritivorous loricariids, mainly species of Liposarcus and Hypostomus, and
by the ubiquitous callichthyid catfish Hoplosternum littorale that feeds
heavily on chironomid larvae. Huge schools of curimatids, notably
Psectrogaster rutiloides (but also species of Potamorhina, Curimata and
Cyphocharax) feed on the organic detritus at the lake bottom (Bowen,
1984; pers. obs.).
The floodplain lakes of white water rivers are fringed by a wide and
dense carpet of aquatic and semi-aquatic plants, mainly grasses of the
genera Echinochloa, Paspalum and Oryza (Junk and Piedade, 1993). These
floating meadows are probably the richest and most productive aquatic
habitats in the Amazon, with an enormous diversity and abundance of
invertebrates and algae associated to the submerged roots and stems
(Junk, 1970, 1973). Although frequently hypoxic, these habitats harbor
an abundant and rich fish fauna (Henderson and Hamilton, 1995;
Crampton, 1996; Sánchez-Botero and Araújo-Lima, 2001; Petry et al.,
2003), and supposedly function as a predator-safer environment for small-
size fishes (Kramer et al., 1978; Junk et al., 1987; Anjos et al., 2008).
Nevertheless, the floating meadows are frequently visited by the pirarucu
Arapaima gigas, one of the largest freshwater fishes in the world and a
generalized carnivore that eats fish, crabs and insects (EF, pers. obs.). The
ability to breathe air allows the pirarucu to withstand the low oxygen
conditions that predominate among the aquatic grasses (Kramer et al.,
1978).
The floating meadows can occupy circa 30% of the lake area
(Goulding et al., 2003), but surprisingly very few species feed directly on
the grass leaves, stems and roots. The few exceptions include the leaf-
eating giant doradid catfish Lithodoras dorsalis (but that also feed on fruits
in the flooded forest), and the anostomids Schizodon fasciatus and
S. vittatus (Santos, 1982). Other anostomids like Rhytiodus microlepis and
R. argenteofuscus are known to browse on the fine grass roots during the
flooding season (Santos, 1982). Although there is no explanation for the
low direct consumption of all that plant matter, some authors argue that
it represents a strongly seasonal resource, and during the dry season (when
almost all floating meadows are gone) the fishes suffer from food scarcity.
Nevertheless, fishes are known to withstand extended periods without
food, relying on fat reserves accumulated during the favorable season
(Junk, 1985).
The periphyton and invertebrates found among the submerged roots
and leaves of the aquatic plants are taken as food by a large number of
small fishes that live in this habitat, even in the scarce Oryza wild rice
banks found in some black water lakes (Araujo-Lima et al., 1986). The
cichlids of the genus Mesonauta are one of the most common fish species
living in the floating meadows, where they feed on algae, phytoplankton
and small invertebrates such as Conchostraca and Ostracoda. Juvenile
individuals of important commercially exploited food fishes of the genera
Colossoma, Triportheus, Brycon, Mylossoma, Semaprochilodus and
Prochilodus use this habitat for food and shelter during their first months
of life. Small tetragonopterine characins such as the ubiquitous

Ctenobrycon hauxwellianus also live among the aquatic grass beds, feeding
on small arthropods and seeds. Catfishes are present in large numbers,
including small mollusk- and insect-eating doradids (e.g. Doras spp.,
Nemadoras humeralis, and Pterodoras granulosus; Nelson et al., 1999), the
omnivorous auchenipterids Parauchenipterus spp. and the periphytivorous
loricariids Hypoptopoma gulare and Rineloricaria castroi (JZ, pers. obs.).
Needlefishes (mainly Potamorrhaphis guianensis, Belonidae; Plate 1.1) are
commonly found near the lake margins and close to the aquatic plants,
slowly moving just below the water surface in search of invertebrates that
fall in the water; small fishes are also occasionally ambushed by
needlefishes.
Deeply inside the oxygen-depleted, compact dead grass mats that
form in lakes is, the home of snake-like fishes such as swamp eels
(Synbranchus spp.) and electric knife fishes (Gymnotiformes). Species of
Brachyhypopomus (Hypopomidae) and Eigenmannia (Sternopygidae) feed
on small aquatic invertebrates, whereas the larger Sternopygus spp. (up to
40 cm long) includes small fishes as an important item on its diet. The very
large (up to 2 m long) electric eel Electrophorus electricus may also be found
in those places, hunting for fishes that are knocked-out by its powerful
electric shocks. The lungfish Lepidosiren paradoxa (Dipnoi:
Lepidosirenidae) is also commonly found amidst the rotten grass beds,
eating snails, insects and probably small fishes. The prey are brought to the
lungfish's mouth by a powerful suction, and immediately smashed and
crushed with its impressive set of fused teeth. Other fishes present in the
rotten grass beds include mainly hypoxia-tolerant species like the
predacious erythrinid Hoplias malabaricus (a fish and insect eater), young
pacus of the genus Mylossoma (that forage in the periphyton for algae,
microcrustaceans and insect larvae; Leite and Araujo-Lima, 2000, 2002),
and cichlids like the invertebrate feeding Cichlasoma amazonarum
(Ferreira, 1981; Sánchez-Botero and Araujo-Lima, 2001).
Flooded Forests
The seasonally available environment of the flooded forests represents a

keystone in the ecology of some large, commercially important food fishes
in the Brazilian Amazon (Gottsberger, 1978; Goulding, 1979, 1980;
Kubitzki and Ziburski, 1994; Piedade et al., 2003). The flooded forests of
black water rivers are locally named 'igapós', and that of white water rivers
as 'várzea' forests. These amphibious forests occupy an area of circa of 30%

of the floodplains in Brazilian Amazon (Goulding et al., 2003), and
represent an important pathway for the allochthonous primary production
to the aquatic environment.
The best examples of such fish-forest interaction are the large pacus
and several piranha species (Characidae, Serrasalminae), and at least two
large doradid catfishes, Lithodoras dorsalis and Pterodoras granulosus
(Goulding, 1980). The tambaqui Colossoma macropomum, one of the main
target fish species of the Amazon fisheries, depends largely on the fruits
and seeds produced in the flooded forests to survive (Waldhoff et al.,
1996). The fruits and seeds that fall in the water are readily consumed by
the tambaqui (Plate 1.1), including hard-shelled palm fruits that are
broken by the fish's powerful jaws and cheek muscles (which exhibit a
remarkable crushing power of up to 200 kg cm - 2 ; Araújo-Lima and
Goulding, 1997). Aside from its role in the energy transfer between the
terrestrial and aquatic habitats, the tambaqui and several other fishes
(e.g. Triportheus spp., Brycon spp., Mylossoma spp., Parauchenipterus spp.,
Auchenipterichthys spp., Lithodoras dorsalis, Pterodoras granulosus) may act
as seed dispersers for many plant species of the flooded forests, thus
contributing to the maintenance of its plant species diversity (Gottsberger,
1978; Goulding, 1980; Kubitzki and Ziburski, 1994; Ferreira, 2000;
Mannheimer et al., 2003; Piedade et al., 2003). Piranhas such as the large
and ferocious looking Serrasalmus rhombeus (up to 40 cm total length and
2 kg) also feed heavily on seeds in the flooded forests, but represent mainly
a seed predator since it masticate most of the food consumed (pers. obs.).
The trophic relations in the flooded forests are not restricted to the
fruits and seeds consumption by the fish fauna. Invertebrates that seek
refuge from the floods in the vegetation also fall in the water and are
readily consumed by many fish species such as Brycon spp., Triportheus
spp., Chalceus spp., and many small to medium-sized characins during the
day, and by Auchenipterichthys spp. and small cetopsids at night. Other
species like the arrowanas Osteoglossum bicirrhosum and O. ferreirai even
jump out of the water (sometimes up to two meters high) to reach large
insects or spiders hanging from tree branches (Smith, 1979; Aragão, 1984;
Goulding, 1989). Furthermore, the prochilodontids Semaprochilodus spp.
and Prochilodus spp. enter the flooded forest to browse the periphyton
growing on the submerged tree trunks, which represents a very important
food source in the oligotrophic black water systems (Ribeiro, 1985).
The relationships between fishes and flooded forests probably

represent the most remarkable biological interaction in the aquatic
environments of the Amazon, but not only for its ecological significance.
The huge amount of fruits and seeds that are produced during the flooding
season sustains an equivalently large biomass of commercially valuable
fishes, which enhances its importance as an energy pathway and for the
regional fisheries (Araújo-Lima et al., 1998). The knowledge of these
intricate ecological interactions and their impact on the regional economy
may represent an effective way to preserve the magnificent flooded forests
and the amazing diversity of aquatic environments in the Amazon.
FOREST STREAMS: HIGH DIVERSITY IN A FOOD-LIMITED

ENVIRONMENT
The Amazon forests are drained by an endless number of small streams of

clear and black waters (Plate 1.2). Despite being unnoticed under the
dense forest canopy that covers most of the region, these hidden
watercourses contribute with a large portion of the water that the mighty
Amazon River empties every year in the Atlantic Ocean (Goulding et al.,
2003). These streams help to maintain the typical high levels of relative
humidity inside the forest and so conditioning the presence of very
characteristic plant assemblages in the stream valleys (Ribeiro et al.,
1999).
The smallest water courses that drain the upland forests in the
Amazon are first - to third-order streams which do not exhibit the seasonal
flood pulse that dominate the dynamics of the large river-floodplain
systems (Junk et al., 1989). Instead, these streamlets show unpredictable
floods of short duration (from some hours to a few days), dispersed along
the lengthy Amazonian rainy season. This unpredictability does not allow
for the development of the remarkable feeding specializations based on the
consumption of fruits and seeds, as recorded in the floodplain habitats
(Goulding, 1980). Nevertheless, the small and shallow forest streams
harbor temporally stable, well defined fish assemblages.
These streams are characteristically oligotrophic (i.e., with a low
autochthonous primary productivity) and apparently sustain a low
biomass of fishes (e.g. Henderson and Walker, 1986; Walker, 1987; Silva,
1993; Mendonça et al., 2005). Most of the time the sunlight barely reaches
the stream surface, and the very low levels of nutrients in the water
contribute to impair the growth of aquatic plants of almost all sorts (with
the exception of some red algae species of the genus Batrachospermum; JZ,
Plate 1.2
(Upper left) General view of a typical forest stream, showing the clear water, predominantly
sandy substrate and dense canopy cover. Photo: L.N. Carvalho.
(Upper right) The ubiquitous forest stream dweller Pyrrhulina brevis (Characiformes:
Lebiasinidae) photographed in a field aquarium. Photo: F.P. Mendonça.
(Second row, left) Cryptically colored suckermouth armored catfishes like Rineloricaria
heteroptera (Siluriformes: Loricariidae) graze the periphyton that grows attached to trunks,
submerged roots and scarce macrophyte stands in forest streams. Photo: J. Zuanon.
Plate 1.2 Contd.
pers. obs.). Nevertheless, a single third-order stream, approximately 5 m

wide and 1 m deep, may contain up to 50 fish species of five different
orders (Anjos, 2005; Mendonça et al., 2005; JZ, pers. obs.). Just to allow
a comparison, this species richness corresponds to one fourth of the whole
European freshwater fish fauna and ca. 10% of the freshwater fishes of
North America (Goulding et al., 2003).
The fish fauna of the small Amazonian streams is seemingly composed
of a process of species addition from the headwaters to the larger water
courses (Vannote et al., 1980). The nutrient-limited environment of the
first-order streams is occupied almost exclusively by fishes that feed on
allochthonous sources of food, such as terrestrial insects and other small
invertebrates that fall from the trees. Ants and termites are common food
items in the stomach contents of these stream fishes and are caught as
soon as they reach the water surface. The Gasteropelecids Carnegiella
strigatta (Plate 1.2) and C. marthae, and the Characid Gnathocharax
steindachneri have mastered this strategy aided by their upturned mouths
and powerful pectoral fins, which allow a really fast access to the food that
fall in the water surface. Small invertivorous such as the characins
Hemigrammus pretoensis and Hyphessobrycon agulha, as well as the
lebiasinids Pyrrhulina brevis (Plate 1.2) and Copella nigrofasciata figure
among the most abundant fish species in the headwater streams in
Brazilian Central Amazon. Although insects constitute the dominant food
item in the diet of most species in these streams, the use of diverse feeding
tactics, different foraging grounds and activity periods (day vs. night)
allow for the coexistence of several fish species with similar diets (e.g.
Soares, 1979; Silva, 1993; Sabino and Zuanon, 1998).
Plate 1.2 Contd.

(Second row, right) The catfish Mastiglanis asopos (Siluriformes: Heptapteridae)
performing the drift trap foraging tactic during the night at the stream bottom. Photo:
J. Zuanon.
(Third row, left) A small group of the dwarf cichlid Apistogramma hyppolitae (Perciformes:
Cichlidae) sifting the detritus layer that accumulates in the litter banks for edible particles
and invertebrates. Photo: J. Zuanon.
(Third row, right) An adult Rivulus compressus (Rivulidae: Cyprinodontiformes)
camouflaged among the leaf litter in a desiccating pond at Reserva Florestal Adolpho
Ducke, Manaus, Amazonas State, Brazil. Photo: J. Zuanon.
(Bottom left) The powerful pectoral fins of the marbled butterfly fish Carnegiella strigata
(Characiformes: Gasteropelecidae) allow the fish to quickly reach allochthonous
invertebrates fallen at the water surface. Photo: J. Zuanon.
(Bottom right) The traira or wolf characin Hoplias cf. malabaricus (Characiformes,
Erythrinidae), a top predator in small streams, ambushes its prey among submerged
plants. Photo: J. Zuanon.
Reduction of body size (= miniaturization; Weitzman and Vari, 1988)

seems to constitute a very common trait among fishes that inhabit food -
limited environments. In fact, one of the few recognizable common
characteristics of stream fishes in Brazil is the small adult size of most
species (up to 15 cm standard length; Castro, 1999). This condition is also
observed in the black waters of the enormous Negro River, where
nutrients and, ultimately, food are very scarce in the aquatic environment.
The low primary productivity and scarcity of food are expectedly reflected
in the stream trophic chain, where detritivorous fishes are rare to almost
absent. The trophic link between the producers and the ichthyofauna is
through the leaf-processing invertebrates, which constitute an important
food source for many fish species.
The food chain in forest streams is characteristically simple and short
(Walker, 1987). The mentioned rarity of detritivorous fishes is
accompanied by the inexistence of phytoplanktivorous and strictly
zooplanktivorous species. As already mentioned, the low light and
nutrient-poor waters of the forest streams do not sustain large enough
populations of planktonic algae and microcrustaceans to allow for the
existence of plankton-feeding specialists. Similarly, the almost absence of
aquatic macrophytes impairs the occurrence of plant (leaf) eating fishes.
So, among the fishes that inhabit forest streams the primary consumers are
mainly represented by periphytivorous loricariids of the genera
Rineloricaria, Acestridium, Farlowella, Ancistrus and Parotocinclus. The most
abundant loricariids in the forest streams seem to be species of
Rineloricaria, especially R. lanceolata and R. heteroptera (Plate 1.2). These
species are usually found at stream stretches where the forest canopy is
somewhat open, so allowing the presence of stands of bog plants like
Thurnia sphaerocephala (Thurniaceae), where the suckermouth catfishes
graze the periphyton that grows in the leaf blades.
Other fish species inhabit the slow-flowing stream margins, taking
advantage of the calmer conditions to browse on algae and nip on
associated tiny invertebrates along the clay banks and on submerged
trunks and tree branches. These micro-omnivorous fishes are represented
by some Nannostomus species, notably the beautiful and widespread
N. marginatus in the smaller streams. In streams of third order and up, this
niche is frequently occupied by some cichlid species of the genus
Mesonauta, the ubiquitous Amazonian flag cichlid. The streamlined
characins Iguanodectes variatus and I. geisleri are also frequently observed
browsing on algae attached to thin tree roots that emerge from the banks
and from the stream bottom, making conspicuous head-twisting
movements while foraging.
Mainly invertivorous fishes that feed on allochthonous insects,
spiders, mites and invertebrates, as well as immature aquatic insects
compose the secondary consumers. This trophic group commonly
includes several characiform fishes (e.g. species of Bryconops,
Hyphessobrycon, Hemigrammus, Moenkhausia, Poptella, Axelrodia,
Pyrrhulina, Copella and others) and perciforms (mainly cichlids of the
genera Apistogramma and Aequidens), as well as the siluriforms Helogenes
marmoratus and Denticetopsis seducta (Cetopsidae), Tatia spp.
(Auchenipteridae), Ituglanis amazonicus (Trichomycteridae), Mastiglanis
asopos and Brachylanis cf. microphthalmus (Heptapteridae). The presence
of several species of fishes exploring the same kind of food resource may
seem contradictory in an oligotrophic environment. This situation seems
to be allowed by some sort of resource sharing, and the activity period
(diurnal vs. nocturnal) is possibly the most common and simple way to
accomplish this. Characins and cichlids compose the majority of the
diurnally active fishes in the forest streams, whereas the night-time is
dominated by catfishes (Siluriformes) and knifefishes (Gymnotiformes)
(Lowe McConnell, 1987). Nevertheless, some fishes exhibit a higher
trophic specialization and feed on fewer prey types. This is the case of the
Auchenipterid Tetranematichthys quadrifilis, which eats mainly shrimps and
constitute one of the few carcinophagous (crustacean-eating) fish species
in the Amazon (although insects may be occasionally preyed on by the
catfish).
Another form of resource sharing includes the vertical stratification of
foraging grounds. Several fishes feed on aquatic invertebrates that live
over and amidst the sandy bottom of the streams. During the day,
generalized sit-and-wait foragers like species of the genus Characidium
(Crenuchidae) exploit the bottom dwelling and drifting invertebrates
along the shallow stream margins. In fast-flowing stretches, the minute
ghost candiru (Stauroglanis gouldingi, Trichomicteridae) feeds on very
small aquatic insect larvae and harpacticoid copepods that accumulate in
the grooves formed behind sand ripples (Zuanon and Sazima, 2004b). At
night, the sit-and-wait invertivorous niche is occupied by the heptapterids
Imparfinis pristos and Mastiglanis asopos that feed on drifting prey while
poised on the stream bottom surface (Plate 1.2). Species of the sand
knifefish Gymnorhamphichtys forage by probing the upper layer of the sand

with their tubular snouts where they prey on small chironomid
(bloodworm) larvae and harpacticoid copepods (Zuanon et al., 2006). In
larger streams, species of Satanoperca and Geophagus (Cichlidae) sift food
from the bottom by taking a mouthful of substrate and sorting the edible
portions (aquatic invertebrates, organic particles) in the oral chamber.
Finally, the minute, eel-like trichomycterid species of the genus
Pygidianops goes deeper and literally 'swims' amidst the sand searching for
tiny aquatic invertebrates, and apparently never ventures out of its cover
(Schaefer et al., 2005; JZ, pers. obs.).
The litter banks are also shared by several fish species of different
groups. The dwarf cichlids of the genus Apistogramma seek shelter and
forage on the upper layer of the banks, picking up small aquatic
invertebrates and sifting through small amounts of organic detritus during
the day (Plate 1.2). At night, the widespread banjo catfishes of the genus
Bunocephalus (such as B. coracoideus and B. verrucosus, Aspredinidae)
forage by slowly moving over the leaf substrate pushed by jets of water
expelled through the gill openings. Insect larvae and other small aquatic
invertebrates are found with the use of its barbels and quickly sucked from
the substrate. Another common inhabitant of the litter banks is the
crenuchid Elachocharax pulcher (as well as the occasionally syntopic
E. mitopterus and E. junki); surprisingly, these small characiforms (up to
20 mm SL) are nocturnal but very little is known about its ecology. The
eel-like fishes that occupy the muddy and hypoxic deeper layers complete
the vertical stratification of the litter banks. The swamp eels of the genus
Synbranchus are commonly found under these conditions, along with
electric fishes of the genera Gymnotus, Microsternarchus, Hypopygus and
Brachyhypopomus (Gymnotiformes). Although little information is
available about the natural history of these species, it seems that most of
them leave the litter banks at night to forage in open waters for small fishes
and aquatic invertebrates. But analogous to the sand-diving Pygidianops
trichomycterids in the sand banks, the red-bodied heptapterid species of
the genus Phreatobius keeps deep inside the litter banks, moving amidst
the thick packs of dead leaves and (supposedly) feeding on minute
invertebrates (Muriel-Cunha and de Pinna, 2005; JZ, pers. obs.).
The top predators among the forest stream fishes are represented
mainly by the wolf characins of the Family Erythrinidae (Géry, 1977). In
the smaller (first- and second-order) streams, the most abundant
carnivore is the 'jeju' Erythrinus erythrinus, a fish that attains ca. 15 cm SL

and feeds on small fishes and insects. The larger Hoplerythrinus
unitaeniatus and Hoplias cf. malabaricus (Plate 1.2) stand among the most
common and abundant crepuscular to nocturnal fish predators in the
larger streams, as well as in several South American water bodies. Juvenile
specimens of the piscivorous Rhamdia catfishes (Heptapteridae) may be
occasionally found venturing upstream in small forest streams, whereas
the wide-mouthed Batrochoglanis raninus (Pseudopimelodidae) is
frequently found in the crevices of submerged logs lurking for prey fishes
at night. During the day, individuals of the dogtooth characin
Acestrorhynchus falcatus (Acestrorhynchidae) cruises at mid water
pursuing small tetras (Characidae) in second order and larger streams.
Temporary Ponds along the Streams: To Eat or to be

Eaten?
During the rainy season, several ponds may be formed in flat-bottomed

valleys along the forest streams. Although insects and frogs are the main
colonizers of the ponds, these small and (usually) ephemeral
environments are also capable of sustaining a particular assemblage of
fishes that differs from a simple subset of the stream ichthyofauna. It is
supposed that the marginal ponds constitute a more productive
environment for the growth of larvae and juvenile fish, like the floodplain
lakes associated to larger rivers. In fact, a large portion of the fishes found
in marginal ponds of forest streams in Central Amazon is composed by
juvenile specimens (Pazin et al., 2006). Nevertheless, some fish species are
far more common in ponds than in the stream channels, and there are
trophic differences among these habitats.
The ponds are colonized mainly by means of opportunistic migrations
of fishes during the sporadic floods of the streams. In other instances, some
air-breathing species actively migrate through runoff waters or even moist
land to the nearby ponds. This group of species includes the predatory
characiforms Erythrinus erythrinus and Hoplias cf. malabaricus, the
siluriforms Callichthys callichthys and Megalechis thoracata, and the
cyprinodontiform Rivulus compressus. The most obvious advantage of
being the first colonizers is the access to newly available food sources and
shelter. Aquatic immature insects like bloodworms (Diptera,
Chironomidae), leaf-cased caddis fly (Trichoptera) and microcrustaceans
(mainly Ostracoda and Cladocera) readily colonize the ponds and are
heavily consumed by the fishes. On the other hand, the highly predatory
dragonfly nymphs (Odonata) also figure among the earlier colonizers but
attracted by the predictable abundance of food.
The trophic chain in the ponds seems to be slightly different from that
in the stream channel. The ubiquitous Pyrrhulina brevis and Copella
nigrofasciata again constitute a large portion of the fishes foraging on
insects and other invertebrates that fall into the water surface. Several
Rivulus species (mainly R. compressus in the Central Amazon; Plate 1.2)
occupy the uppermost layer of the ponds, staying almost motionless at the
surface and close to the pond margin, preying chiefly on small ants. Small
cichlids (mainly Apistogramma and juvenile specimens of Aequidens
pallidus) and the beautiful crenuchid Poecilocharax weitzmani feed on small
aquatic invertebrates among the leaf litter that accumulates in the pond
bottom. The callichthyids Megalechis thoracata and Callichthys callichthys
also forage on the bottom, probing the substrate with their barbels and
feeding on a compound of organic particles and associated micro-
invertebrates, thus inserting a detritivorous component in the food chain.
Again, the main piscivorous fishes in this aquatic habitat are the
erythrinids and Synbranchus, which seems to be very efficient predators in
the shallow, leaf-packed environment of the ponds.
Although more productive and (supposedly) less competitively
stressful than the stream channel, colonizing the marginal ponds present
some risks to the survival of the fishes. The main problem is related to the
survival during the dry season, when most of the ponds dry completely and
may turn into a deadly seasonal trap. Evading the ponds at the end of the
rainy season and just before they become completely isolated from the
stream seems to be the commonest strategy employed by opportunistic
occupants of the ponds, like most of the small characins. The
disconnection from the stream does not represent a problem for the air-
breathers Callichthys and Megalechis, which can migrate back to the
channel by dry land. A similar emergency escape strategy is shown by
species of Rivulus that just jump away from the ponds to the stream
(although R. compressus can withstand the dry period partially buried in
the mud or moist leaf litter). The leaf-litter specialists like the eel-like
Phreatobius catfishes can migrate to deeper layers of the litter bed where
some water (or moister conditions) remains throughout the dry season.
Some electric fishes (several species of Gymnotus) and swamp eels
(Synbranchus) can bury themselves and survive the desiccation of the
ponds inside the muddy bottom or in moist galleries. During that time,
these species do not eat and survive on the fat reserves accumulated along
the rainy season. Annual killifishes that specialize in ephemeral
environments and complete the short (less than a year) life cycle in the
pond (by means of burying the eggs in the soft pond substrate) are not
common in the forest environments of the Central Amazon.
As mentioned before, the productive environment of the marginal
ponds may turn in a very risky environment for fishes, since they are
occupied by some large and voracious predators like Pipa toads (Pipidae)
and the small fish-eating twist-necked turtle Platemys platycephala
(Chelidae). Furthermore, the desiccating ponds frequently end up with
reversed predator-prey roles in the food chain: invertebrates as predators
(mainly dragonfly larvae and giant water bugs of the family
Belostomatidae), and vertebrates as prey (fishes and amphibian larvae). In
conclusion, the different abilities of fish to stand in such an ephemeral
habitat play a major role in the constitution of fish assemblage in ponds,
which may change along time as a result of the complex interactions
among predators and prey in that aquatic environment.
RAPIDS AND WATERFALLS: GRAZING ROCKS FOR A

LIVING
Besides the huge floodplains and the lengthy white water rivers that form
the dominant elements in the region, the Amazon Basin comprises other
aquatic environments that harbor some specialized assemblages of fishes.
A very different kind of aquatic habitat exists in the region where the
Amazon's sedimentary floodplain contacts the Guyana and the Central
Brazilian Plateaus. These transition zones are far from smooth: there, a
series of rapids and waterfalls clearly mark the limits of two very different
aquatic environments. Downstream from the rapids these rivers show a
predictable annual flood pulse that strongly influences the dynamics of the
rivers themselves and of their fish fauna. Upstream the riffle zone the
landscape is composed by a long stretch of rocky outcrops and turbulent
waters, where stands of podostemaceans plants carpet the rocks under the
strongest water flow.
Most of the rapids and waterfalls located at the edge of the two main
plateaus are formed by clear water rivers that carry very few suspended
solids, since they drain very old, continuously washed granitic rocks (Plate
1.3). Although clear water rapids constitute the more common situation
in the Brazilian Amazon, rapids zones also occur in rivers of different water
Plate 1.3
(Upper left) General view of some rapids in the Xingu River near the city of Altamira, Pará
State, Brazil. Note the clear water typical of most rivers that drain the Central Brazilian
plateau. Photo: J. Zuanon.
(Upper right) The underside of submerged rocks in Xingu River is colonized by several
numerous invertebrates like sponges, gastropods, and insect larvae that are utilized as
food by several rapids-dwelling fishes. Photo J. Zuanon.
Plate 1.3 Contd.
types. Black water rapids exist in the upper reaches of the Negro River
near the Brazilian-Colombian border and in a few other tributaries. White
water rapids and waterfalls are extremely rare: they occur only in the
Madeira River, extending from the vicinity of Porto Velho (Rondônia
State) upstream to the Bolivian border. Both black- and white-water
rapids harbor a lower diversity of strictly rheophilic fishes when compared
with clear water rivers (JZ, pers. obs.), which seems to result from
differences in the local primary productivity. In black waters, the light
penetration is limited by the large amount of dissolved organic matter that
darkens the water and impairs the growth of periphyton in the submerged
rocks (Goulding et al., 1988). In white water rapids the light is scattered
by the huge amount of suspended sediments carried by the Madeira River,
and the incident light barely penetrates the upper (20 cm) layer of water
(Goulding et al., 2003). These facts indicate that food (periphyton) is
probably the main limiting factor to the fish fauna in black and white water
river rapids, although faunal limitations cannot be ruled out.
The rapids constitute highly structured habitats, where rocks of
different sizes provide lots of shelters for fishes and other aquatic life. The
hard rocky substrate allows for the establishment of a dense cover of algae
and other sessile organisms like sponges, bryozoans and podostemaceans
plants. Nevertheless, these characteristics are accompanied by a strong
Plate 1.3 Contd.

(Second row, left) The zebra pleco Hypancistrus zebra (Siluriformes: Loricariidae)
photographed in a field aquarium. The bold color pattern rendered this species very
attractive to the aquarium trade, occasionally driving it to an endangered conservation
status. Photo: J. Zuanon.
(Second row, right) A close-up view of Sartor respectus (Characiformes: Anostomidae)
photographed in field aquarium just after capture in the Xingu River. Note the upturned
mouth and the protruding red-tipped teeth that are employed to scrap off sessile
invertebrates from submerged rocks. Photo: J. Zuanon.
(Third row, left) The riffle specialist anostomid Leporinus julii (Characiformes:
Anostomidae) browsing the periphyton of submerged rocks in the Xingu River. Photo:
J. Zuanon.
(Third row, right) Omnivorous generalists like Leporinus aff. fasciatus (Characiformes:
Anostomidae) are commonly found foraging behind rocks and so protected from the direct
action of the current. Note the dark sponges where the fish is feeding. Photo: J. Zuanon.
(Bottom left) The odd-looking pacu Ossubtus xinguense (Characiformes: Characidae:
Serrasalminae) photographed just after capture. This endemic species of the Xingu River
in Brazil feeds on algae and podostemacean plants that grow attached to the rocks in the
rapids. Note its strongly down-turned mouth. Photo: J. Zuanon.
(Bottom right) The rapids-dwelling cichlid Retroculus xinguensis (Perciformes: Cichlidae)
photographed just after capture. Note the long snout and thick lips that allow this species
to forage deeply in sand-pebbles substrate of the rapids. Photo: J. Zuanon.
water turbulence that limits the presence of most of the fish species that
abound in the calmer downstream sections of those rivers. To cope with
these environmental conditions, several morphological, physiological and
behavioral solutions have evolved in different fish groups.
The fish fauna in the clear water rivers rapids is characteristically rich
and diverse. More than a hundred species may occupy a relatively short
(100 km) stretch of the Xingu River rapids (Zuanon, 1999), and similar
numbers may be found in the Tocantins, Trombetas and other rivers in the
Brazilian Amazon (pers. obs.). This impressive species richness is
apparently supported by the high primary productivity in the rapids, where
periphyton constitutes the main food or foraging ground for the fish fauna.
The diverse characteristics of the rocky substrate and water flow
dynamics, coupled with the morphological diversity of the fishes, have
allowed the coexistence of many fish species. The consumption of a same
food type by so many fishes seems possible by the use of a variety of feeding
tactics and behavioral traits.
Variations over a Theme: Mouth Morphology, Teeth Types

and Trophic Characteristics of Rapids-dwelling Fishes
One of the most successful fishes dwelling in the rapids are the Loricariids
or armored suckermouth catfishes. Although usually known as important
detritivorous or periphytivorous, these odd-shaped catfishes are, in fact,
versatile foragers that exploit a large diversity of food types. Loricariids use
their sucker mouths and specialized teeth to graze on several kinds of food,
which result in diverse feeding habits: detritivory, periphyitivory,
omnivory, and even invertivory. Some loricariid species go even further
and seems to show specialized carnivorous habits, feeding on hard-shelled
invertebrates like mollusks and caddis fly larvae, and even sponges. This
feeding tactic known as durophagy (Hoogerhoud, 1987; Turingan, 1994)
is exhibited by members of the genus Scobinancistrus, which possess a few
strong teeth that seem to be employed as pincers to scrap off food from the
rocky surface. On the other hand, the flat-bodied Ancistrus ranunculus
feeds on minute algae and cyanobacteria that seem to be obtained by
suspending the fine detritus layer settled on the slit-like cracks on the
rocks inhabited by these catfish (Müller et al., 1994; Zuanon, 1999). The
delicate and numerous teeth of A. ranunculus and other loricariids seem
adequate to feed on loosely settled unicellular diatomaceous algae that
grow amidst the firmly attached tufts of filamentous blue-green algae
(Cyanophyceae), by 'combing' the algae tufts (Müller et al., 1994; Rapp

Py-Daniel and Zuanon, 2005).
Although predominantly herbivorous, some loricariids include a
significant portion of animal food in their diets. The boldly patterned
catfish Hypancistrus zebra (Plate 1.3) seems to be a sedentary forager that
explores small areas around its shelter on the rocks, where it feeds on
periphyton, chironomid larvae and microcrustaceans. The 'v'-shaped
dentary teeth of the zebra loricariid seem to allow the selection of small
prey animals amidst the detritus layer or periphyton that cover the rocks.
Finally, some species of loricariid shows a bizarre diet composed by wood
fragments gathered from submerged trunks and logs. The wood-eating
Panaque nigrolineatus and Hypostomus cochliodon scrap small pieces of wood
aided by the strong and spoon-shaped teeth in both jaws, and are
apparently capable of digesting the cellulose matter aided by symbiotic
invertebrates living on their guts (Nelson et al., 1999).
Scraping food from the rocks is also the main feeding strategy of a very
different group of fish. The leporins and allies of the family Anostomidae
are torpedo-shaped characiform fishes that abound in the rapids and
constitute the more conspicuous inhabitants of the fast flowing rocky
stretches of clear water rivers. The widespread Leporinus friderici and the
boldly patterned L. tigrinus and L. cf. fasciatus (Plate 1.3) occupy the
periphery of the rapids where they feed on a variety of invertebrates, algae
and other organic debris. These species do not present evidence of
morphological adaptations related to the life in the rapids and seem to
occupy those habitats by means of a simple strategy: to avoid struggling
against the faster, more turbulent waters of the rapids. These two species,
as well as several fishes of various taxonomic groups use the patch of
calmer waters formed downstream of rocks and other obstacles to
occasionally access some feeding grounds in the middle of the rapids. A far
more specialized way to avoid the direct effects of the currents is showed
by Sartor respectus, a small anostomid species with an upwardly directed
mouth (Plate 1.3). The very large dentary teeth of this fish emerge out of
the mouth and are utilized to scrap off hard-bodied invertebrates
(bryozoans, sponges) from the ceilings of dark shelters under the rocks
(Santos and Rosa, 1998; Zuanon, 1999) (Plate 1.3). These fishes
apparently do not venture out of their rocky shelters and probably
constitute one of the more specialized species that inhabit the rapids
stretches of clear water rivers.
Nevertheless, there are truly rheophilic specialists among the

anostomids that live in the rapids. Leporinus julii (Plate 1.3) is a long and
slender species that possess a down-turned mouth furnished with a series
of aligned sharp teeth that allow it to browse and graze the algae that
carpet the rocks in the shallow, faster flowing stretches of the rapids. The
streamlined body allows the fish to roll on its sides and reach and graze on
tufts of algae that grow in the small nooks and crannies of the rock surface.
The thick margins of the pelvic, anal and lower lobe of the caudal fins are
utilized as a tripod that helps the fish to hold position against the current
while foraging.
Another group of Characiformes constitute an important component
of the fish fauna in the rapids: the leaf-cutting and algae-browsing pacus
(Characidae: Serrasalminae). Species of Mylesinus and Tometes occupy the
stronger flowing stretches of the rapids, where they feed mainly on the
leaves of podostemaceans plants that grow on the rock surface. A further
step towards the life strictly in the rapids is shown by Ossubtus xinguense,
which have a down-turned mouth and scraps algae in flat rocks at the
bottom of the rapids (Plate 1.3). This endemic species of the Xingu River
probably represents the most extreme example of morphological
adaptation to the rheophilic life among the serrasalmines.
Although unexpectedly, some cichlids also inhabit the rapids of clear
water rivers in the Amazon. The above-mentioned strategy of avoiding
the direct effects of the strong currents is also employed by several species
of Crenicichla that hunts for small fishes and aquatic invertebrates among
the rocks at the bottom of rapids. Similarly, some peacock basses of the
genus Cichla have been observed hunting for prey in the turbulent zone
downstream of small waterfalls in Trombetas and Xingu rivers (pers. obs.).
Nevertheless, some cichlids can be considered as strictly rheophilic.
Retroculus xinguensis (Plate 1.3) digs the gravel and sand substrate in fast-
flowing river stretches, where it feeds on chironomids, mayfly and caddis
fly larvae (Moreira and Zuanon, 2002). The small pike cichlids of the
genus Teleocichla also live in the rapids and feed on aquatic invertebrates
(Zuanon, 1999). Of these, Teleocichla centisquama is a strongly rheophilic
species that forages at the top of submerged boulders directly exposed to
the strong water current, where it picks small insect larvae and micro
crustaceans at the rock surface (Zuanon, 1999; Zuanon and Sazima,
2002).
After night fall, the bottom of the rapids is occupied by several

carnivorous species that feed on fishes and aquatic invertebrates under
the rocks, including small catfishes of the genera Microglanis

(Pseudopimelodidae) and Myoglanis (Heptapteridae), and juvenile
specimens of the swamp eel Synbranchus spp. (Synbranchidae). Some very
large individuals of Hoplias macrophthalmus are frequently seen ambushing
preys under rocky shelters in the rapids, and the weakly electric eel
Archolaemus blax (Sternopygidae) emerges from small spaces and cracks in
the rocks to forage for insect larvae and small fishes.
In conclusion, trophic relationships play a major role in the structure
of fish assemblages in the rapids of Amazonian rivers. The high primary
productivity, coupled with slight variations in mouth and teeth
morphology and different feeding strategies, allow for the presence of a
highly diverse fish fauna. The presence of many endemic species in the
rapids also points out to the need for special conservation strategies since
those areas are highly threatened by the construction of hydroelectric
power plants in the Brazilian Amazon.
Aknowledgements
We wish to thank L.N. Carvalho, M. Goulding and I. Sazima for criticisms

and suggestions to the text; to M. Goulding, L.N. Carvalho, and F.P.
Mendonça for allowing the use of some photographs that illustrate this
chapter; and to INPA, WWF-Brasil, Fundaçã o 'O Boticário' de Proteçã o
à Natureza FAPEAM
, ,
Fundaçã ,
o Djalma Batista Fundaçã o Rio Madeira,
CNPq, CAPES, ORSTOM/CNRS, Unicamp, Eletronorte, and FURNAS
for logistic and financial support along the last 25 years of ecological
studies in several Amazonian rivers. We are also grateful to INPA's
fishermen for the help during fieldwork along the last 25 years, and for
sharing valuable information about the natural history of Amazonian
fishes.
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— Venez donc un peu que je vous explique, vous Rose, la
Maternelle ; y a longtemps que j'ai envie de vous causer… Venez
donc là, dans le coin.
Elle bombait ses épaules et avançait le menton comme Adam
quand il va se battre ; son intonation copiait celle des provocations
en usage dans le quartier : « Viens donc un peu, su' l' boulevard, si
t'es pas un faignant! »
Je la suivis, moitié de gré, moitié parce qu'elle me tenait au
coude. Elle se mit à me parler dans la figure.
— Non, elle n'ira plus à vot' école ma fille… c'est pas la peine,
pour apprendre qu'il faut rester dans la débine comme père et mère
et se tenir bien tranquille, en crevant de faim tout comme eusses et
surtout pas oublier de dire merci… Mais c'est pas vrai, vos histoires!
il ne suffit pas d'être poli… Et qu'est-ce que t'avais l'air de rigoler en
me regardant, avec ton intérêt moral? L'intérêt c'est de bouffer… J'y
ai été à l'école moi, est-ce que ça m'a empêchée de crever la
misère?… Ah! oui, j'ai fait comme ma mère, pour sûr!… Et quand ma
gosse me répétait vos boniments d'école, je croyais entendre mes
premiers patrons : de l'ordre, de la propreté, du respect, de
l'obéissance, de la politesse… Oui! et des dix-huit heures de travail
et mal nourrie, et pas de pitié, pas de bon Dieu, jusqu'à ce qu'on
vous flanque dans le ruisseau… Et v'là que c'est toujours les mêmes
boniments que de mon temps! mais je veux autre chose!… Dis donc,
la maternelle, est-ce que tu crois que c'est toujours les mêmes qui la
danseront!… Dis donc, chienne de garde, chienne d'administration,
me v'là moi, devant ta baraque en pierres de taille, et v'là ma
gosse… est-ce que tu crois que ça va recommencer? Je te le
demande?… Qué que tu dis?… Tu veux pas me répondre… De quoi
que tu te mêles alors?… On n'a pas besoin de toi, laisse donc les
malheureux : tu n'auras pas ma gosse pour ton école de crève-la-
faim!… Va-t'en de not' passage!…
Et, du geste le plus irréconciliable qu'eût jamais précipité la
maternité en révolte, elle me chassa de sa misère.
FIN
Fontenay-aux-Roses (Seine). — Imp. Louis Bellenand. 9-04

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No claim to original U.S. Government works

Printed on acid-free paper

International Standard Book Number-13: 978-1-57808-375-6 (Hardback)

Understanding the biology of the innumerable number of aquatic species

investigators cover topics as diverse as feeding ecology of fish in their

J.E.P. Cyrino, D.P. Bureau and B.G. Kapoor

1. Feeding Ecology of Fishes in the Brazilian Amazon - 1

4. Classical and Modern Concepts in Fish Digestion 85

7. Ontogeny and Physiology of the Digestive System of 281

8. Tracer Studies in Fish Larvae 349

Conceiçã o Luís E.C.

Dinis Maria Teresa

Hahn Norma Segatti

Portella Maria Célia

Zambonino-Infante José Luis

Feeding Ecology of Fishes in

Jansen Zuanon* and Efrem Ferreira #

The Amazon system is composed by a multitude of rivers with varying

habitats. This chapter focuses on the main aquatic environments found in

Plate 1.1 Contd.

water habitats, floating meadows and the flooded forests. Although

Plate 1.1 Contd.

substrate (Goulding and Ferreira, 1984), whereas groups of the widespread

Roeboides are supposed to rely on their body translucency and scattered

predators and plankton-eating schooling fishes. Among the commonest

of life. Small tetragonopterine characins such as the ubiquitous

The seasonally available environment of the flooded forests represents a

as 'várzea' forests. These amphibious forests occupy an area of circa of 30%

The relationships between fishes and flooded forests probably

FOREST STREAMS: HIGH DIVERSITY IN A FOOD-LIMITED

The Amazon forests are drained by an endless number of small streams of

pers. obs.). Nevertheless, a single third-order stream, approximately 5 m

Plate 1.2 Contd.

Reduction of body size (= miniaturization; Weitzman and Vari, 1988)

knifefish Gymnorhamphichtys forage by probing the upper layer of the sand

carnivore is the 'jeju' Erythrinus erythrinus, a fish that attains ca. 15 cm SL

Temporary Ponds along the Streams: To Eat or to be

During the rainy season, several ponds may be formed in flat-bottomed

RAPIDS AND WATERFALLS: GRAZING ROCKS FOR A

Plate 1.3 Contd.

Variations over a Theme: Mouth Morphology, Teeth Types

(Cyanophyceae), by 'combing' the algae tufts (Müller et al., 1994; Rapp

Nevertheless, there are truly rheophilic specialists among the

After night fall, the bottom of the rapids is occupied by several

the rocks, including small catfishes of the genera Microglanis

We wish to thank L.N. Carvalho, M. Goulding and I. Sazima for criticisms

Araujo-Lima, C.A.R.M., L.P.S. Portugal and E.J.G. Ferreira. 1986. Fish-macrophyte

Muriel-Cunha, J. and M. de Pinna. 2005. New data on cistern catfish, Phreatobius

Fontenay-aux-Roses (Seine). — Imp. Louis Bellenand. 9-04